The CANADIAN FIELD-NATURALIST A JOURNAL OF FIELD BIOLOGY AND ECOLOGY Promoting the study and conservation of northern biodiversity since 1880 i Ps ath) \. Volume 133, Number 1 ¢ January—March 2019 Ottawa Field-Naturalists’ Club Club des naturalistes d’Ottawa The Ottawa Field-Naturalists’ Club FOUNDED 1863 (CURRENT INCORPORATION 1879) Patron Her Excellency the Right Honourable Julie Payette, C.C., C.M.M., C.O.M., C.Q., C.D. Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encour- age Investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Honorary Members Ronald E. Bedford Michael D. Cadman J. Bruce Falls Robert E. Lee Allan H. Reddoch Charles D. Bird Paul M. Catling Peter W. Hall John Mcneill Joyce M. Reddoch Fenja Brodo Francis R. Cook Christine Hanrahan Theodore Mosquin Dan Strickland Irwin M. Brodo Bruce Di Labio C. Stuart Houston Robert W. Nero John B. Theberge Daniel F. Brunton Anthony J. Erskine Ross A. Layberry E. Franklin Pope Sheila Thomson 2019 Board of Directors President: Diane Lepage Annie Bélair Edward Farnworth Dwayne Lepitzki Henry Steger 1st Vice-President: Jakob Mueller Fenja Brodo Catherine Hessian = Bev McBride Ken Young Recording Secretary: Elizabeth Moore Robert Cermak Anouk Hoedeman Gordon Robertson Eleanor Zurbrigg Treasurer: Ann Mackenzie Owen Clarkin Diane Kitching Jeff Saarela To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofnc.ca. For information on club activities, go to www.ofnc.ca. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this jour- nal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Website: www.canadianfieldnaturalist.ca/index.php/cfn Editor-in-Chief: Dr. Dwayne Lepitzki Assistant Editor: Dr. Amanda Martin Copy Editors: Sandra Garland and Dr. John Wilmshurst Layout: Robert Forsyth Book Review Editor: Dr. Barry Cottam Online Journal Manager: Dr. Bill Halliday Subscription Manager: Eleanor Zurbrigg Author Charges Manager: Ken Young Associate Editors: Dr. Ron Brooks Dr. Jennifer R. Foote Dr. Donald F. McAlpine _ Dr. Jeffery M. Saarela Dr. Carolyn Callaghan Dr. Graham Forbes Dr. Garth Mowat David C. Seburn Dr. Paul M. Catling Thomas S. Jung Dr. Marty Obbard Dr. Jeffrey H. Skevington Dr. Francois Chapleau Chair, Publications Committee: Dr. Jeffery M. 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It is available to libraries at $40 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be sent to subscriptions@canadianfieldnaturalist.ca or mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2 Canada. Canada Post Publications Mail Agreement number 40012317. Return postage guaranteed. The Thomas H. Manning fund, a special fund of the OFNC, established in 2000 from the bequest of northern biologist Thomas H. Manning (1911-1998), provides financial assistance for the publication of papers in the CFN by independent (non-institutional) authors, with particular priority given to those addressing Arctic and boreal issues. Qualifying authors should make their application for assistance from the fund at the time of their initial submission. Cover: Rock Ptarmigan (Lagopus muta townsendi) on Amchitka Island, Alaska, in late May to very early June. Male Rock Ptarmigan on mainland Alaska and northern Canada during this same period are mostly white, a difference that has been used to describe subspecies of Rock Ptarmigan. See the article in this issue by Clait Braun et al., pages 49-55. Photo: Steve Ebbert, 10 June 2015. The Canadian Field-Naturalist Characteristics of Wolverine (Gulo gulo) dens in the lowland boreal forest of north-central Alberta MICHAEL E. JOKINEN'*, SHEVENELL M. WEBB’, DOUGLAS L. MANZER?’, and ROBERT B. ANDERSON? ‘Alberta Conservation Association, 817-4th Avenue South, Lethbridge, Alberta T1J OP6 Canada *Maine Department of Inland Fisheries and Wildlife, 650 State Street, Bangor, Maine 04401 USA -Alberta Conservation Association, P.O. Box 1139 Provincial Building, Blairmore, Alberta TOK OEO Canada “Corresponding author: mike.jokinen@ab-conservation.com Jokinen, M.E., S.M. Webb, D.L. Manzer, and R.B. Anderson. 2019. Characteristics of Wolverine (Gulo gulo) dens in the lowland boreal forest of north-central Alberta. Canadian Field-Naturalist 133(1): 1-15. https://doi.org/10.22621/cfn. v13311.2083 Abstract We investigated Wolverine (Gu/o gulo) denning ecology in the boreal forest of northern Alberta. During winters 2015/2016 and 2016/2017, we used live traps to capture four female Wolverines and fitted them with global positioning system (GPS) collars programmed to take a location every two hours. We determined reproductive status at capture and GPS location data were used to identify den sites. One female denned in one of the two years, one female denned in two consecutive years, and two females did not den during the study. Seven of the eight Wolverine den sites were in mature or old Black Spruce (Picea mariana) stands, where dens consisted of a hollow, moss-covered mound originating from a partially uplifted root mass caused by a leaning or fallen tree. One den was located under decayed logging debris with an overstorey dominated by dense deciduous regeneration. Maximum snow depth recorded (December—March) at weather stations in the study area was 32-51 cm. Spring snow coverage was scarce in our study area (<1%) and always associated with ice cover on lakes and large ponds; mean distance from dens to nearest spring snow coverage was 15.19 km (SD = 2.73, n= 8). Female Wolverines appear to be using locally-available denning structures in the lowland boreal forest, despite a lack of deep snow, persistent spring snow cover, or large boulders documented in other studies. Key words: Alberta; boreal forest; den; lowlands; snow; Wolverine Introduction Wolverines (Gulo gulo) are well adapted to cold, snowy environments with their compact body, large paws, dense, frost-resistant fur, and capacity to store significant body fat (Banci 1994). Because Wolverines give birth in winter, females must find suitable den sites that are protected from predators, disturbance, cold temperatures, and melting spring snow (Magoun and Copeland 1998). Most verified Wolverine dens were under 1-5 m of snow (Pulliainen 1968; Magoun and Copeland 1998), suggesting that a deep snowpack of- fers important benefits throughout the denning sea- son (Magoun and Copeland 1998). The majority of Wolverine den locations documented around the world (n = 562 dens) overlapped areas with persis- tent spring snow; a small subset of dens that were outside this mapped area of persistent spring snow cover (hereafter, the spring snow coverage) were vis- ited and later confirmed to be snow dens (Copeland et al. 2010). Deep snow and/or persistent spring snow cover has been associated with Wolverine dens throughout ©The Ottawa Field-Naturalists’ Club their distribution (Magoun and Copeland 1998; Cope- land et al. 2010; May et al. 2012), but few dens have been described in low elevation, forested habitats. The majority of published information on Wolverine dens is from regions where deep snow was associ- ated with steep, rugged terrain, and large boulders in Norway (May ef al. 2012), woody debris and boul- ders in British Columbia (Krebs and Lewis 2000), long complex tunnels (Magoun and Copeland 1998) and drainage features in Alaska (Magoun ef al. 2017), and fallen trees or boulders in Idaho (Copeland 1996; Magoun and Copeland 1998). A Wolverine denned under large boulders and downed trees in the low-ele- vation boreal forest of Ontario (n = 1 den; Dawson et al. 2010) and females used boulder complexes in mature, mixed-coniferous boreal forests in Sweden (n = 49 dens; Makkonen 2015). Given a lack of steep terrain and large boulders, a shallow snowpack, and relatively early spring snowmelt in the lowland boreal forest of northern Alberta (Webb ef a/. 2016), it was unclear what resident Wolverines were using for den- ning structures. 2 THE CANADIAN FIELD-NATURALIST Similar to Wolverines, American Black Bear (Ur- sus americanus) gives birth in winter and need to select den sites that will keep cubs dry, warm, and safe. In the northern boreal forests, most black bear dens are excavated, typically beneath ground level, under the roots of standing or partially blown-down trees, into hillsides, or into riverbanks (Fuller and Keith 1980; Klenner and Kroeker 1990). American Black Bear dens are typically in more upland forest stands, and peatland is avoided (Tietje and Ruff 1980). We hypothesized that in northern boreal landscapes, Wolverine dens located in upland habitat with mature forest cover and deeper snowpack would provide the best protection and insulation available, while more lowland, wet areas would not be used. Although long-term fur harvests and images cap- tured at camera traps suggest a reproducing popu- lation of Wolverines in northern Alberta (Webb ef al. 2016), very little is known about denning ecol- ogy. Documenting den structures, snow conditions near dens, and duration of use, particularly in areas outside of the expected distribution of spring snow cover, could help clarify the relationship between Wolverines and snow and be useful information for timber harvest planning. Currently, Alberta’s tim- ber harvest guidelines list Wolverine dens under the “other species/sensitive site” section of the docu- ment, suggesting a forested buffer distance of 100 m (Alberta Agriculture and Forestry 2016); yet, there is no description of how to identify a potential Wolverine den. Our objectives were to: (1) document the general forest characteristics and specific struc- tures associated with Wolverine den sites; (2) char- acterize snow, land cover, and industrial disturbance surrounding Wolverine den sites; and (3) summar- ize female Wolverine movements during the denning period (February—May). Methods The study area, roughly 4600 km? in size, is lo- cated ~500 km north of Edmonton and 100 km north- east of Red Earth Creek in north-central Alberta (S7°N, 114°W; Figure 1). The landscape is typical of Alberta’s boreal region (Natural Regions Committee 2006), with a mosaic of aspen (Populus spp.)-domin- ated and aspen/White Spruce (Picea glauca (Moench) Voss) mixedwood forests in the uplands and exten- sive areas of Black Spruce (Picea mariana (Miller) Britton, Sterns & Poggenburgh) treed fens and bogs in the surrounding wetlands. Approximately 42% of the study area is comprised of wetlands (fens, bogs, swamp, open water, and marsh), which were pre- dominantly peatland forms (fens or bogs; 30% of the study area; AEP 2015). Mean elevation of overlap- ping townships within the study area is 616.98 m (SD Vol. 133 = 89.56, n = 75 townships) and ranged from 500 to 800 m. Summers are short and cool, and winters are cold with snow typically covering the ground from November to mid-April. Mean August temperature in the study area was 13.68 + 1.86 (SD) °C (mean max- imum August temperature = 18.2°C, n = 5 weather stations, 2003-2009; ACIS 2015). The study area supported low numbers of Moose (Alces americanus) and White-tailed Deer (Odo- coileus virginianus), and had a limited number of American Beaver (Castor canadensis), Gray Wolf (Canis lupus) occurred in small numbers when com- pared to other regions of the province. Caribou (Ran- gifer tarandus) are rare, but known to occur in the northern portion of the study area. American Black Bear, Canada Lynx (Lynx canadensis), American Marten (Martes americana), Fisher (Pekania pen- nanti), Ermine (Mustela erminea), Snowshoe Hare (Lepus americanus), Red Squirrel (Tamiasciurus hud- sonicus), Spruce Grouse (Falcipennis canadensis), and Ruffed Grouse (Bonasa umbellus) were common. The study area is remote and uninhabited, with little human activity due to limited access and exten- sive wetlands. The industrial footprint is small and comprised primarily of oil and gas development (e.g., all-season gravel roads, seismic lines from past exploration, and well-sites), with active forest har- vesting occurring only in the extreme southern por- tion of the study area. Many of the seismic lines had experienced considerable regrowth of alder (Alnus spp.) and other shrubs. Active wells are visited on a regular basis by oil field staff, while unmaintained wells in the area (some of which were reclaimed and having shrub regrowth) receive little to no winter visitation based on our observations while working there. Gravel road and well-site density (including active and unmaintained wells) was 0.04 km/km7? and 0.13 wells/km7?, respectively. Large wildfires were the primary disturbance in the area and approximately one-third of the study area had burned in the past 50 years (1961-2016). We used baited run pole camera traps during win- ters 2014/2015 (n = 8 run poles), 2015/2016 (n = 7 run poles), and 2016/2017 (n = 14 run poles) to docu- ment the presence of individual Wolverines based on unique markings (Magoun ef a/. 2011). During win- ters 2015/2016 and 2016/2017 (November—March), we live-trapped Wolverines using 10 and 17 log box traps, respectively (Copeland et al. 1995). The run poles and live traps were spaced ~S—10 km apart and were baited with beaver carcasses. Traps were outfit- ted with TT3 trap transmitters (Vectronic Aerospace, Berlin, Germany), which instantly sent an email mes- sage via satellite communication when a trap was triggered. On the advice of a wildlife veterinarian, oO Edmonton e CJ y -P | Fl Mar-Aug 2016 | ‘ Poke a = ||| Perennial lake ee Wildfires: 1961 - 2016 115° W JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 3 }| F2 Nov—Jan 2017 ee: OL 4, F3 Mar—Apr 2017 114° W FicurE 1. Wolverine (Gulo gulo) den locations (stars) and 100% minimum convex polygon home ranges for three female Wolverines from 2015-2017 in north-central Alberta, Canada (inset). Wolverines were immobilized using a jab stick (Dan-Inject, Borkop, Denmark) loaded with keta- mine hydrochloride, 100 mg/ml (Ketalean; Bimeda- MTC Animal Health Inc., Cambridge, Ontario) and medetomidine hydrochloride, 1 mg/ml (Cepetor; Mod- ern Veterinary Therapeutics, Miami, Florida, USA) at a dosage of 10.6-11.9 mg/kg and 0.1-0.12 mg/kg, respectively. Wolverines were equipped with Tellus ultralight global positioning system (GPS) collars (Followit, Lindesberg, Sweden) that were programmed to take a location every two hours. Atipamezole hydrochloride 5 mg/ml (Revertor; Modern Veterinary Therapeutics) was hand injected to reverse the effects of the sedative. The animals were returned to the trap on a bed of spruce boughs until fully recovered and then released. Collars uploaded data to a secure website via sat- ellite communication, but there was typically a 2-3 day time lag until locations became available. We vis- ually inspected GPS collar data to identify potential reproductive den sites. Potential dens had a repeated pattern of collar locations within 100 m of each other and movements to/from a localized area, in addition to associations with long periods of GPS time-outs when we assumed females were underground in the den and satellites were not able to get a fix (February—April). The primary den was the first den we documented and secondary dens were subsequent dens used by female Wolverines (Makkonen 2015). We used the terms primary and secondary dens, similar to Makkonen (2015), because collaring sometimes occurred after kits were born; therefore, we could not be certain that the primary den was actually the natal den. We used a geographic information system (ArcMap 10.4, Esri, California, USA) for all spa- tial calculations. We created a 5 km buffer around each den (estimated average female home range dur- ing the denning season; Makkonen 2015) and calcu- lated density of gravel roads and well-sites (active and unmaintained). We measured distance of each den site to nearest gravel road and well-site rounded to the nearest whole number. We used multiple sources of data to characterize the study area climate. During winter 2016/2017, we established winter weather sta- tions (n = 12) that were 10—20 km apart to measure local climate variables throughout the study area. Air temperature was recorded every hour using a Kimo KT50 compact temperature logger (Chevry-Cossigny, Seine-et-Marne, France). The temperature logger was not able to record temperatures below —40°C; how- 4 THE CANADIAN FIELD-NATURALIST ever, these were infrequent events. Snow depth was recorded by field staff on a weekly to biweekly basis using a stationary metal metre stick. Study weather stations were established in areas avoiding direct sun- light and unnatural tamping, drifting, or interception of snow. In addition to the winter weather stations we established, we summarized long-term (2005-2017) mean monthly temperature (°C) at the nearest (<20 km) five government-maintained weather stations surrounding the study area (1.e., Trout Mountain/ Peerless Lake, Chipewyan Lake, Loon River, Panny River, and Picadelly; ACIS 2015). We used the spring snow coverage data from Copeland et al. (2010), which was estimated across the Wolverine’s circumboreal range using MODIS, to classify 500 x 500 m pixels over seven years (2000— 2006). For each year, pixels received a one when the raster image was classified continuously as snow without any bare ground during the approximate end of the Wolverine denning period (24 April—15 May), the total number of years with continuous snow cover until mid-May was summed to get a value between one and seven for each pixel (Copeland ef al. 2010). We created a 5 km buffer around each den site and calculated percent of area with spring snow coverage. We also used snow depth data from the Canadian Meteorological Centre (CMC) which was derived using interpolation models that incorporated actual daily snow measurements from weather stations, meteorological aviation reports, and special aviation reports from the World Meteorological Organization information system (Brasnett 1999; Brown and Brasnett 2010). We summarized long-term (1998— 2014) mean monthly snow depths for CMC locations within our study area. We also inferred snow condi- tions using remote cameras and ground and aerial observations during a field visit in April 2017. We created a 500 m buffer around each den site to characterize upland land cover (circa 2010; Castilla et al. 2014) and wetlands (circa 2015; AEP 2015). Land cover near dens included coniferous forest, broadleaf/deciduous forest, mixed forest, grassland, and shrubland. Wetland classes near dens included swamp, fen, and bog. We also overlapped den sites with the Derived Ecosite Phase, which is a represen- tation of the vegetation, soil, and moisture condi- tions (wetland and upland; Figure 2) based on Alberta Vegetation Inventory and LiDAR (circa 2017; Alberta Agriculture and Forestry 2017). We collected additional details related to forest structure and ecological classification at den sites during November and December 2017. Forest struc- ture data were collected at five, 5.64 m radius plots. One plot was established at the den and the additional four plots were 30 m from the den in the four cardinal Vol. 133 directions. Plot trees were identified to species and diameter at breast height was measured using a steel diameter tape for all trees >5 m in height. Tree heights were measured with a clinometer and tree ages were determined using an increment borer, typically from the two trees having the largest diameter within each plot. We defined stand age class as young (20—49 years), mature (50-119 years), and old (=120 years), similar to Stelfox (1995). Each plot was classed to an ecosite phase, which is an Alberta-based field guide that subdivides forest types using site characteris- tics (moisture and nutrient regime), plant community type, soil type, and forest productivity information (Beckingham and Archibald 1996). Internal den dimensions were strictly based on a visual estimate as we did not want to enlarge the entrance and alter den structures. Means + SD are reported for all parameters, un- less otherwise indicated. Results The nature of the terrain (bogs and extensive wetlands), limited our ability to operate in the field beyond March. The transition from frozen to thawed ground occurred quickly (early April) and access to our remote field camp and the bulk of the study area was impractical. Therefore, we were unable to collect weather station data or monitor dens and litters dur- ing the denning months of April and May. Female Wolverines We captured four females (Fl, F2, F3, and F4) over two winters (2015/2016 and 2016/2017). A year prior to our live captures, we identified F1 from cam- era images at run poles. Fl’s home range during the denning season (March—May) was similar (859 km7?, 100% minimum convex polygon [MCP], 1 = 746 locations) to her overall home range from 21 March to 2 August 2016 (869 km’, 100% MCP, = 1046 loca- tions; Figure 1). We had no evidence that she was lac- tating or denning. We captured F2 during two winters. F2’s home range 19 December 2015 to 15 May 2016 was 2254 km? (100% MCP, n = 1642 locations; Figure 1), which was similar to her home range during the denning sea- son that year (February—May; 2219 km’, 100% MCP, n = 1163 locations). She showed no sign of lactation or denning during this first winter and we suspect that she may have been a young female that ultim- ately took over the neighbouring FI’s territory. We recaptured F2 on 30 November 2016 and again on 21 February 2017, and discovered she was lactating. Based on her subsequent movements and GPS time- outs, we believe she gave birth to her first litter of kits on or shortly after 22 February 2017, ~9 km from where she was captured. Her collar largely timed-out JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST Bi) Wetlands * Wolverine dens FiGurE 2. Upland and wetland matrix surrounding F3 (a) and F2 (b) den locations during 2016 and 2017 in north-central Alberta—only one den (F3 den 2, 2016) was within the upland category. over a week-long period starting on 23 February and continued to time-out on a regular basis over the next several weeks. We monitored her movements until 9 April 2017 (premature collar failure) and documented her primary and secondary den (Figure 2b). F2 dem- onstrated strong fidelity to the primary den during the first four weeks (Table 1). The greatest movement she made was ~12.5 km from her secondary den on 23 March 2017 to a location she had visited earlier in the winter, to feed on remnants of an American Black Bear hide. F2 used her primary den 22 February—13 March and secondary den until at least 26 March; the distance between the two dens was ~700 m. We suspect that F2 had another den (26 March—9 April), but we were not able to locate a third den when we searched a cluster of locations in late April. At a loca- tion where F2 had spent time (1-9 April), we did find three mounds of dead spruce limbs that had recently been broken off the lower section (~0.8 m) of spruce trees. The breaking and piling of limbs appeared deliberate and bed-like, similar to the observation reported by a Finnish Wolverine hunter/trapper in Pulliainen (1968). F2’s home range from November to January was 484 km’(100% MCP, n= 555 locations), and 90 km? (100% MCP, n = 309 locations) while she was denning (February—April; Figure 1). F2’s mean daily movements from February to April were <5 km, providing further evidence of raising young in 2017, especially when compared to the previous year when she did not raise young and her daily movements were 8-15 km during the same time period (Table 1). We captured F3 during two winters. During the first winter, our staff set up trail cameras close to her secondary den and she moved immediately after- wards to a third den (Figure 2a); camera images documented F3 and her three kits leaving the den on the evening of 19 April 2016. Based on this experi- ence, we chose not to visit female den sites during the denning period as we would not be able to deter- mine whether the use of multiple dens was natural or influenced by researchers. We did receive collar loca- tion data for F3 after 4 May 2016, which was the last day she occupied den 4. Over the next 27 days, F3 Spent six days at one GPS cluster location and three TABLE 1. Summary of daily movements (number of days, mean + SD km) made by female Wolverines (Gulo gulo) in each month of the denning season during 2016 and 2017 in north-central Alberta. Female_Year February Fl 2016 — — 11 F2 2016 29 8.22+7.12 31 F2 2017 7 2.674 4.22 31 F3_ 2016 — — 9 F3 2017 — — 10 March April May 8.58 + 6.10 30 11.18 + 8.80 31 =: 11.19+5.13 10.92 + 7.26 30 15.19+ 8.16 15 13.03+4.95 3.62 + 5.03 8 458+43.31 — — 6.89 + 6.65 30 10.23 + 8.50 31 8.48+8.12 911+45.64 23 13.83 + 10.69 — — 6 THE CANADIAN FIELD-NATURALIST days at two additional locations but these remaining GPS clusters were not visited. F3’s home range dur- ing the denning season (March—May) was 315 km? (100% MCP, n = 500 locations), which was similar to her overall home range 22 March—16 August 2016 (338 km’, 100% MCP, n = 878 locations; Figure 1). We recaptured F3 on 21 March 2017 and she was lac- tating; indicating that F3 had litters in two consecu- tive years. F3 used her primary den until 9 April and then occupied a secondary den 400 m away until at least 23 April (premature collar drop; Figure 2a). Her home range 21 March—27 April 2017 was 406 km? (100% MCP, n = 193 locations; Figure 1). Distance between F3’s 2016 and 2017 primary dens was ~8 km. Although home range size was similar between years, F3 moved further distances in 2017 compared to 2016, and daily movements were much greater than the denning F2 (Table 1). We captured F4 twice during March 2017, but she showed no signs of lactation in camera images or while we handled her 2 March and 26 March 2017. Her collar malfunctioned and we were not able to determine home range. Den descriptions Because we did not not disturb females while dens were in use and due to the challenges of work- ing in the study area during April and May, most den sites were confirmed the following winter season. We found that using repeated patterns of GPS collar locations in combination with long periods of GPS sid . . Vol. 133 time-outs to be an effective method of estimating den site locations. In 2016, F3’s primary den was 20 m off our estimated GPS location and was confirmed with fresh Wolverine tracks leading in and out of the den. F3’s secondary den in the regenerating cutblock was 10 m off our estimated location and was confirmed by very high frequency (VHF) signal and trail camera images. In 2017, F2’s primary den was 10 m off our estimated location and was confirmed with packed snow/paths leading into the den. The remaining den locations that were visited the following season were an average of 21 m off the point derived from GPS clusters and time-outs. Alternative den structures in the immediate area of the estimated den locations were limited. Seven of the eight Wolverine dens (n = 3 primary, n=5 secondary) were in the hollow created by a par- tially uplifted root mass (i.e., root ball, root wad; here- after uplifted root mass) of a leaning or fallen spruce tree. Seven of eight dens were located in mature (S0— 119 years) or old (+120 years) Black Spruce stands. Two of the seven dens were in mossy formations ori- ginating from an uplifted root mass where the trees had decayed, while the other dens were braced by the roots of intact leaning or fallen spruce trees. Root mass dens require little to no excavation by a Wolverine because a natural cavity is created when a thick moss blanket separates from the soil below as the shallow roots of a leaning or fallen tree upheave. Essentially, the lateral roots form the skeleton of the den, which supports a dense mat of soil and moss FiGurE 3. Wolverine (Gulo gulo) F3 2016 primary den was ina partially uplifted root mass of a leaning spruce tree. The den entrance 1s located along the upper side of the tree trunk in the centre of the den cavity. Photo: Michael Jokinen. JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 7 ie uF ee ir ris Ficure 4. An example of a Wolverine (Gu/o gulo) den underneath a partially uplifted root mass (F3 den 4, 2016) in the low- ils = land boreal forest of north-central Alberta. The den entrance is located at the exposed root; the tree is lying on the ground (upper right) while the lateral roots opposite the entrance have curved, creating a natural cavity. Photo: Michael Jokinen. creating the den walls (Figures 3 and 4). It is import- ant to note that these root dens are not wind throw trees characterized by roots that have been pulled out of the ground and are left standing on end. Such trees also existed within the study area, but exposed stand- ing roots do not create the mound and associated cav- ity that the Wolverines used in our study. Estimated internal den dimensions were slightly variable in size, but den size was ultimately deter- mined by the extent of the root heave (~1 m x 1 m). A soccer ball-sized opening (~30 cm) often created the den entryway and most dens had alternate openings or potential escape routes in the walls. No material was brought into the dens by Wolverine, but spruce cone bracts often lined the floors. Cone seeds have been reported in Wolverine scat (Copeland 1996). However, we observed Red Squirrel caching of intact cones and cone feeding sites, where stripped cones lay beside piles of cone bracts in and around the den location. We did not observe animal remains or Wolverine scat inside or outside the dens. Snowshoe Hare sign was widespread around denning areas when we visited sites in November and December 2017. One of the eight dens was located under decayed logging debris, which appeared to have been within or adjacent to a landing area used during previous forest harvesting activities. At the time of the ob- served den use, the overstorey was dominated by dense deciduous regeneration; the landing area and within-block roads were no longer apparent on the ground. We estimated that the cutblock was 27 years old based on tree aging and historical imagery from Google Earth (Google, Mountainview, California, USA), which suggested that the block was harvested in 1987. We could not determine the interior charac- teristics of this den without destroying the integrity of the structure. Ecosite classification at den sites Three primary and three secondary dens were re- visited in November and December of 2017 to col- lect forest structure data. Two of F3’s secondary dens from 2016 were not included in this den site forest assessment; however, they were similar in struc- ture (uplifted root mass) and were dominated by old spruce forest based on observations made dur- ing a September 2016 visit. The study area is located 8 THE CANADIAN FIELD-NATURALIST within the transition zone of the Boreal Mixedwood and Boreal Highlands ecological areas (Beckingham and Archibald 1996). The Boreal Mixedwood and Highlands are ecologically similar, but the Highlands are slightly cooler (1.7°C cooler in summer) and have higher precipitation in both summer and winter (28 mm higher in summer; winter comparison not avail- able; Beckingham and Archibald 1996). Based on the ecosite field guide of Beckingham and Archibald (1996), three of five dens (not includ- ing the den in the regenerating aspen stand) were an ecosite of Common Labrador Tea (Rhododendron groenlandicum (Oeder) Kron & Judd)/horsetail (Equisetum spp.) in the Boreal Mixedwood ecological area and two were an ecosite of horsetail and White Spruce in the Boreal Highlands. Of the three Boreal Mixedwood den locations, all sample plots but one (treed poor fen) were identified to a Labrador Tea/ horsetail phase. The most common indicator spe- cies that we found at these ecosites included Black and White Spruce, alder, Labrador Tea, and horse- tail. All but one sample plot (Labrador Tea-hygric Black Spruce-Jack Pine (Pinus banksiana Lambert)) at the two dens located in the Boreal Highland eco- logical area were identified to a horsetail and White Spruce ecosite phase. Indicator species at the two dens in this ecosite were similar to those found at Boreal Mixedwood ecosites. The conifer forest bor- dering the regenerating deciduous cutblock, in which F3’s secondary den was located, appeared to consist primarily of a Labrador Tea/horsetail ecosite class of the Boreal Mixedwood. Table 2 lists the tree spe- cies, count and average tree diameter, height, and age measured at sample plots. Disturbance, land cover, and climate The elevation of dens ranged from 535 to 687 m above sea level (601.5 + 52.6 m, n = 8; Table 3). F2’s dens were at similar elevations to the mean eleva- tion of the surrounding township (681 m). F3 denned in the same township over two consecutive winters (township elevation = 557 m). Dens were typically far Vol. 133 from roads and wells (Table 3); however, that could simply reflect available habitat within the study area. Gravel road density within a 5 km buffer of each den was 0—0.18 km/km/? (0.07 + 0.08 km/km?, n= 8 dens). Well density (active and unmaintained) within the 5 km buffer of each den was 0.04—0.08 wells/km/? (0.06 + 0.02 wells/km?, n = 8 dens). Conifer forest was the dominant land cover within the 5 km buffer for six of the eight dens (range: 50—100%). One den was 54% deciduous for- est, 21% mixed forest, 21% conifer forest, and 3% shrub within this buffer area. The area surrounding the logging debris den had been classified as 65% shrub (primarily regenerating Populus spp. within a cutblock), 30% conifer forest, 3% grassland, and 2% deciduous forest; however, the regenerating cut- block had reached heights >10 m by 2016. There was a wide range in the amount of wetland within 500 m of each den (range: 10-74%; 33.7 + 21.28%, n = 8 dens). The wettest den (74% wetland) was classi- fied as 53% swamp, 18% fen, and 3% bog within 500 m (F2’s primary den). Six of the eight dens, however, had 10-35% wetland (mostly peatlands) within 500 m. Moreover, based on the Derived Ecosite Phase data, six of eight dens fell within the wetland cat- egory. This category is described as hydric/poor dominated by shrubby, treed bog vegetation (Alberta Agriculture and Forestry 2017). Wolverine dens were 4—7 km to the nearest study weather station. Mean snow depths for each month were 32.4 + 12.6 cm in December, 37.6 + 11.1 cm in January, 41.4 + 14.7 cm in February, and 34.0 + 17.8 cm in March (n = 12 stations; Table 4). Maximum snow depth recorded (December—March) at individ- ual weather stations was 32-51 cm. Hourly temper- atures in the study area increased by the latter half of March (16-29 March, daily —3.6°C), as compared to the first half of the month (1-15 March, —16.8°C). Mean monthly temperatures increased slightly with each month, while monthly ranges were highly vari- able: December —14.4 + 6.8°C (range —36.0 to 2.9°C), TABLE 2. Forest stand structure (count, mean + SD) associated with Wolverine (Gu/o gulo) dens (n = 6) during 2017 in the lowland boreal forest of north-central Alberta. Den Tree DBH* (cm) Tree height (m) Tree age (yrs) Stem count? F3_1_ 2016 32. + 491+4204 10 76247 10 116.9+44.9 23 Sb, 8 Sw, 1 Lt F3_ 2 2016 83." (33 2105 10 1A S35 1 26622" .0:7 19 Pb, 18 Aw, 15 Sw F2_1 2017 113° 23.4+12.2 10 13.4431 10" 705: 4227 87 Sb, 19 Sw, 7 Lt F2 2 2017 84 3064159 10 16.0+4.1 10 85.8+4287 51 Sb, 22 Sw, 10 Lt F3_1_ 2017 70 = =42.84257 10 190+6.1 LO 121 9 375 70 Sb H3: 2.2017 40 576+25.5 10 24.2442 10 114.6+ 18.3 40 Sb *Diameter at breast height (DBH). ‘Trees in plot >5 m tall. Species: Trembling Aspen (Aw; Populus tremuloides Michaux), Balsam Poplar (Pb; Populus bal- samifera L.), Black Spruce (Sb; Picea mariana (Miller) Britton, Sterns & Poggenburgh), Tamarack (Lt; Larix laricina (Du Roi) K. Koch), and White Spruce (Sw; Picea glauca (Moench) Voss). 2019 JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 9 TABLE 3. General summary of Wolverine (Gu/o gulo) dens found in the lowland boreal forest during 2016 and 2017 in north- central Alberta. Entrance aspect Den Date occupied Elevation (m) F3_1_ 2016 mid Feb—9 Apr* 561 S F3_2 2016 10 Apr—19 Apr 590 S F3_3_ 2016 20 Apr—23 Apr 607 E F3 4 2016 24 Apr—4 May 615 NW F291 2017 22 Feb—13 Mar 673 N F2. 2.2017 14 Mar—26 Mar 687 SE F2 3 2017. 27 Mar—9 Apr? _ = F3 1 2017 mid-Feb—9 Apr* 544 W F3_2 2017 10 Apr—23 Apri 535 S Nearest road (km) Nearest active wellsite (km) 2.0 2.0 0.4 0.4 1.0 0:9 1.0 0.9 1250) 10.4 12.0 10.3 10.0 10.9 10.0 11.0 *F3 denning start date is approximate, as she was collared after kits were born in both instances. *Collar failure or premature collar drop. {Unconfirmed den location. January —14.0+ 10.2°C (range —40.0 to 10.5°C), February —12.7+ 10.1°C (range —40.0 to 19.2°C), and March —10.6+ 10.8°C (range —38.8 to 14.4°C; Table 4). Mean monthly temperatures were similar between long-term data from nearby government stations (2005-2017) and monthly study station temperatures measured during winter 2016/2017 (Table 4). CMC model grid points were 7-13 km from Wolverine den sites and indicated that snow depths are typically shallow in our study area (December-— March, 21.66 + 1.77 cm, range 19.74—25.03 cm, n = 10 stations; Brown and Brasnett 2010). Snow depths interpolated for points within the study area were slightly higher than mean monthly snow depth trends in the boreal forest of Alberta (February: 25.57 cm, March: 24.24 cm, n = 686 stations; Webb ef al. 2016). Spring snow coverage (Copeland et a/. 2010) was limited (0.38%) and patchy (mean size 1.6 + 2.68 km’, n = 11 patches) in our study area. There were no instances of spring snow coverage predicted near Wolverine dens. Mean distance from dens to nearest spring snow coverage was 15.19 + 2.73 km (n = 8). All patches of the spring snow coverage in the study area corresponded to lakes or large ponds that would be expected to retain at least some ice cover beyond when snow in the forest had melted. We used trail cameras to document spring snow conditions for F3’s primary and secondary dens in 2016. Her primary den was completely snow-cov- ered on 30 March 2016 and 20 days later the snow had all melted. There was no snow cover surround- ing the area of F3’s secondary den on 19 April 2016. We visited the study area 25—27 April 2017 to retrieve dropped radio collars and observed patchy snow cov- er across the entire region, from the air (Figure 5) and on the ground. We used an Argo (New Hamburg, Ontario, Canada) to access the area of F2’s primary den (2017) as she had not used this den for several weeks and patchy snow cover was encountered at the time. We did not locate F2’s secondary den until November 2017 as we were not confident that she was finished using the den during our April visit. We flew over (Figure 5) and hiked within 1 km of F3’s 2017 dens while retrieving her dropped radio collar and encountered sparse snow cover throughout the area. Discussion Wolverine pregnancy is largely dependent on body TABLE 4. Mean temperature and snow depths (+ SD) recorded at study weather stations (2017), government weather sta- tions (2015-2017), and Canadian Meteorological Centre (CMC) locations (1998-2014) during December—May in north- central Alberta. Weather station December January Study stations Temperature (°C) -144+6.8 -14.0+ 10.2 n=12 Snow depth (cm) 32.4+ 12.6 37.6+ 11.1 n=12 Government stations and CMC estimates Temperature (°C) =15 3) 2057 -18.1+0.8 n=5 Snow depth (cm) 147+1.2 213 165 n= 10 February March April May A272 10,1 S10,622510.8 — _ 4144147 34.0+ 17.8 — — —13.9+0.8 7 oO QS £02 8.7+40.2 26.5+2.1 24.2426 5.6+1.2 0.2+0.1 10 THE CANADIAN FIELD-NATURALIST Vol. 133 FiGureE 5. Snow cover near Wolverine (Gu/o gulo) F3 primary den on 27 April 2017 in north-central Alberta. Photo: Michael Jokinen. condition and winter food availability (Persson 2005), because of delayed implantation (Banci 1994). It has been hypothesized that dens that provide females and their offspring with secure shelter from disturbance (e.g., predation, weather, people), thermal insulation (Magoun and Copeland 1998), and access to adequate food resources (Inman et a/. 2012) may be more likely to produce successful litters. Nearly all documented Wolverine dens in the world have been associated with deep snow (Magoun and Copeland 1998) and/or persistent spring snow cover (Copeland et al. 2010), however, Wolverines have not been studied equally across their range (Banci 1994), particularly in North America. We recognize that our sample size of den- ning females was small and that reproductive suc- cess was not measured for those females; however, our study detected a Wolverine denning strategy that is largely undescribed. We documented Wolverine dens in low elevation forests lacking boulders and deep or persistent spring snow, where a core, resi- dent population has supported Wolverine harvests for over 30 years (Webb et a/. 2016). Our results pro- vide further evidence that Wolverines are adapted to exploiting cold, low productivity environments, but females appear to be selecting denning habitat that differs from what we hypothesized and what has been reported elsewhere. In addition to shallow snow cover, our study area had other unique differences from other Wolverine studies. Ungulates can be important in the diet of female Wolverines (Banci 1994; Inman ef al. 2012), yet ungulates were in low abundance in our study area and in much of the boreal forest, where smaller prey including American Beaver, Snowshoe Hare, and grouse are more common. Female Wolverine in north- ern British Columbia were positively associated with rugged terrain in alpine environments, where Hoary Marmot (Marmota caligata) and Columbian Ground Squirrels (Urocitellus columbianus) were common (Krebs et al. 2007). Although the Omineca region of British Columbia is at similar latitude, our study area does not support this prey or terrain selection. Not unlike the difference between northern mountain and boreal ecotypes of Woodland Caribou (Wood and Terry 1999; ASRD & ACA 2010), Wolverines in our study area must meet their needs in a very differ- ent environment. Although we lacked data on win- ter food availability, we documented one female that 2019 denned in two consecutive years, with three kits con- firmed to be alive at ~4—6 weeks of age in the first year. Snowshoe Hare and Canada Lynx sign was common during our study. Based on Canada Lynx harvests, Snowshoe Hare cycle peaks in Alberta have occurred around 1980, 1990, 2000, and 2010 (Webb et al. 2013). Snowshoe Hare numbers were increasing during our study (N. Kimmy pers. comm. 30 January 2019). The habitat within our study area was highly mosaic, likely a result of frequent fires and abun- dant wetlands. Female Wolverines may rely on hunt- ing small prey, such as Snowshoe Hare (Banci 1994; Scrafford and Boyce 2015), and this varied land- scape may provide hares the forage, concealment, and thermal cover to persist in relatively good num- bers throughout the various habitat types (Hodges 2000; Gigliotti et a/. 2018). Krebs et al. (2018) state that the Snowshoe Hare is one of the few prey species available to predators during the winter in the bor- eal forest. All avian and mammalian predators in the boreal forest eat Snowshoe Hare (Krebs ef al. 2018). Wolverine and Canada Lynx harvest data have shown that a pattern may exist between Wolverine harvest and the Snowshoe Hare cycle (Webb ef al. 2013; Boonstra et al. 2018). By denning within mature con- ifer, female Wolverines in the boreal forest may have access to a prey source in close proximity. Inman et al. (2012) suggest there may be a con- nection between food storage, persistent spring snow cover, and Wolverine denning requirements. If deep snow may provide an opportunity for food cach- ing in other settings, it begs the question: How are Wolverines meeting this need in a landscape where snow is far less abundant? We do not have the data required to answer this question, but local knowledge may have provided a hypothesis worth testing with future studies. On three independent occasions, trap- pers in our study area observed a Wolverine having depredated a harvested Canada Lynx from a trap, bringing it into an adjacent peatland area, and cach- ing it. In each case, the trapper reported seeing that the Wolverine had dug through the snow and down into the organic peat layer, then buried the carcass up to 45 cm below the surface with a mixture of snow, moss, and other vegetation. In one case, the trapper reported that the Wolverine had urinated on top of the location before leaving; another reported finding it very challenging to dig into the cache as the infill had frozen solid. Scrafford and Boyce (2015) also documented Wolverines caching in bogs in northern Alberta. We observed instances where it appeared that a wolverine had returned to a peatland cache and excavated and fed on food remnants (M.E.J. unpubl. data). Burying foods into bogs may help preserve excess food for later use (Verhoeven and Liefveld JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 11 1997, Moldowan and Kitching 2016) or hinder com- petitors from locating it. Future research into boreal Wolverine ecology should seek to test this hypothesis. Wolverine dens have been documented under wind-drifted snow, large boulders, and trees in areas with deep snow (Magoun and Copeland 1998; Krebs and Lewis 2000; Copeland et al. 2010; May et al. 2012; Makkonen 2015; Magoun ef a/. 2017), but these features are lacking in the boreal forests of north- ern Alberta. Instead, most dens in our study (n = 7) were under partially uplifted root masses of leaning or fallen trees in older spruce forests, while one den was under decayed logging debris in an ~30 year old regenerating deciduous forest. We realize that our sample size of two denning females and their choice of denning structure could be a result of individual preference. However, Scrafford and Boyce (2015) also found Wolverines denning in an uplifted root mass and timber slash pile near Rainbow Lake in north- western Alberta. Approximately 42% of our study area is comprised of various wetland forms, includ- ing a majority made up by peatlands, with a mean ele- vation of 600 m. Makkonen (2015) notes that no dens were found in peat bogs, despite their abundance on the boreal landscape in Sweden, but Wolverines had access to and used large boulders at higher elevations for denning. Pulliainen (1968) found that half of the Wolverine dens in the boreal forest of Finland were associated with standing or fallen spruce trees; how- ever, the dens and tunnels were established under the length of a fallen tree and were always under deep snow cover (>I m). In contrast, maximum snow depth in our study rarely exceeded half a metre and was meaningfully absent for the final third of the denning season. American Black Bears use a variety of den struc- tures across their range, but adequate thermal cover is critical for successful reproduction in northern cli- mates. The most common black bear den was under the roots or stumps of standing or partially blown down trees in the boreal forest of Ontario (Kolenosky and Strathearn 1987), Manitoba (Klenner 1982), and Alberta (Tietje and Ruff 1980), and the den cham- ber was similar in size to what we measured inside Wolverine dens (~1 m3). Contrary to black bear dens, however, the Wolverine dens we investigated were not deliberately lined with other materials (e.g., grass, moss, leaves, twigs; Klenner 1982). Instead, most of the Wolverine dens had cone bracts inside that had been discarded by feeding Red Squirrels. Squirrel middens have also been associated with marten den sites (Ruggiero et al. 1998) and Western Toad (Anaxyrus boreas) hibernation sites (Browne and Paszkowski 2010), where it has been suggested that they may provide some thermal benefit. Marten 12 THE CANADIAN FIELD-NATURALIST will utilize root masses of fallen trees for winter rest sites (Gilbert et al. 1997) and den sites can occur underground (Bull and Heater 2000). Browne and Paszkowski (2010) note that Western Toad hiberna- tion sites in north-central Alberta were also located within peat hummocks and decayed root channels. Mosses are the prevalent ground cover in the wetland environments of the boreal forest. Instead of deep snow (>1 m) providing thermal protection (Magoun and Copeland 1998), it is possible that the thick moss layer insulates Wolverine dens from cold temperatures and excess moisture. Snow accumula- tion in our study area averages only 30-40 cm, but when combined with the thick, mossy root layer, these den structures may provide adequate thermal insula- tion. Moss was traditionally used by Laplanders and other circumpolar people for bedding and insulation in both dwellings and clothing (Kimmerer 2003). Various species of moss have been shown to have thermal properties that insulate and limit the fluctua- tion of soil temperature and moisture (Soudzilovskaia et al. 2013). Marchand (2014) suggests that under 40-50 cm of snow, air temperature fluctuations have little influence on subnivean conditions. We sus- pect that typical late winter snow depth in our study area, 1n combination with the layer of moss, may also approximate those conditions. The ecosites in which our dens were located are naturally wet and are rated as having high excess moisture (Beckingham and Archibald 1996). Even though these ecosites have elevated water tables near the ground surface, the den cavities are shallow and not far below the mossy forest floor. Because snow cover is relatively light and the den floor close to ground level, the probability of the den flooding dur- ing spring melt would be low. Wind-throw hazard (i.e., potential for trees to become partially or completely uprooted) is rated as medium-high/high for the ecosites where dens were found in our study area (Beckingham and Archibald 1996). The potential for ready-to-move-in den struc- tures in this forest type is therefore greater. The lat- eral roots and soil lining create a barrier, although the walls are relatively thin (~15—30 cm) and appear fra- gile even when snow-covered. The root mass walls provide limited protective shielding from potential danger, so females may be more susceptible to dis- turbance. However, this did not seem to result in them moving denning sites more frequently, as other stud- ies documented similar number of dens per female as we did (Magoun and Copeland 1998). The prox- imity of a den structure to potential human disturb- ance is likely important (Banci 1994). Our study area was remote and most dens were located far from roads and trails, where encounters with people would Vol. 133 be rare. In addition to potential direct disturbance at the den, Scrafford et a/. (2018) suggested that roads may negatively influence Wolverines by altering both habitat use and movement rates through habitat near roads. However, the density of roads near den sites in our study was an order of magnitude less than that of Wolverine home ranges in their study, suggesting that these females may be less impacted by roads. In addi- tion, ungulates were not abundant in our study area, so wolf numbers were not high. This may lessen the need to have a secure den structure as would be pro- vided by a snow cave or large boulders. Forest companies seeking to provide long term Wolverine denning habitat within low elevation bor- eal forests have been operating with a paucity of information, trying to determine how to apply what is known about dens from a mountain environment to one largely devoid of boulders and a deep, per- sistent snow pack. Although our observations are limited, these females, and those of the Scrafford and Boyce (2015) study, provide a glimpse into the unique denning ecology of boreal Wolverines. Until more detailed information can be obtained, forest companies should retain mature representative sam- ples of high-wind-throw-risk ecosites within their planning area. In some cases, forest harvesting may have the potential to create future suitable denning habitat when structure is left behind (e.g., brush piles, log landings). Although the availability of par- tially wind thrown trees may not be limiting on the boreal landscape, their suitability for den sites may be influenced by the degree of disturbance in the sur- rounding area. In the absence of deep snowpack, Wolverines in our study area have found a way to persist in the lowland boreal forest. Our small sample size lim- its our ability to draw robust conclusions. As such, our observations and speculation about potential eco- logical processes should be viewed as the basis for hypotheses that can be tested with further study. In a landscape lacking deep snowpack and large boul- ders, we speculate that Wolverines are able to meet their needs through locally available features such as the cavity created by a partially uplifted root mass, the thermal properties of thick moss, and the caching opportunities provided by deep peat accumulations. Wolverines are resourceful and may be more flexible in their denning requirements than documented by studies in other landscapes. Author Contributions Writing — Original Draft: M.J., S'W., and R.A.; Writing — Review & Editing: M.J., S.W., and R.A.; Conceptualization: M.J., SW., D.M., and R.A.; In- vestigation: M.J., S.W., and R.A.; Methodology: M.J., 2019 S.W., and R.A.; Formal Analysis: M.J., S.W., and R.A.; Funding Acquisition: R.A., D.M., and S.W. Acknowledgements A special thank you goes to Neil Kimmy and Bill Abercrombie (Alberta Trappers’ Association mem- bers) for assisting with project establishment. A huge thank you goes to the Kimmy family for providing field accommodation and field support, while Duncan Abercrombie and Dan Mclean of Animal Damage Control contributed with their time and expertise in the field. John Hallett, Registered Professional Forester, Alberta Conservation Association (ACA), made a special contribution by providing forestry and ecosite classification expertise and Corey Rasmussen, ACA, was an essential member of the immobiliz- ation crew. A special thank you to all ACA staff (there were many) who participated in the field. Matt Scrafford, University of Alberta, and staff at Animal Damage Control, contributed by establishing the first Wolverine live traps within the study area. Mark Boyce, University of Alberta, who has been a collab- orator on much of our Wolverine work in the province, loaned us equipment. We thank Dr. Michelle Oakley, Doctor of Veterinary Medicine (DVM), and Dr. Mark Johnson, DVM, for their immobilization training and guidance and Dr. Glenn Meyers, DVM, for providing immobilization agents. We also appreciate the spring snow coverage GIS data from Jeffrey Copeland. This study was supported by Alberta Conservation Association, Alberta Environment and Parks, Al- berta Trappers’ Association, Alberta-Pacific Forest Industries Inc., Crowsnest Conservation Society, Daishowa-Marubeni International Ltd., McGill Uni- versity, Roadrunner Leasing and Sales Ltd., Shell FuellingChange, TD Friends of the Environment, and the University of Alberta. Capture and handling protocols and run pole camera traps were approved by the following Government of Alberta Research Permit and Collection Licences (#56202, 56203, 56900, 56901, 57157, 57158, 58403, 58404). Literature Cited ACIS (Alberta Agriculture and Forestry, AgroClimatic Information Service). 2015. Weather stations data view- er. Accessed 20 January 2019. http://agriculture.alberta. ca/acis/. AEP (Alberta Environment and Parks). 2015. Alberta Wetland Classification System. Accessed 1 February 2018. https://geodiscover.alberta.ca/geoportal/catalog/ search/resource/details.page? uuid={A73F5A E1-4677- 4731-B3F6-700743A 96C97}. Alberta Agriculture and Forestry. 2016. Alberta timber harvest planning and operating ground rules framework for renewal. Alberta Forest Management Branch. Ac- cessed 29 January 2018. https://wwwl.agric.gov.ab.ca/ $department/deptdocs.nsf/all/formain15749/$FILE/ JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 13 TimberHarvestPlanning-OperatingGroundRules Framework-Dec2016.pdf. Alberta Agriculture and Forestry. 2017. Derived Ecosite Phase Version 1. Accessed 1 March 2019. https://open. alberta.ca/opendata/derived-ecosite-phase#summary. ASRD & ACA (Alberta Sustainable Resource Develop- ment and Alberta Conservation Association). 2010. Status of the Woodland Caribou (Rangifer tarandus caribou) in Alberta: Update 2010. Alberta Sustainable Resource Development. Wildlife Status Report No. 30 (Update 2010). Edmonton, Alberta, Canada. Accessed 1 March 2019. https://open.alberta.ca/dataset/OScdc28e- 5fbf-4906-9adf-eefbb26a2d1e/resource/9dd98304- Oddc-40cb-b94c-44158ca4bad8/download/4782681- 2010-status-woodland-caribou-alberta-update-2010. pdf. Banci, V. 1994. Chapter 5: Wolverine. Pages 99-127 in The Scientific Basis for Conserving Forest Carnivores: American Marten, Fisher, Lynx, and Wolverine in the Western United States. Edited by L.F. Ruggiero, K.B. Aubry, S.W. Buskirk, J.-L. Lyon, and W.J. Zielinski. U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. Fort Collins, Colorado, USA. Beckingham, J.D., and J.H. Archibald. 1996. Field Guide to Ecosites of Northern Alberta. Natural Resources Cana- da, Canadian Forest Service, Northwest Region Forestry Centre, Edmonton, Alberta, Canada. Boonstra, R., S. Boutin, T.S. Jung, C.J. Krebs, and S. Taylor. 2018. Impact of rewilding, species introductions and climate change on the structure and function of the Yukon boreal forest ecosystem. Integrative Zoology 13: 123-138. https://doi.org/10.1111/1749-4877.12288 Brasnett, B. 1999. A global analysis of snow depth for numerical weather prediction. Journal of Applied Me- teorology 38: 726-740. https://doi.org/10.1175/1520-04 50(1999)038<0726:agaosd>2.0.co;2 Brown, R.D., and B. Brasnett. 2010. Canadian Meteo- rological Centre (CMC) daily snow depth analysis data. Environment Canada, National Snow and Ice Data Cen- tre, Boulder, Colorado, USA. Accessed 1 February 2018. https://nsidc.org/data/NSIDC-0447. Browne, C.L., and C.A. Paszkowski. 2010. Hibernation sites of Western Toads (Anaxyrus boreas): characteriza- tion and management implications. Herpetological Con- servation and Biology 5: 49-63. Bull, E.L., and T.W. Heater. 2000. Resting and denning sites of American martens in Northeastern Oregon. Northwest Science 74: 179-185. Castilla, G., J. Hird, R.J. Hall, J. Schieck, and G.J. McDermid. 2014. Completion and updating of a land- sat-based land cover polygon layer for Alberta, Canada. Canadian Journal of Remote Sensing 40: 92-109. https://doi.org/10.1080/07038992.2014.933073 Copeland, J.P. 1996. Biology of the wolverine in cen- tral Idaho. M.Sc. thesis, University of Idaho, Moscow, Idaho, USA. Copeland, J.P., E. Cesar, J.M. Peek, C.E. Harris, C.D. Long, and D.L. Hunter. 1995. A live trap for wolverine and other forest carnivores. Wildlife Society Bulletin 23: 535-538. 14 THE CANADIAN FIELD-NATURALIST Copeland, J.P., K.S. McKelvey, K.B. Aubry, A. Landa, J. Persson, R.M. Inman, J. Krebs, E. Lofroth, H. Golden, J.R. Squires, A. Magoun, M.K. Schwartz, J. Wilmot, C.L. Copeland, R.E. Yates, I. Kojola, and R. May. 2010. The bioclimatic envelope of the wolverine Gulo gulo: do climatic constraints limit its geographic distribution? Canadian Journal of Zoology 88: 233-246. https://doi.org/10.1139/Z09-136 Dawson, N.F., A.J. Magoun, J. Bowman, and J.C. Ray. 2010. Wolverine, Gulo gulo, home range size and den- ning habitat in lowland boreal forest in Ontario. Cana- dian Field-Naturalist 124: 139-144. https://doi.org/10. 22621 /cfn.v124i2.1052 Fuller, T.K., and L.B. Keith. 1980. Summer ranges, cover type use and denning of black bears Ursus americanus near Fort McMurray, Alberta, Canada. Canadian Field- Naturalist 94: 80-83. Accessed 1 February 2018. https:// biodiversitylibrary.org/page/28088961. Gigliotti, L.C., B.C. Jones, M.J. Lovallo, and D.R. Die- fenbach. 2018. Snowshoe hare multi-level habitat use in a fire-adapted ecosystem. Journal of Wildlife Manage- ment 82: 435—444. https://doi.org/10.1002/jwmg. 21375 Gilbert, J.H., J.L. Wright, D.J. Lauten, and J.R. Probst. 1997. Den and rest-site characteristics of American Marten and Fisher in northern Wisconsin. Pages 135— 145 in Martes: Taxonomy, Ecology, Techniques, and Management. Edited by G. Proulx, H.N. Bryant, and P.M. Woodard. The University of Alberta Press. Edmonton, Alberta, Canada. Hodges, K.E. 2000. The ecology of Snowshoe Hares in northern boreal forests. Pages 117-161 in Ecology and Conservation of Lynx in the United States. Edited by L.F. Ruggiero, K.B. Aubry, S:W. Buskirk, G.M. Koehler, C.J. Krebs, K.S. McKelvey, and J.R. Squires. Depart- ment of Agriculture, Forest Service, Rocky Mountain Research Station. Fort Collins, Colorado, USA. Inman, R.M., A.J. Magoun, J. Persson, and J. Mattis- son. 2012. The wolverine’s niche: linking reproductive chronology, caching, competition, and climate. Journal of Mammalogy 93: 634-644. https://do1.org/10.1644/ 11-mamm-a-319.1 Kimmerer, R.W. 2003. Gathering Moss. Oregon State Uni- versity Press, Corvallis, Oregon, USA. Klenner, W. 1982. Seasonal movements, home range util- ization, and denning habits of black bears (Ursus ameri- canus) in western Manitoba. M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba, Canada. Klenner, W., and D.W. Kroeker. 1990. Denning be- havior of black bears Ursus americanus, in western Manitoba. Canadian Field-Naturalist 104: 540-544. Accessed 1 February 2018. https://biodiversitylibrary. org/page/34347100. Kolenosky, G.B., and S.M. Strathearn. 1987. Winter den- ning of Black Bears in east-central Ontario. Bears: their biology and management. Pages 305-316 in Papers from the Seventh International Conference on Bear Research and Management, Plitvice Lakes, Yugoslavia. Krebs, C.J., R. Boonstra, and S. Boutin. 2018. Using ex- perimentation to understand the 10-year snowshoe hare cycle in the boreal forest of North America. Journal of Animal Ecology 87: 87-100. https://doi.org/10.1111/13 Vol. 133 65-2656.12720 Krebs, J.A., and D. Lewis. 2000. Wolverine ecology and habitat use in the North Columbia Mountains: progress report. Pages 695-703 in Proceedings of Biology and Management of Species and Habitats at Risk. Edited by L.M. Darling. University College of the Cariboo, Kamloops, British Columbia, Canada. Krebs, J.A., E.C. Lofroth, and I. Parfitt. 2007. Multiscale habitat use by wolverines in British Columbia, Canada. Journal of Wildlife Management 71: 2180-2192. https:// doi.org/10.2193/2007-099 May, R., L. Gorini, J. van Dijk, H. Broseth, J.D.C. Linnell, and A. Landa. 2012. Habitat characteristics associated with wolverine den sites in Norwegian mul- tiple-use landscapes. Journal of Zoology 287: 195-204. https://doi.org/10.1111/j.1469-7998 .2012.00907.x Magoun, A.J., and J.P. Copeland. 1998. Characteristics of wolverine reproductive den sites. Journal of Wildlife Management 62: 1313-1320. https://doi.org/10.2307/380 1996 Magoun, A.J., C.D. Long, M.K. Schwartz, K.L. Pilgrim, R.E. Lowell, and P. Valkenburg. 2011. Integrating motion-detection cameras and hair snags for wolverine identification. Journal of Wildlife Management 75: 731— 739. https://doi.org/10.1002/jwmg.107 Magoun, A.J., M.D. Robards, M.L. Packila, and T.W. Glass. 2017. Detecting snow at the den-site scale in wol- verine denning habitat. Wildlife Society Bulletin 41: 381-387. https://doi.org/10.1002/wsb.765 Makkonen, T. 2015. Den site characteristics of female wol- verine (Gulo gulo) in Scandinavian forested landscape. M.Sc. thesis, University of Oulu, Oulo, Sweden. Marchand, P.J. 2014. Life in the Cold: an Introduction to Winter Ecology. Fourth Edition. University Press of New England, Hanover, New Hampshire, USA. Moldowan, P.D., and H. Kitching. 2016. Observation of an Eastern Wolf (Canis sp. cf. Lycaon) caching food ina Sphagnum bog in Algonquin Provincial Park, Ontario. Canadian Field-Naturalist 130: 351-354. https://do1.org/ 10.22621/cfn.v13014.1930 Natural Regions Committee. 2006. Natural regions and subregions of Alberta. Compiled by D.J. Downing and WW. Pettapiece. Government of Alberta. Pub. No. T/852. Accessed | January 2018. https://open.alberta.ca/ publications/0778545725. Persson, J. 2005. Female wolverine (Gulo gulo) repro- duction: reproductive costs and winter food availabil- ity. Canadian Journal of Zoology 83: 1453-1459. http:// dx.do1.org/10.1139/z05-143 Pulliainen, E. 1968. Breeding biology of the wolverine (Gulo gulo) in Finland. Annales Zoologici Fennici 5: 338-344. Ruggiero, L.F., E. Pearson, and S.E. Henry. 1998. Characteristics of American marten densites in Wy- oming. Journal of Wildlife Management 62: 663-673. https://doi.org/10.2307/3802342 Scrafford, M.A., T. Avgar, R. Heeres, and M.S. Boyce. 2018. Roads elicit negative movement and _habitat- selection responses by wolverines (Gulo gulo luscus). Behavioral Ecology 29: 534-542. https://doi.org/10.1093 /beheco/arx182 2019 JOKINEN ET AL.: WOLVERINE DENS IN BOREAL FOREST 15 Scrafford, M.A., and M.S. Boyce. 2015. Effects of in- dustrial development on wolverine (Gulo gulo) ecol- Sphagnum. Acta Botanica Neerlandica 46: 117-130. Webb, S.M., R.B. Anderson, D.L. Manzer, B. Aber- ogy in the boreal forest of northern Alberta. Wolverine Project Progress Report — Winter 2014/2015. Accessed 1 February 2018. http://wolverinefoundation.org/wp- content/uploads/2011/02/Scrafford-and-Boyce_2015_ Wolverine-Project-Progress-Report.pdf. Soudzilovskaia, N., P. Bodegom, and J. Cornelissen. 2013. Dominant bryophyte control over high-latitude soil temperature fluctuations predicted by heat transfer traits, field moisture regime and laws of thermal insula- tion. Functional Ecology 27: 1442-1454. https://do1.org/ 10.1111/1365-2435.12127 Stelfox, J.B. 1995. Relationships between stand age, stand structure, and biodiversity in aspen mixedwood forests in Alberta. Jointly published by Alberta Environmental Centre (AEC V95-R1), Vegreville, Alberta, and Canadi- an Forest Service (Project No. 0001A), Edmonton, Al- berta, Canada. Accessed 1 February 2018. http://cfs. nrcan.gc.ca/pubwarehouse/pdfs/19534. pdf. Tietje, W.D., and R.L. Ruff. 1980. Denning behavior of black bears in boreal forest of Alberta. Journal of Wild- life Management 44: 858-870. https://doi.org/10.2307/ 3808314 Verhoeven, J.T.A., and W.M. Liefveld. 1997. The eco- logical significance of organochemical compounds in crombie, B. Bildson, M.A. Scrafford, and M.S. Boyce. 2016. Distribution of female wolverines relative to snow cover, Alberta, Canada. Journal of Wildlife Manage- ment 80: 1461-1470. https://doi.org/10.1002/jwmg. 21137 Webb, S., D. Manzer, R. Anderson, and M. Jokinen. 2013. Wolverine harvest summary from registered trap- lines in Alberta, 1985—2011. Technical Report, T-2013- 001, produced by the Alberta Conservation Association, Sherwood Park, Alberta, Canada. Accessed 11 Janu- ary 2019. https://www.ab-conservation.com/downloads/ report_series/wolverine_harvest_in_alberta_1985-2011. pdf. Wood, M.D., and E.L. Terry. 1999. Seasonal movements and habitat selection by Woodland Caribou in the Omi- neca Mountains, north-central British Columbia Phase 1: The Chase and Wolverine Herds (1991-1994). Peace/ Williston Fish and Wildlife Compensation Program, Report No. 201. Prince George, British Columbia, Canada. Accessed 1 March 2019. http://www.env.gov. bc.ca/wildlife/wsi/reports/4737_WSI_4737_RPT_ OMINECA_1991_ 1994. PDF. Received 31 July 2018 Accepted 28 February 2019 The Canadian Field-Naturalist Note Wolf (Canis sp.) attacks life-like deer decoy: insight into how wolves hunt deer? THOMAS D. GABLE!" and DANIEL P. GABLE? ‘University of Minnesota, 2003 Upper Bufford Circle, St. Paul, Minnesota 55108 USA 23176 E Siebert Road, Midland, Michigan 48642 USA “Corresponding author: thomasd.gable@gmail.com Gable, T.D., and D.P. Gable. 2019. Wolf (Canis sp.) attacks life-like deer decoy: insight into how wolves hunt deer? Canadian Field-Naturalist 133(1): 16-19. https://doi.org/10.22621/cfn.v133i1.2044 Abstract We know of no documented observations of wolves (Canis sp.) detecting and then attacking a White-tailed Deer (Odocoi- leus virginianus) during spring, summer, or fall. We describe an observation of a wolf attacking a life-like, two-dimensional deer decoy in November 2017 near Killarney Provincial Park, Ontario, Canada. The wolf appeared to locate the decoy by sight rather than sound or scent, suggesting that the profile of a deer is sufficient to trigger an attack by a wolf. Key words: Wolf; Canis; carnivore; hunting behaviour; predation; predator-prey; White-tailed Deer; Odocoileus virgini- anus; Killarney Provincial Park White-tailed Deer (Odocoileus virginianus) are the primary prey of wolves (Canis sp.) throughout much of the southern boreal ecosystem in North America (Potvin et al. 1988; Benson et al. 2017; Gable et al. 2018). How and where wolves hunt and kill deer during winter is well understood because of the ease of observing wolf-hunting behaviour and lo- cating kill sites from the air (Mech and Frenzel 1971; Fuller 1989; Mech et al. 2015). However, equiva- lent information for the snow-free months is rare, as wolves and deer primarily co-occur in densely for- ested areas (Demma et al. 2007). For example, there are no estimates of wolf kill rates of White-tailed Deer (adults or fawns) during spring to fall, and lit- tle information exists about where and how wolves successfully hunt and kill deer during this period (Demma et al. 2007; Mech ef al. 2015). In a compre- hensive review of wolf—deer interactions, Mech et al. (2015) provided descriptions of eight such inter- actions during the snow-free season. However, all of these observations occurred after the wolf or wolves had already detected and attempted to chase deer. To our knowledge, there are no observations that dem- onstrate how wolves find deer during spring to fall. Herein, we document a wolf (Canis sp. according to Rutledge et al. 2016) attacking a life-like deer decoy that provides rare insight into how wolves locate and detect deer during this period. During the first week of November, D.P.G. was hunting White-tailed Deer on McGregor Island (46° 04'49""N, 81°35'18"W), about 2 km west of Killarney Provincial Park, Ontario, Canada. Before hunting, D.P.G. set-up a life-like, two-dimensional decoy of a squatting doe (“Estrous Betty”, Montana Decoys, Hummelstown, Pennsylvania, USA). The decoy con- sisted of a life-size photograph of a deer with an inter- nal wire frame, ~1.3 cm thick, for support (Figure 1). The decoy was oriented in an east—west direction so that profile views of the decoy could be seen from the north or south (Figure 2). D.P.G. also left doe urine (details on manufacturer not available) on a branch 1.5 m off the ground 1 m north of the decoy. At about 1515, after setting up the decoy and dis- pensing the doe urine, D.P.G. situated himself ina tree stand on a rocky point 23 m west of the decoy. The stand faced east and overlooked a 100-m wide valley dominated by mature Sugar Maple (Acer saccharum Marshall) forest between two steep rock ridges (north and south of the stand; Figure 2). On both sides of the valley at the base of the ridges were prominent deer trails running east to west. Immediately to the west of the tree stand was a dense Balsam Fir (Abies balsamea (L.) Miller) lowland. About 50 m north of the northern ridge was a 0.5—1.0 km wide channel of A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. ©The Ottawa Field-Naturalists’ Club GABLE AND GABLE: WOLF ATTACKS DEER DECOY 17 of Killarney Provincial Park, Ontario, Canada, during the first week of November 2017. Photo: Daniel Gable. water; this channel surrounds McGregor Island. The maple forest that the stand overlooked had minimal understorey for about 150 m before transitioning to marshy lowland, which abutted a small shallow cove that was connected to the main water channel. D.P.G. accessed the stand by parking his boat at the north- western opening of this cove (~300 m east by north- east of the stand). There was no snow cover during this period. The sky was overcast with moderate (8—16 km/h) winds blowing from the west/southwest. At 1600, D.P.G. noticed a wolf about 150 m east by south- east of the stand trotting along the deer trail on the southern edge of the valley (Figure 2). Given the pos- ition of the decoy and the structure and arrangement of the trees, the wolf would have been unable to see the decoy when D.P.G. first spotted the wolf. We later verified this by walking to the wolf’s location. The wolf continued at the same pace, moving east to west, until it was about 70 m southeast of the decoy (Figure 2). Without stopping, the wolf turned abruptly and started travelling directly toward the decoy. As the wolf approached, it appeared to be intently focussed on the decoy; however, it maintained a trotting pace for another 30 m. When about 40 m from the decoy, the wolf suddenly sprinted toward the decoy and, when only a few metres away, lunged at it, latching onto its neck, leaving punctures in the fabric of the decoy. The force of the contact ripped the decoy from the ground and caused the wolf and decoy to tum- ble for about 10 m (total time 2-3 s). After the wolf had stopped its fall, it promptly stood up and jumped back about 10 m. It stood looking at the decoy for a few seconds with both ears and tail lowered. Within a few more seconds, the wolf ran quickly over the steep ridge to the south and disappeared from view. We know of no other observation of a wolf trav- elling, detecting, and then attacking a deer or deer facsimile during the snow-free season. Although the decoy was not an actual deer, it looked exactly like a deer (Figure 1) and behaved (stood still staring at the wolf) as deer do when approached by predators (DeYoung and Miller 2011; Mech ef al. 2015). Given this and the observed changes in the wolf’s behaviour after it appeared to detect the deer, we believe that the wolf was convinced the decoy was a deer. As a result, we assert that the wolf’s behaviour on detecting and approaching the decoy provides insight into how this 18 THE CANADIAN FIELD-NATURALIST CC ee SPP ee LLL TL Lie Deer trails . Sanmne, . “a. Cr "es henge ntti tae, “Semennny * Vol. 133 ane a® a? Parry geneuue — gee™ eueeeee™ ” ote "egnat® Open maple Y | forest Wind direction Where wolf was first observed FiGurE 2. Route taken by a wolf (Canis sp.) that detected and then attacked a life-like decoy of a White-tailed Deer (Odo- coileus virginianus) near Killarney Provincial Park, Ontario, Canada, in early November 2017. wolf, and likely other wolves, may locate deer. The wind direction and consistent wind flow would have made the doe urine difficult, and likely impos- sible (Conover 2007), for the wolf to detect during its approach, which strongly suggests that the wolf lo- cated the decoy visually. Wolves are thought to be adept at visually detecting slight movements, which likely helps in locating prey (Harrington and Asa 2003), but our observation suggests that wolves are capable of detecting motionless prey from consider- able distances. We estimate that the wolf detected the decoy about 70 m away, although detection was likely aided by the minimal understorey and daylight conditions. Although wolves likely rely on scent to locate deer when hunting (Mech ef a/. 2015), it appears they can also use visual detection, even if not associated with odour, sound, or any other cues. Dense vegetation throughout most of wolf—deer range likely limits vis- ual detection of deer during the summer. However, events that reduce forest or understorey cover (e.g., forest fires, clear-cuts) could enhance the ability of wolves to detect deer and increase encounter rates between wolves and deer (Whittington et a/. 2011) and possibly wolf kill rates (Sand et a/. 2005; Vander Vennen et al. 2016). Mech ef al. (2015: 26) noted that “when wolves detect deer, they usually proceed slowly and deliber- ately, ever on the alert”. However, this wolfapproached relatively rapidly after detecting the decoy, closing a ~70 m distance in a matter of seconds. Once 30 m away from the decoy, the wolf apparently decided that the deer (1.e., the decoy) was indeed vulnerable, possibly because it did not move, and sprinted toward it. Wolves generally assess the vulnerability of deer by approaching, chasing, and testing them. Most deer are not vulnerable to predation because they are in sufficiently good physical condition to easily out-run and evade wolves; therefore, most hunting attempts are short lived as wolves realize their efforts are futile (Mech et al. 2015). Our observation provides the only information we are aware of about how at least one wolf approached and attacked what it thought was an adult White- tailed Deer during the snow-free season. Thus, whether the observation is the exception or represents normal behaviour is unknown. Still, it does provide new insight into the predatory behaviour of wolves. 2019 The lack of information on wolf predation of deer— both fawns and adults—during the snow-free season is Surprising given the amount of research on wolves, deer, and their interactions. Because of this, we rec- ommend intensive research on wolf—deer interactions during summer as has been done recently with cari- bou (Rangifer tarandus, e.g., Whittington et al. 2011; Latham et al. 2013; Mumma eft al. 2017). Indeed, as the range of White-tailed Deer continues to expand northward (Dawe and Boutin 2016), thereby increas- ing the area that wolves and deer co-occur, such information will only become more valuable and rel- evant for the conservation and management of both species (Latham et al. 2011). Acknowledgements We thank L. David Mech for reviewing an earlier version of this manuscript and providing constructive comments to improve it. Literature Cited Benson, J.F., K.M. Loveless, L.Y. Rutledge, and B.R. Patterson. 2017. Ungulate predation and ecological roles of wolves and coyotes in eastern North America. Eco- logical Applications 27: 718-733. https://doi.org/10.10 O2/eap.1499 Conover, R. 2007. Predator—prey Dynamics: the Role of Olfaction. CRC Press, New York, New York, USA. Dawe, K.L., and S. Boutin. 2016. Climate change is the primary driver of white-tailed deer (Odocoileus vir- ginianus) range expansion at the northern extent of its range; land use is secondary. Ecology and Evolution 6: 6435-6451. https://doi.org/10.1002/ece3.2316 Demma, D.J.,S.M. Barber-Meyer, and L.D. Mech. 2007. Testing global positioning system telemetry to study wolf predation on deer fawns. Journal of Wildlife Man- agement 71: 2767-2775. https://do1.org/10.2193/2006-382 DeYoung, R.W., and K.V. Miller. 2011. White-tailed deer behavior. Pages 147—180 in Biology and Management of White-tailed Deer. Edited by D.G. Hewitt. CRC Press, Boca Raton, Florida, USA. Fuller, T.K. 1989. Population dynamics of wolves in north- central Minnesota. Wildlife Monographs 105: 3—41. Gable, T.D., S.K. Windels, J.G. Bruggink, and S.M. Barber-Meyer. 2018. Weekly summer diet of gray wolves (Canis /upus) in northeastern Minnesota. Amer- ican Midland Naturalist 179: 15-27. https://doi.org/10. 1674/0003-0031-179.1.15 Harrington, F.H., and C.S. Asa. 2003. Wolf communica- tion. Pages 66-103 in Wolves: Behavior, Ecology, and Conservation. Edited by L.D. Mech and L. Boitani. University of Chicago Press, Chicago, Illinois, USA. GABLE AND GABLE: WOLF ATTACKS DEER DECOY 19 Latham, A.D.M., M.C. Latham, K.H. Knopff, M. Heb- blewhite, and S. Boutin. 2013. Wolves, white-tailed deer, and beaver: implications of seasonal prey switch- ing for woodland caribou declines. Ecography 36: 1276— 1290. https://doi.org/10.1111/j.1600-0587.2013.00035.x Latham, A.D.M., M.C. Latham, N.A. Mccutchen, and S. Boutin. 2011. Invading white-tailed deer change wolf—caribou dynamics in northeastern Alberta. Jour- nal of Wildlife Management 75: 204-212. https://doi. org/10.1002/jwmg.28 Mech, L.D., and L.D. Frenzel, Jr. 1971. Ecological studies of the timber wolf in northeastern Minnesota. Research paper NC-52. United States Department of Agriculture Forest Service, North Central Forest Experimental Station, St. Paul, Minnesota, USA. Accessed 18 July 2019. https://www.nrs.fs.fed.us/pubs/rp/rp_nc052.pdf. Mech, L.D., D.W. Smith, and D.R. MacNulty. 2015. Wolves on the Hunt: the Behavior of Wolves Hunting Wild Prey. University of Chicago Press, Chicago, Illinois, USA. Mumma, M.A., M.P. Gillingham, C.J. Johnson, and K.L. Parker. 2017. Understanding predation risk and individual variation in risk avoidance for threatened boreal caribou. Ecology and Evolution 7: 10266-10277. https://doi.org/10.1002/ece3.3563 Potvin, F., H. Jolicoeur, and J. Huot. 1988. Wolf diet and prey selectivity during two periods for deer in Quebec: decline versus expansion. Canadian Journal of Zoology 66: 1274-1279. https://doi.org/10.1139/z88-186 Rutledge, L.Y., J.M. Fryxell, K. Middel, B.N. White, and B.R. Patterson. 2016. Patchy distribution and low effective population size raise concern for an at-risk top predator. Diversity and Distributions 23: 79-89. https:// doi.org/10.1111/ddi.12496 Sand, H., B. Zimmermann, P. Wabakken, H. Andrén, and H.C. Pedersen. 2005. Using GPS technology and GIS cluster analyses to estimate kill rates in wolf—un- gulate ecosystems. Wildlife Society Bulletin 33: 914— 925. https://doi.org/10.2193/0091-7648(2005)33[914:ugt agc]2.0.co;2 Vander Vennen, L.M., B.R. Patterson, A.R. Rodgers, S. Moffatt, M.L. Anderson, and J.M. Fryxell. 2016. Diel movement patterns influence daily variation in wolf kill rates on moose. Functional Ecology 30: 1568-1573. https://doi.org/10.1111/1365-2435.12642 Whittington, J., M. Hebblewhite, N.J. Decesare, L. Neu- feld, M. Bradley, J. Wilmshurst, and M. Musiani. 2011. Caribou encounters with wolves increase near roads and trails: a time-to-event approach. Journal of Applied Ecology 48: 1535-1542. https://doi.org/10.1111/ j.1365-2664.2011.02043.x Received 3 February 2018 Accepted 16 January 2019 The Canadian Field-Naturalist Birds of Mansel Island, northern Hudson Bay ANTHONY J. GASTON Science and Technology Branch, Environment and Climate Change Canada, Carleton University, Ottawa, Ontario KIA 0OH3 Canada; email: tonygastonconsult@gmail.com Gaston, A.J. 2019. Birds of Mansel Island, northern Hudson Bay. Canadian Field-Naturalist 133(1): 20—24. https://doi.org/ 10.22621/cfn.v13311.2153 Abstract A recent review of bird distributions in Nunavut demonstrated that Mansel Island, in northeastern Hudson Bay, is one of the least known areas in the territory. Here, current information on the birds of Mansel Island is summarized. A list published in 1932 included 24 species. Subsequent visits by ornithologists since 1980 have added a further 17 species to the island’s avifauna. The list includes 17 species for which breeding has been confirmed and 10 for which breeding is considered prob- able. The island seems to support particularly large populations of King Eiders (Somateria spectabilis) and Tundra Swans (Cygnus columbianus) and the most southerly breeding population of Sabine’s Gull (Xema sabini) and Red Knot (Calidiris canuta, probably). Key words: Mansel Island; Hudson Bay; birds; breeding Introduction At 3180 km?, Mansel Island, Qikiqtaaluk Region, Nunavut, is the 28th largest island in Canada. It is one of three large islands in northern Hudson Bay, the others being Southampton and Coats Islands. Although the birds of Coats and Southampton Is- lands have been documented (Sutton 1932a; Gaston and Ouellet 1997), those of Mansel Island are com- paratively poorly known. Only one publication pro- vides information on the avifauna of the island: a list prepared by G.M. Sutton (1932b) based on speci- mens provided to him by A.T. Swaffield, the Hudson Bay manager who established the trading post at Swaffield Harbour, near the northern tip of the is- land, in 1929. At its nearest point, Mansel Island is 56 km from the mainland of Quebec (Figure 1). The topography is mostly low elevation (maximum 138 m), without any prominent hills or gullies except for a shallow central valley running east—west across the island. Underlying bedrock throughout is Silurian limestone, which is covered, over large parts of the island, by raised beach deposits of Holocene age. There are ex- tensive wetlands throughout, especially in the south- west portion of the island. Sutton (1932b: 41) com- mented: “an exceedingly flat, dull-gray piece of land”. Dry areas support low-growing shrubs, including willow (Salix spp.), cranberry (Vaccinium spp.), and Four-angled Mountain Heather (Cassiope tetragona (L.) D. Don), as well as the tussock forbs, Entire- leaved Mountain Avens (Dryas integrifolia Vahl) and Purple Mountain Saxifrage (Saxifraga oppositifolia L.). Marshes support extensive sedge (Carex spp.) meadows. The Hudson Bay post on the island closed in 1945, and there has been no permanent habitation on the island since then, although people from the nearby Inuit community of Ivujivik, Nunavik, sometimes visit in summer to hunt Caribou (Rangifer tarandus) and Polar Bear (Ursus maritimus; Gaston et al. 1985). Sutton’s list comprised 24 species, but only 17 of them were collected in summer and, hence, potential breeders, and no evidence of breeding was included (Sutton 1932b). Species were collected at various dates between September 1929 and June 1930. Sutton com- mented on their likely breeding status, but there was no definite evidence available to support his sugges- tions. Subsequent ground surveys, all of only one or two days’ duration, have added another 17 species to the island’s list, and breeding has been confirmed for some. Although this information is based on very brief visits, it is assembled here to give an up-to-date summary of what little is known about the avifauna of Mansel Island. Methods Subsequent to Swaffield’s collection, three ground surveys have been carried out by ornithologists. In July 1984, R. Decker visited the island for one day by helicopter, landing at several sites. Information A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. ©The Ottawa Field-Naturalists’ Club 2019 rk * : ‘Mansel Island HUDSON BAY QUEBEC ONTARIO GASTON: BIRDS OF MANSEL ISLAND DA FicurE 1. a. Location of Mansel Island in Hudson Bay. b. Localities visited in 1992 and 2016. Source: Mansel Island, Nuna- vut, 61°59'23.31"N, 79°56'12.54"W. Google Earth Pro 7.3.2.5776. Imagery date: 13 December 2015. Data provider: Landsat/ Copernicus 2018. Accessed: 30 July 2018. from his survey was incorporated into the Land Use Information Series map of Mansel Island (Environ- ment Canada 1970), which includes a list of “avian species which occur or are thought likely to occur within this map-area’, but I have only included spe- cies definitely sighted on the island during the survey. On 8 and 9 August 1992, A.J.G., V. Johnston, and I. Storm landed from the MV. Teregluk near the east- ernmost point of the island and spent 10 h ashore sur- veying an area of lakes, ponds, and marshes adjacent to a shallow bay (Figure 1). On 20 and 21 June 2016, Y. Aubry, M. Robert, F. Shaffer, and C. Marcotte carried out systematic surveys of breeding birds in two areas (Figure 1), using the protocol of the Program for International and Regional Shorebird Monitoring (PRISM; Bart and Johnston 2012). In addition to total bird counts, Species presence or absence was recorded by l-ha squares. They also touched down at several other sites to make additional observations. In addition, on 12 July 1984, an aerial survey (Cessna 337) was carried out by R. Decker along the entire coastline and over selected parts of the interior. I did not have access to the original data, but some information from this survey was incorporated into a general survey of larger birds in Foxe Basin and northern Hudson Bay (Gaston et al. 1986). Results and Discussion Combining the species listed by Sutton (1932b) with subsequent surveys yields 41 species reported from Mansel Island to date, of which definite evi- dence of breeding, in the form of nests or flightless young, has been obtained for 17 species. A further 10 Species were considered by at least one survey to be “probably breeding” (Table 1). Major concentrations of Arctic Terns (Sterna paradisaea) and Common Eiders (Somateria mollissima) were noted on the aer- ial surveys of 1984, with an estimated 1000 pairs of Common Eiders on Awrey Island and several col- onies of 50-75 pairs of Arctic Terns on the east and southwest coasts (Gaston et al. 1986). Because of the timing of surveys, breeding could be confirmed for fewer than half of the species recorded during the breeding season. The 1992 survey Vol. 133 © i Ud §q ud THE CANADIAN FIELD-NATURALIST jaaee u = aeaaoeN es ud Jsnjeys poulquio’) 22 ud ud Od SON GLUT AK StS Ss SS or oS OS OO Se ANS ee OSS SoS en CR ee oS o Ou d ud N popiossr = SMBS SADAINS WON N ie Ol SC € € uaes ‘ON Juasoid sarenbs ‘ON 910c WSTad NM ON ee Ol (6 Cc eas 18 ud Od Od ud ud snjqei1s 766] SSUNIYSIS Ol Oc OC LV udeS ‘ON ame ud (snye}js) p86l FOIE = AcEeeT UOHNS Tea Te Tea JOIN Jouruns Jouruns Jouruns Jourwuns Jouruns Jouruns Tea Jouruns Jouruns Jouruns Jouruns Jouruns Jouruns Jouruns Jouruns Jouruns Tea (uoseas) (D1AUO] D1A/]) BINA, P2T[IG-JOL (SNPNDIIBUO] SNIADAOIAAJG) 1IBIeL PI[Ie}-suo'T (SNd1JISDADA SNIADAOIAAIS) IIBICL ITYISVICY (vapsippivd DusajS) UII], DIV (SNUDIUOSYJIWS SNADT) |[NH SULLIIY (snasogiadAy snavT) [IND snoonepH (vau.inga vjlydosn) {ND AIOAT (WIgDs DUAX) |[ND S,ouIges (sniapaynf sndoapjoy) sdoreyeyg poy (snjoqo] sndoavjpy) sdoseyeyd Ppoyxoou-poy (sauduajul DiaDUualy) SUO\sUINT, Appny (buldjp siapije)) urjund (s1yjoa1asn{f s1apyv) Jodidpues padumn-syy (DUIJ1ADU SLAPIJDD) Jadidpues siding (oyjisnd siipyoD) Jadidpues poyewyediwuas (DINU S1IpIVD) UY POY (sndoapyd sniuawnn) [2a1quity A (snJpujodimas snispoADYD)) IAO]g poyewyediwias (vaIUIWOP SIJPIAN]Z) JdA0|d-UdSpyOH uesI3aWYy (vjoavjonbs sipian]q) 1A0|g Potf[oq-yoelq (42WU1 DIADD) UOO'T UOWTWIOD (volfiond D1IADD) U00'T syle (DJD]J21S DIADD) UOO'T poyeoly}-poy (sndosp] sndosvT) uesiwieid MOTI (S1jp19b]8 snapwujn.7) LeWU[N] UIIYIION (sijpuady pjnsuvnjD) Yond psytej-suo TJ (syiqpjaads p1dajouos) Jopiq SUTy] (DUISSI]JOW D1AAJDUOG) Jopiy UOWIUIOD (4OJDALIS SNBAIP ) JOSUCBIDI POISCIIq-Poy (adojauad spuy) [reg UIOYION (snupiqunjos snusd)) UeMS eIpuNny, (ojoludag DjudAg) yueIg (sisuapouDdd DJUDAG) ISOOD epeuedy (1uOSUIYIINY DJUDAG) ISOODH BUI;YIVD (SUdISAaJNAADI AASUP) 9SOOD MOUS sorsadg ‘PURIST OY} WOLF SIQISTA Ud9Q DALY ISI SPIIG dy} SIOYM POPNIOUI de SPIOIOI BIS-IV ‘S]9S BLP INOJ UO pase puUrIS] JasuULY Ie ..SNILIS SUIP9EIG pUB SUOTJAIOSQO PIIg *[ ATAV$ 2019 GASTON: BIRDS OF MANSEL ISLAND 23 was conducted after most shorebirds would have com- pleted breeding, and breeding could not be confirmed in that season for any shorebird species. Those species Combined statust PR PO B (17), PR (10) ri for which breeding could be confirmed were those that have longer breeding periods. Conversely, surveys in = 2016 found the island partly covered in snow, which oS poo a ae presumably delayed breeding for many species, mak- 3 5 ing the surveys earlier than ideal. Breeding could be a confirmed for only five species, although it was con- 2 i sidered probable for another 15 species. Among spe- s 2 Ro a cies for which breeding was confirmed, Canada and 7 Cackling Geese (Branta canadensis, Branta hutchin- g soni), Northern Pintail (Anas penelope), and Dunlin o| 3 00 a] 9 (Calidris alpinus) are not shown as breeding on Man- = S N sel Island by Richards and Gaston (2018). 2 z Only seven species were reported by all four sur- Py 5 veys: Cackling Goose, Tundra Swan (Cygnus colum- E 5 bianus), King Eider (Somateria spectabilis), Long- 5 + + =| tailed Duck (Clangula hyemalis), Red-throated Loon sf (Gavia stellata), Sabine’s Gull (Xema sabini), and e Herring Gull (Larus argentatus). Black-bellied Plover Z (Pluvialis squatarola), American Golden-plover (Plu- vialis dominica), and Arctic Tern were recorded on pee ae les all three post-1930 surveys. According to the 2016 a 5 = a {4 survey, the most widespread species (Seen in nine ay or more survey squares) were Canada Goose, King B= 5 Eider, and Herring Gull. In 1992, 38 pre-flying King ch| & nN a S Eider ducklings were seen in four separate creches, 7 S i i along with nine adults and the species was the second 5 most widespread on the 2016 survey. These observa- ng | 2 tions suggest that Mansel Island may be an import- E 2 = 5 ant breeding area for this species. Likewise, Tundra B =| < Swan, as well as being seen on all surveys, was the S most widespread species reported on the aerial sur- vey in 1984. Mansel Island appears to support a sig- nificant population of this species. Overall, the avifauna of Mansel Island is very similar to that of the better-known Coats Island, im- mediately to the west (Gaston and Ouellet 1997). Like Coats, it supports Caribou but apparently not lemmings (Dicrostonyx and Lemmus spp.; Gaston et al. 2012). The absence of the latter probably deter- mines the lack of specialist lemming predators, such as Snowy Owl (Nyctea scandiaca) and Long-tailed Jaeger (Stercorarius longicaudus). The very flat top- ography, lacking cliffs, may determine the absence of Peregrine Falcon (Falco peregrinus) and Common Raven (Corvus corax) and the relative paucity of (season) Fall Sutton 1932b Decker 1984 Summer Summer 24 probable breeding, PO +Combined breeding evidence from all surveys to give likeliest status. definite evidence of breeding, PR $A geolocator-tracked Ruddy Turnstone spent the summers of 2014 and 2015 on Mansel Island and showed evidence of incubation in 2015 (R. Porter pers. comm. August 2018). tA satellite-tracked Red Knot was recorded in summer on Mansel Island by Lathrop et al. (2018). Black Guillemot (Cepphus grylle) Horned Lark (Eremophila alpestris) Lapland Longspur (Calcarius lapponicus) Snow Bunting (Plectrophenax nivalis) S 3 ~ NX Ss & oD ~~ i S Snow Bunting (Plectrophenax nivalis), all common X : : : < 3 z mn on adjacent parts of mainland Quebec (Gaston ef al. R ee 2 1985). However, the breeding of Sabine’s Gull and the S) 2c 3 probable breeding of Red Knot (Calidris canutus) on =I ee ie = Mansel Island represent the most southeasterly ex- ; 8 z E Si! tension of these species’ known ranges in Canada ell w Ao el & (Richards and Gaston 2018). 24 THE CANADIAN FIELD-NATURALIST Acknowledgements Iam very grateful to Yves Aubry for providing me with information on the 2016 surveys, to R. Porter of the Delaware Bay Shorebird Project for information on satellite-tracked birds, and to my companions in the 1992 visit, Vicky Johnston, Ilya Storm, and the crew of the MV. Teregluk. Literature Cited Bart, J.R., and V.H. Johnston. 2012. Arctic Shorebirds in North America: a Decade of Monitoring. Studies in Avian Biology 44. University of California Press, Ber- keley, California, USA. Environment Canada, Lands Directorate, Conservation and Protection. 1970. Mansel Island, District of Kee- watin, Northwest Territories (map). Land use infor- mation series. Surveys and Mapping Branch, Energy, Mines and Resources Canada, Ottawa, Ontario, Can- ada. Accessed 22 May 2019. http://sis.agr.gc.ca/cansis/ publications/maps/nluis/250k/lu/nluis_250k_lu_35el_ 45hijpg. Gaston, A.J., D.K. Cairns, R.D. Elliot, and D.G. Noble. 1985. A natural history of Digges Sound. Report 46. Canadian Wildlife Service, Ottawa, Ontario, Canada. Gaston, A.J., R. Decker, F.G. Cooch, and A. Reed. 1986. The distribution of larger species of birds breeding on Vol. 133 the coasts of Foxe Basin and northern Hudson Bay. Arctic 39: 285-296. https://doi.org/10.14430/arctic2089 Gaston, A.J., M. Gavrilo, and C. Eberl. 2012. Ice bridg- ing as a dispersal mechanism for Arctic terrestrial ver- tebrates and the possible consequences of reduced sea ice cover. Biodiversity 13: 182-190. https://doi.org/10. 1080/14888386.2012.719177 Gaston, A.J., and H. Ouellet. 1997. Birds and mammals of Coats Island, NWT. Arctic 50: 101-118. https://doi. org/10.14430/arcticl1094 Lathrop, R.G., L. Niles, P.A. Smith, M. Peck, A. Dey, R. Sacatelli, and J. Bognar. 2018. Mapping and mod- eling the breeding habitat of the Western Atlantic Red Knot (Calidris canutus rufa) at local and regional scales. Condor 120: 650-665. https://doi.org/10.1650/ condor-17-247.1 Richards, J., and A.J. Gaston. 2018. The Birds of Nuna- vut. University of British Columbia Press, Vancouver, British Columbia, Canada. Sutton, G.M. 1932a. Birds of Southampton Island. Carne- gie Institute, Washington, DC, USA. Sutton, G.M. 1932b. Notes on a collection of birds from Mansel Island, Hudson Bay. Condor 34: 41—43. https:// dot.org/10.2307/1363790 Received 5 November 2018 Accepted 12 February 2019 The Canadian Field-Naturalist Note Behaviour of a porcupine (Erethizon dorsatum) swimming across a small boreal stream THOMAS S. JUNG Department of Environment, Government of Yukon, Whitehorse, Yukon Y1A 2C6 Canada and Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H1 Canada; email: thomas jung@ gov yk.ca; tyung@ualberta.ca Jung, T.S. 2019. Behaviour of a porcupine (Erethizon dorsatum) swimming across a small boreal stream. Canadian Field- Naturalist 133(1): 25-27. https://doi.org/10.22621/cfn.v133i1.2107 Abstract The swimming behaviour of North American Porcupine (Erethizon dorsatum) is largely unrecorded, even though much of its habitat is bisected by innumerable rivers and streams. Moreover, the literature is inconsistent regarding how readily por- cupines take to the water and how well adapted they are for swimming. I observed a porcupine swimming across a relatively placid and shallow braid in the Klondike River (Yukon, Canada), after it had aborted three apparent attempts to swim at a relatively fast-flowing, deep channel upstream. This observation provides evidence of porcupine swimming across moving water and suggests that they may be reluctant to do so and selective of where they cross rivers and streams. Key words: Behaviour; Erethizon dorsatum; North American Porcupine; swimming Observations of North American Porcupine (Ere- thizon dorsatum) swimming are rare in the literature, suggesting that it may be uncommon behaviour. Yet, much of their range is within the boreal forest (Woods 1973; Roze and Ilse 2003), which is interspersed and divided by numerous water bodies. The few obser- vations reported involve swimming in ponds and lakes (Dean 1950; Woods 1973; Roze 2009), with no observations of them crossing rivers or streams. An unusual observation of a Bull Trout (Sa/velinus con- fluentus) embedded with porcupine quills provided circumstantial evidence of a porcupine swimming in moving water (Cott and Mochnacz 2007). The willingness of porcupines to swim is unclear, particularly across rivers and streams. Some author- ities suggest that porcupines are not averse to swim- ming (Roze and Ilse 2003; Roze 2009), and that swimming is an important means for them to ac- cess seasonal food resources. For instance, there are observations of porcupines feeding on water lilies (Nymphaeaceae) in shallow ponds and swimming to retrieve food items that they then bring to shore to consume (Dean 1950; Roze and Isle 2003). Moreover, their quills may also be adapted, in part, to help them swim; specifically, Roze and Ilse (2003: 376) surmised that “their watertight, sponge-filled inte- riors aid in floatation, enhancing the porcupine’s ©The Ottawa Field-Naturalists’ Club swimming capabilities”. Alternatively, Woods (1973: 4) opined that “they do not like to swim”, although he conceded that they have been observed crossing small water bodies. In an early “experiment”, Murie (1926: 112) noted: One day I tried to make a porcupine swim across a narrow stream. I shoved it toward the water with a stick and intercepted it whichever way it turned. Nothing could induce it to swim, al- though I almost shoved it bodily into the water. It came straight toward me, rather than cross the stream, and I finally gave up the attempt. Here, I provide an observation of a porcupine swimming across a small boreal stream and note its apparent indecision in doing so. While angling ona braid of the Klondike River, ~15 km east of Dawson City, Yukon, Canada (64.059°N, 139.433°W), I observed a porcupine approaching and, eventually, swimming across the river. At ap- proximately 1705 Pacific Daylight Time, on 6 July 2018, an apparently full-grown porcupine emerged from tall shrubs on the far side of the stream. I did not know its age or sex. The porcupine came to the shore (point A in Figure 1) and, after about 15 s of apparently sniffing toward the far shore, it stepped about 30 cm into the stream, immersing its front 26 THE CANADIAN FIELD-NATURALIST Vol. 133 , names Pe eee Se —. ee) asc ee era ae get a a we ~ > = . — - = : _ > : - a al _ = a SS Sa = — a ‘ — = a = = Figure 1. Photograph of the site where a North American Porcupine (Erethizon dorsatum) swam across a braid in the Klondike River, Yukon, Canada. At sites A, B, and C, the porcupine stepped into the stream but did not cross it; the dashed line (D) indicates where it swam across the stream. Photo: T.S. Jung. legs. However, rather than swim across the stream, it backed out of the water and sniffed across the stream again. It immediately moved about 6 m downstream and repeated the same actions at point B (Figure 1). The porcupine moved another 15 m downstream along the shore to point C (Figure 1) and again en- tered the stream, this time without apparently sniff- ing the far shore, and it waded deeper until its belly and both legs were under water; however, it again re- turned to the shore within approximately 30 s. The porcupine then moved into the shrubs and was not seen for about 5 minutes. I then observed it ~35 m downstream of point C, at point D (Figure 1), where it entered the water and swam across the stream, after standing in the stream with both legs and its belly under the water for about 1 minute. The porcu- pine reached the far shore after swimming for about 2 minutes, and then entered the forest on the other bank and was no longer observed. I do not know why the porcupine crossed the stream. It was on a small island in the Klondike River that was largely covered with willow (Salix spp.) and alder (A/nus spp.), whereas, the other side of the stream was covered by mature boreal forest, domin- ated by Balsam Poplar (Populus balsamifera L.) and White Spruce (Picea glauca [Moench] Voss) trees. It may have been attracted to something not available on the island at that time. Points A—C, where the porcupine entered the stream but did not cross it, were in the section of the stream with the swiftest water and a relatively deep channel (~1.2 m deep). In contrast, point D (Figure 1), where the porcupine entered and crossed the stream, was immediately downstream of the riffle, and the water there was more placid and only about 0.5 m deep. However, the stream here was about 30 m wide, compared to about 8 m wide at the riffle (points A—C). It appeared that the porcupine was hesitant to enter the stream and cross the riffle and selected a location to cross where the stream was comparatively slow flowing. This observation suggests that porcupines may not be strong swimmers and seek areas with slow-moving water to cross rivers and streams. This observation is of scientific value from two perspectives. First, to the best of my knowledge, this is the first record of a porcupine crossing a stream or river, despite the fact that this must be relatively com- mon behaviour for porcupines given the innumerable streams and rivers in the boreal forest, even if it is not regularly observed by humans. Second, given the apparent indecision of the animal about whether to cross the stream, this observation suggests that some porcupines may be averse to swimming, supporting the assertion of Woods (1973). In addition, this obser- vation suggests that porcupines may be selective in terms of where they cross rivers and streams, avoid- ing deep, turbulent water in favour of more placid and shallow sections. Although the porcupine swam across the stream with apparent ease, its head and body were quite low in the water; thus, waves and rif- fles may pose a substantial risk of drowning. A swift current could also quickly take a porcupine down- stream during a crossing into hazards, such as rough water or waterfalls. 2019 Acknowledgements Dwayne Lepitzki, Garth Mowat, and an anonym- ous reviewer kindly provided comments that im- proved this manuscript. Literature Cited Cott, P.A., and N.J. Mochnacz. 2007. Bull trout, Sa/velinus confluentus, and North American porcupine, Erethizon dorsatum, interaction in the Mackenzie Mountains, Northwest Territories. Canadian Field-Naturalist 121: 438-439. https://doi.org/10.22621/cfn.v12114.523 Dean, H.J. 1950. Porcupine swims for food. Journal of Mammalogy 31: 94. Murie, O.J. 1926. The porcupine in northern Alaska. Jour- JUNG: SWIMMING PORCUPINE 27 nal of Mammalogy 7: 109-113. Roze, U. 2009. The North American Porcupine. Second Edi- tion. Cornell University Press, Ithaca, New York, USA. Roze, U., and L.M. Ilse. 2003. Porcupine, Erethizon dors- atum. Pages 371-380 in Wild Mammals of North Amer- ica: Biology, Management, and Conservation. Second Edition. Edited by G.A. Feldhamer, B.C. Thompson, and J.A. Chapman. The John Hopkins University Press, Baltimore, Maryland, USA. Woods, C.A. 1973. Erethizon dorsatum. Mammalian Spe- cies 29: 1-6, Received 16 July 2018 Accepted 7 February 2019 The Canadian Field-Naturalist More Mountain Chickadees (Poecile gambelt) sing atypical songs in urban than in rural areas STEFANIE E. LAZERTE!”", KRISTEN L.D. MARINI, HANS SLABBEKOORN‘*, MATTHEW W. REUDINK?, and KEN A. OTTER! 'Natural Resources and Environmental Studies, University of Northern British Columbia, 3333 University Way, Prince George, British Columbia V2N 4Z9 Canada ?Current address: Department of Biology, Brandon University, 270- 18th Street, Brandon, Manitoba R7A 6A9 Canada >Department of Biological Sciences, Thompson Rivers University, 805 TRU Way, Kamloops, British Columbia V2C 0C8 Canada “Institute of Biology, Leiden University, Sylviusweg 72, 2333 BE Leiden, the Netherlands “Corresponding author: sel@steffilazerte.ca LaZerte, S.E., K.L.D. Marini, H. Slabbekoorn, M.W. Reudink, and K.A. Otter. 2019. More Mountain Chickadees (Poecile gambeli) sing atypical songs in urban than in rural areas. Canadian Field-Naturalist 133(1): 28-33. http://doi.org/ 10.22621/cfn.v13311.1994 Abstract Urbanization results in novel ecosystems with unique challenges. These may lead to problems during song learning or development and could result in the singing of atypical songs. During studies of Mountain Chickadees (Poecile gambeli) and urbanization in British Columbia, Canada, we observed males singing atypical songs along an urbanization gradient. We found that eight of 78 males consistently sang atypical songs and the odds of singing atypical songs increased with urbanization. We explored several explanations including habitat quality, population density, and bioacoustics. Future stud- ies investigating causes and consequences of atypical singing will clarify effects of urbanization on Mountain Chickadees. Key words: Mountain Chickadee; Poecile gambeli, Paridae; communication; atypical songs; urbanization; urbanization index Introduction Among songbirds, unusual songs are those that differ from species-specific local song types. These unusual songs may be (a) rarely heard ‘special’ songs (such as whisper songs), (b) juvenile songs, the result of early song development, (c) uncommon mimicry of other species, or (d) dialectal songs in an abnormal geographic location (Borror 1968). Unusual songs that do not fit these four categories are considered atypical (e) and may be the consequence of errors in learning or developmental problems (Borror 1968). Occasionally, young males make ‘mistakes’ when learning their songs. Perhaps they have few tutors, or cannot hear their tutors well, or perhaps their tu- tors are a closely related species (e.g., Black-capped Chickadees [Poecile atricapillus| and Carolina Chickadees [P. carolinensis] learn each other’s songs; Sattler et al. 2007). There may also be developmen- tal problems, as poor-quality habitat can lead to poor- quality songs (e.g., poor-quality Black-capped Chick- adee songs appear less dominant to both males and females; Grava et al. 2012, 2013a), which, in extreme cases, could be considered atypical. Alternatively, ©The Ottawa Field-Naturalists’ Club changes to habitat acoustics may result in young males incorrectly hearing their tutors’ songs or act- ively modifying their own song to reduce interfer- ence and increase transmission (e.g., Slabbekoorn and den Boer-Visser 2006). Many situations leading to atypical songs may occur as a result of urbanization. Urbanization cre- ates a novel ecosystem with unique challenges for many species. Among birds, urbanization can lead to changes in habitat quality that may be positive (e.g., increased food availability from bird feeders; Robb et al. 2008) or negative (e.g., habitat loss, competi- tion with invasive species, or environmental pollut- ants; McKinney 2002), and may influence population dynamics. Urbanization can also lead to altered habi- tat acoustics (e.g., echoes and reverberation from buildings and pavement; Warren et a/. 2006) and anthropogenic noise pollution, which can interfere with vocal communication through masking of lower frequencies (Patricelli and Blickley 2006; Shannon et al. 2015). Mountain Chickadees (Poecile gambeli) live in montane forests in western North America. They are 2019 found in urban areas, although they occur at lower densities than they do in rural areas (LaZerte 2015; S.E.L. and K.L.D.M. pers. obs.) and may thus be less urban-adapted than Black-capped Chickadees. Here we present a short exploration of the relationship between atypical songs and urbanization in Mountain Chickadees using the combined data from Marini (2016) and LaZerte (2015). Methods We analyzed recordings of 78 adult male Moun- tain Chickadees vocalizing at dawn in the spring dur- ing nest-building and egg-laying (2012 through 2015). These recordings were obtained from two studies in- vestigating effects of urbanization: communication and individual condition (Marini et al. 2017a, n = 42), and vocal plasticity (LaZerte et al. 2017, n = 36). Recordings were made in and around the cit- ies of Williams Lake (n = 12; 52.129°N, 122.138°W), Kamloops (n = 60; 50.676°N, 120.341°W), and Ke- lowna (n = 6; 49.884°N, 119.493°W), British Colum- bia (BC), Canada. Each male was recorded a max- imum of once per year. We used site territoriality to distinguish among males within a year, but sites in Kamloops were revisited between years. Known duplicate recordings of males (if the male was banded or identified by distinctive atypical singing) were omitted. Habitat urbanization was evaluated as a con- tinuous index (low = rural, high = urban) by compar- ing satellite Google Earth images (Google Inc. 2012) of territories (defined as a circular area 150 m in diam- eter around the recording location of the focal male) and scoring the amount of natural vegetation (natural grass or trees) versus urban ground cover (pavement, buildings, or lawn; for more details see LaZerte et al. 2017; for scripts and tutorial see https://github.com/ steffilazerte/urbanization-index). The lowest habitat urbanization value (—0.95) reflected sites with 100% natural vegetation (no pavement, no buildings, no lawns). The highest value (2.01) reflected sites with only 11% natural vegetation cover, and 89% pave- ment, buildings, or lawn. We only included samples with a minimum of five minutes of vocalization and 25 songs (as Mountain Chickadees use both songs and calls during the dawn chorus; McCallum et al. 1999; Grava et al. 2013b). Part of LaZerte et al.’s (2017) experimental protocol involved exposing males to five minutes of experi- mental noise. Although they found no effect of this exposure on song variation, we excluded all songs recorded during the noise exposure period and in the five minutes following. Mountain Chickadees in BC typically sing songs with 3—5 notes in descending order (Grava eral. 2013b; Figure la). We therefore defined songs as atypical if LAZERTE ET AL... ATYPICAL URBAN MOUNTAIN CHICKADEE SONGS 29 they were monotone (multiple notes sung on a sin- gle frequency; Figure 1b top), contained a reverse frequency change (ascending note[s] as opposed to descending; Figure 1b middle), or contained novel notes (e.g., anote with an extreme upwards frequency sweep; Figure 1b bottom). We used categorical desig- nations for songs as opposed to measuring song char- acteristics because our data were obtained from two prior studies. In one study, songs had been categor- ized, but there were no compiled data on individual songs. Although atypical songs are unusual, it is not uncommon for an individual to occasionally sing a few atypical songs. Therefore, we classified males as atypical singers only if they consistently sang atyp- ical songs (>80% of all songs recorded were atypical, most males sang <5% atypical songs). To determine whether the odds of being an atyp- ical singer increased with urbanization, we performed a logistic regression of male singer type (atypical/typ- ical) against the urbanization index using R statis- tical software (version 3.3.2; R Core Team 2016). We calculated bias-corrected and adjusted (BCa) boot- strap 95% CI for coefficients. We performed 10000 replicates using the boot package for R (version 1.3- 20; Angelo and Ripley 2017). Figures were created using the R package ggplot2 (version 2.2.1; Wickham 2009). Spectrograms were created with Hanning window lengths of 1024 using the R packages ggplot2 and seewave (version 2.0.5; Sueur et al. 2008). Results Eight of 78 individuals consistently sang atyp- ical songs. Roughly categorizing urban areas as those with an urbanization index greater than the mean (0) showed that 21% of urban males consistently sang atypical songs whereas only 2% of rural males did (Figure 2a). The odds of a male consistently singing atyp- ical songs increased significantly with the continu- ous urbanization index (Log odds = 1.10, 95% CI = 0.28—2.30, SE = 0.42, z=2.61, P = 0.009; Figure 2b); expressed as an odds ratio, for every 1 unit increase in the urbanization index, males were 3.00 (95% CI = 1.32—9.95) times as likely to be atypical singers. The probability of individuals in the most rural habi- tats being atypical singers was 2.4% (95% CI = 0.3— 11.2%). In the most urban habitats, the probability was 39.0% (95% CI = 11.6—68.0%). Discussion Consistently singing atypical songs was not com- mon; however, the odds of doing so increased with in- creasing urbanization. Because these recordings were collected during the breeding period before juveniles were present, it is highly unlikely that atypical songs 30 Kelowna Williams Lake ol 1 1 | Rese; » ——_) —_ Frequency (kHz) b w i THE CANADIAN FIELD-NATURALIST Vol. 133 b. Atypical Monotone cee fm amt: pass? Reverse frequency drop Ga a Kamloops Novel note: Freq. sweep & reverse freq. drop / & 44 SS t atl | =a exes) T T T T T T T T 0.0 0.5 1.0 1.5 2.0 0.0 0.5 1.0 1.5 2.0 Time (s) FiGurE 1. Variation in Mountain Chickadee (Poecile gambeli) songs in British Columbia, Canada. a. Typical regional variation; all songs show descending frequencies. b. Some examples of atypical songs include monotone songs (top), songs with a reverse frequency drop (middle), and songs with novel notes (bottom). a. 2% atypical 21% atypical Number of individuals Rural Urban Habitat type b. Atypical 4 @) Oo @® O00 0 o 5 a 2 5 Typical 4 een GD O@ anes ce T T T T -1 0 1 2 Urbanization Index Singer type [] Typical || Atypical FiGuRE 2. Male Mountain Chickadees (Poecile gambeli) are more likely to consistently sing atypical songs in urban areas. a. By categorizing urban sites as those with an urbanization index > 0 and rural sites as those with an urbanization index <0, urban sites show 21% of males singing atypical songs versus 2% in rural areas. b. As urbanization increases, the like- lihood of being an atypical singer increases. The line represents the predicted logistic regression, the grey area shows the 95% CI interval around the predicted model. Each point represents a male Mountain Chickadee. The outlier (top left panel b) was recorded in a rural area on the outskirts of Kamloops. There were no sources of water, nor any other obvious sources of noise. It is possibly it could have been a windy location as it was on the side of a hill, but excessive wind was not noted. It was up slope of the train tracks, ~1.5 km away, a distance unlikely to have had an effect. Possibly this individual migrated to the area from an urban area. represent early song development. Further, as these cities are relatively small (the largest, Kamloops, has a population of 90280; Statistics Canada 2017) and are surrounded by rural habitat, it seems unlikely that birds from different populations (and with dif- ferent song types) would have exclusively settled in urban areas, or that these urban habitats are iso- lated enough to facilitate cultural evolution of song (cf. Gammon and Baker 2004; Luther and Derryberry 2012). Consequently, atypical singers in urban areas may result from differences in habitat quality, popula- tion density, or environmental acoustics. Poor-quality habitat may be associated with poor- quality males, either because males in urban habitats do not get enough resources or because only poor- quality males will settle in urban habitats. This, in 2019 turn, may lead to poor-quality song (e.g., nutritional stress hypothesis; Nowicki et a/. 2002; male quality; Grava et al. 2012) which could explain the increase in atypical singers. However, our previous studies of Mountain Chickadees in Kamloops suggest that urban habitat seems to be of at least equivalent qual- ity to rural habitat (Marini et a/. 2017b). Thus, poor- quality habitat may not fully explain the presence of atypical singers we found. Mountain Chickadees are less abundant in ur- ban than in rural areas (LaZerte 2015; S.E.L. and K.L.D.M. pers. obs.). In some species, greater urban population densities affect song variation, by influen- cing male-male interactions (e.g., Eurasian Blackbird [Turdus merula], Ripmeester et al. 2010; Great Tits [Parus major], Hamao et al. 2011). However, it is unclear how reduced competition could lead to singing atypical songs in Mountain Chickadees. Alternatively, low population density may result in fewer tutors or tutors that are farther away, making it difficult for young chickadees to learn songs correctly (similar to Laiolo and Tella 2005). Further, low dens- ities may also result in the direct introduction of un- usual song types by juveniles and less social pressure to conform to local song types (Gammon ef al. 2005; Gammon 2007). Urban areas are often noisy (LaZerte et al. 2015) and more pavement and concrete leads to altered acoustics (Warren eft al. 2006). These changes may interfere with vocal communication leading to ad- justed songs and/or calls. Male Mountain Chickadees are known to adjust their vocalizations in noisy habi- tats and in response to noise exposure (LaZerte et al. 2017). In a study on closely related Great Tits, Slabbekoorn and den Boer-Visser (2006) found that, throughout Europe, urban males sang more atyp- ical song types (songs with fewer or more notes than the typical 2—4) than rural males, and suggested this could be due to noise interference. If, during song learning, only un-masked and well transmitted as- pects of tutor songs are learned properly, changes in bioacoustics could result in atypical songs (Rabin and Greene 2002; Slabbekoorn and den Boer-Visser 2006). Depending on the situation, these atypical songs could be beneficial or detrimental. Atypical songs, which are the result of learning only the least- masked aspects of a normal song (e.g., Mountain Chickadee monotone songs may represent songs which have lost low-frequency notes), could result in less noise-interference and better transmission, and could thus be an adaptation to urban environments. Alternatively, atypical songs may be a symptom of poor learning in urban areas wherein young males settling in urban areas are learning songs incorrectly from tutors that results in poor quality songs. LAZERTE ET AL... ATYPICAL URBAN MOUNTAIN CHICKADEE SONGS 3] While atypical songs were uncommon overall, urban Mountain Chickadees in BC were more likely to consistently sing atypical songs than rural males. However, it is not clear whether these songs repre- sent a response to the urban acoustic environment, or a symptom of low population densities. Studies in progress suggest that atypical songs may transmit bet- ter in noisier conditions than typical songs (S.E.L. un- publ. data). However, Gammon et al. (2005) observed more atypical songs in Black-capped Chickadees in quiet, rural populations as opposed to presumably noisier, urban populations, suggesting a stronger role for population density than urban noise. There are fewer studies on Mountain Chickadees and it is thus less clear how prevalent atypical songs are in more natural landscapes. Possibly, they might be more common than in Black-capped Chickadees, simply because their song varies more among populations than do Black-capped Chickadees (e.g., Grava et al. 2013a). Further studies exploring the interaction be- tween noise and population densities (such as ina 2x2 factorial design, varying density of birds and levels of urban noise) could help clarify the potential mech- anism. The research could be an observational study or a manipulative experiment (e.g., alter population density through removing birds, use audio speakers to vary the amount of urban noise). It is also unclear what consequences these changes may have on com- munication or reproductive success, which further studies may also help to clarify. Author Contributions S.E.L., K.A.O., and H.S. contributed to the de- velopment and design of the LaZerte (2015) study, and K.L.D.M., K.A.O., and M.W.R. contributed to the development and design of the Marini (2016) study. S.E.L. and K.L.D.M. collected data and conducted data cleaning and preparation. S.E.L. conducted the analysis and wrote the manuscript. All authors con- tributed to development of ideas and commented on draft versions of the manuscript. Acknowledgements The assistance of technicians from Thompson Rivers University and University of Northern British Columbia (UNBC) was greatly appreciated. We wish to thank: BC Parks; the cities of Williams Lake, Kel- owna, and Kamloops; Regional District of the Cen- tral Okanagan; Thompson Rivers University; and University of British Columbia Okanagan for al- lowing access to their parks and grounds. An anonym- ous reviewer and David Gammon provided helpful comments on the manuscript. Financial support was provided by The James L. Baillie Memorial Fund of Bird Studies Canada (S.E.L.); Natural Sciences and 32 THE CANADIAN FIELD-NATURALIST Engineering Research Council of Canada through Post Graduate doctoral scholarship (S.E.L.), Indus- trial Postgraduate Scholarship (K.L.D.M.), and Dis- covery grants (K.A.O. and MW.R.); and UNBC through Graduate Entrance Research Awards and a Research Project Award (S.E.L.). Literature Cited Angelo, C., and B. Ripley. 2017. boot: Bootstrap R (S-Plus) Functions. Version 1.3-20. Accessed 30 July 2017. http:// CRAN.R-project.org/package=boot. Borror, D.J. 1968. Unusual songs in passerine birds. The Ohio Journal of Science 68: 129-138. Gammon, D.E. 2007. How postdispersal social environ- ment may influence acoustic variation in birdsong. Pages 183-197 in Ecology and Behavior of Chickadees and Titmice: an Integrated Approach. Edited by K.A. Otter. Oxford University Press, Oxford, United Kingdom. Gammon, D.E., and M.C. Baker. 2004. Song reper- toire evolution and acoustic divergence in a popula- tion of Black-capped Chickadees, Poecile atricapillus. Animal Behaviour 68: 903-913. https://doi.org/10.1016/j. anbehav.2003.10.030 Gammon, D.E., M.C. Baker, and J.R. Tipton. 2005. Cultural divergence within novel song in the Black- capped Chickadee (Poecile atricapillus). Auk 122: 853— 871. https://doi.org/10.1642/0004-8038(2005)122[0853: cdwnsi|2.0.co;2 Google Inc. 2012. Google Earth. Accessed 26 August 2014. http://google.com/earth/. Grava, T., A. Grava, and K.A. Otter. 2012. Vocal per- formance varies with habitat quality in Black-capped Chickadees (Poecile atricapillus). Behaviour 149: 35— 50. https://doi.org/10.1163/156853912x625854 Grava, T., A. Grava, and K.A. Otter. 2013a. Habitat- induced changes in song consistency affect perception of social status in male chickadees. Behavioral Ecology and Sociobiology 67: 1699-1707. https://doi.org/10.1007/ s00265-013-1580-z Grava, A., K.A. Otter, T. Grava, S.E. LaZerte, A. Poesel, and A.C. Rush. 2013b. Character displacement in dawn chorusing behaviour of sympatric Mountain and Black- capped Chickadees. Animal Behaviour 86: 177-187. https://doi.org/10.1016/j.anbehav.2013.05.009 Hamao, S., M. Watanabe, and Y. Mori. 2011. Urban noise and male density affect songs in the Great Tit Parus ma- Jor. Ethology Ecology & Evolution 23: 111-119. https:// doi.org/10.1080/03949370.2011.554881 Laiolo, P., and J.L. Tella. 2005. Habitat fragmentation af- fects culture transmission: patterns of song matching in Dupont’s Lark. Journal of Applied Ecology 42: 1183— 1193. https://doi.org/10.1111/j.1365-2664.2005.01093.x LaZerte, S.E. 2015. Sounds of the city: the effects of urban- ization and noise on Mountain and Black-capped Chick- adee communication. Ph.D. thesis, University of Nor- thern British Columbia, Prince George, British Colum- bia, Canada. https://do1.org/10.24124/2015/bpgub1059 LaZerte, S.E., K.A. Otter, and H. Slabbekoorn. 2015. Relative effects of ambient noise and habitat openness Vol. 133 on signal transfer for chickadee vocalizations in rural and urban green-spaces. Bioacoustics 24: 233-252. https://doi.org/10.1080/09524622.2015.1060531 LaZerte, S.E., K.A. Otter, and H. Slabbekoorn. 2017. Mountain Chickadees adjust songs, calls and chorus composition with increasing ambient and experimental anthropogenic noise. Urban Ecosystems 20: 989-1000. https://doi.org/10.1007/s11252-017-0652-7 Luther, D.A., and E.P. Derryberry. 2012. Birdsongs keep pace with city life: changes in song over time in an urban songbird affects communication. Animal Be- haviour 83: 1059-1066. https://do1.org/10.1016/j.anbehav. 2012.01.034 Marini, K.L. 2016. City life and chickadees: effects of ur- banization on vocal output and reproductive success of the Mountain Chickadee (Poecile gambeli). M.Sc. thesis, Thompson Rivers University, Kamloops, British Columbia, Canada. Marini, K.L., K.A. Otter, S.E. LaZerte, and M.W. Reu- dink. 2017b. Urban environments are associated with earlier clutches and faster nestling feather growth com- pared to natural habitats. Urban Ecosystems 20: 1291— 1300. https://doi.org/10.1007/s11252-017-0681-2 Marini, K.L., M.W. Reudink, S.E. LaZerte, and K.A. Otter. 2017a. Urban Mountain Chickadees (Poecile gambeli) begin vocalizing earlier, and have greater dawn chorus output than rural males. Behaviour 154: 1197-1214. https://do1.org/10.1163/1568539x-00003464 McCallum, D.A., R. Grundel, and D.L. Dahlsten. 1999. Mountain Chickadee (Poecile gambeli). In The Birds of North America Online. Edited by A. Poole. Cornell Laboratory of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.453 McKinney, M.L. 2002. Urbanization, biodiversity, and conservation. BioScience 52: 883-890. https://doi.org/ 10.1641 /0006-3568(2002)052[0883:ubac]2.0.co;2 Nowicki, S., W.A. Searcy, and S. Peters. 2002. Brain de- velopment, song learning and mate choice in birds: a review and experimental test of the “nutritional stress hypothesis.” Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 188: 1003-1014. https://doi.org/10.1007/s00 359-002-0361-3 Patricelli, G.L., and J.L. Blickley. 2006. Avian com- munication in urban noise: causes and consequences of vocal adjustment. Auk 123: 639-649. https://doi.org/ 10.1642/0004-8038(2006)123[639:aciunc]2.0.co;2 R Core Team. 2016. R: a Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. Rabin, L.A., and C.M. Greene. 2002. Changes to acous- tic communication systems in human-altered environ- ments. Journal of Comparative Psychology 116: 137— 141. https://doi.org/10.1037/0735-7036.116.2.137 Ripmeester, E.A.P., J.S. Kok, J.C. van Rijssel, and H. Slabbekoorn. 2010. Habitat-related birdsong diver- gence: a multi-level study on the influence of territory density and ambient noise in European Blackbirds. Behavioral Ecology and Sociobiology 64: 409-418. https://doi.org/10.1007/s00265-009-0857-8 Robb, G.N., R.A. McDonald, D.E. Chamberlain, and S. 2019 Bearhop. 2008. Food for thought: supplementary feed- ing as a driver of ecological change in avian popula- tions. Frontiers in Ecology and the Environment 6: 476— 484. https://doi.org/10.1890/060152 Sattler, G.D., P. Sawaya, and M.J. Braun. 2007. An assess- ment of song admixture as an indicator of hybridization in Black-capped Chickadees (Poecile atricapillus) and Carolina Chickadees (P. carolinensis). Auk 124: 926— 944. https://doi.org/10.1642/0004-8038(2007)124[926:aa osaa]2.0.co;2 Shannon, G., M.F. McKenna, L.M. Angeloni, K.R. Crooks, K.M. Fristrup, E. Brown, K.A. Warner, M.D. Nelson, C. White, J. Briggs, S. McFarland, and G. Wittemyer. 2015. A synthesis of two decades of research documenting the effects of noise on wildlife. Biological Reviews 91: 982-1005. https://do1.org/ 10.1111/brv.12207 Slabbekoorn, H., and A. den Boer-Visser. 2006. Cities change the songs of birds. Current Biology 16: 2326— 2331. https://doi.org/10.1016/j.cub.2006.10.008 LAZERTE ET AL... ATYPICAL URBAN MOUNTAIN CHICKADEE SONGS 33 Statistics Canada. 2017. Kamloops, British Columbia and British Columbia (table). Census Profile. 2016 Census. Statistics Canada Catalogue 98-316-X2016001. Ottawa. Released 29 November 2017. Accessed 22 January 2019. https://www1l2.statcan.gc.ca/census-recensement/2016/ dp-pd/prof/index.cfm?Lang=E. Sueur, J., T. Aubin, and C. Simonis. 2008. Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics 18: 213-226. https://doi.org/10.1080/0952 4622.2008.9753600 Warren, P.S., M. Katti, M. Ermann, and A. Brazel. 2006. Urban bioacoustics: it’s not just noise. Animal Behaviour 71: 491-502. https://doi.org/10.1016/j.anbehav. 2005.07.014 Wickham, H. 2009. ggplot2: elegant graphics for data an- alysis. Springer, New York, New York, USA. Received 27 October 2017 Accepted 3 January 2018 The Canadian Field-Naturalist Body mass as an estimate of female body condition in a hibernating small mammal CAITLIN P. WELLS", JAMES A. WILSON’, DoUGLAS A. KELT', and Dirk H. VAN VUREN! 'Department of Wildlife, Fish, and Conservation Biology, University of California, 1 Shields Avenue, Davis, California 95616 USA ?Department of Biology, University of Nebraska, 6001 Dodge Street, Omaha, Nebraska 68182 USA “Corresponding author: cpwells@ucdavis.edu Wells, C.P., J.A. Wilson, D.A. Kelt, and D.H. Van Vuren. 2019. Body mass as an estimate of female body condition in a hibernating small mammal. Canadian Field-Naturalist 133(1): 34—42. https://doi.org/10.22621/cfn.v13311.2073 Abstract In hibernating squirrels, the amount of energy stored as fat may influence several important demographic traits, but 1s dif- ficult to quantify in living animals. Thus, several non-destructive indices of body condition are used, including simple indi- ces that use body mass and scaled indices that correct body mass for structural size. However, the accuracy of these indices for hibernating squirrels is poorly known. We used measurements of total body electrical conductivity (TOBEC) from adult female Golden-mantled Ground Squirrels (Callospermophilus lateralis) to characterize body composition (lean mass versus fat mass) and condition (fat stores) at multiple stages in the circannual cycle. Body mass explained a high proportion of the variation in fat mass during the emergence and pre-hibernation stages, but less during the reproduction stage. Contrary to expectation, correcting for structural size did not markedly improve the condition index. Our results suggest that body mass is a good estimate of body condition during the periods of emergence and pre-hibernation fattening, and therefore may be useful to predict important components of fitness such as reproductive success and overwinter survival. Key words: Body mass; body condition; condition index; mass-length residuals; fat; ground squirrel; Callospermophilus lateralis Introduction Seasonal variation in energy supply is a central problem for many mammals, which may respond to periods of environmental energy shortage by storing energy, reducing energy expenditure, or both (Hum- phries et a/. 2003). Hibernation, which reduces meta- bolic demands during winter, is one life-history adaptation to seasonal energy scarcity, but sufficient energy stores are essential to its success (Pulawa and Florant 2000). In hibernating squirrels, the amount of energy stored as fat may influence several important demo- graphic traits such as overwinter survival (Murie and Boag 1984; Lenihan and Van Vuren 1996), timing of reproductive maturity (Barnes 1984), male breeding effort (Delehanty and Boonstra 2011), female repro- ductive success (Dobson and Michener 1995; Rieger 1996), offspring sex allocation (Allainé et al. 2000), and natal dispersal (Nunes and Holekamp 1996; Neuhaus 2006). Additionally, estimating fat stores is essential for bioenergetic models of hibernation, which can be used to project distribution changes of hibernating species under changing climatic condi- tions (Humphries et a/. 2002). However, quantifying ©The Ottawa Field-Naturalists’ Club body condition (defined here as fat stores, in grams; Kiell and Millar 1980; Dark et al. 1989) is difficult to do non-destructively. Because determining the effects of body condition on future life-history out- comes requires that the animal survive measure- ment, several non-destructive indices for estimating condition have been developed (Schulte-Hostedde er al. 2005; Peig and Green 2010). These include sim- ple condition indices that use body mass (e.g., Hock 1960), and scaled condition indices that attempt to correct body mass for structural size (e.g., Reid 1988). Many studies use total body mass as a simple condi- tion index, with the implicit assumption that greater mass reflects greater relative fat stores (Barnes 1984; Sauer and Slade 1987; Lenihan and Van Vuren 1996; Neuhaus 2003; Lane et a/. 2011). However, larger ani- mals may have greater mass due to larger structural size (skeleton and associated lean tissue) instead of greater fat stores (Dobson 1992). Thus, some stud- ies use a scaled condition index based on residuals derived from a regression of body mass on structural size, with the expectation that correcting body mass by the structural size of an individual improves the estimate of its condition (Bachman 1993; Dobson 2019 and Michener 1995; Dobson et al. 1999; Allainé et al. 2000). Positive residuals suggest the animal con- tains more tissue (presumably fat) than predicted for a given structural size, while negative residuals sug- gest the animal contains less tissue than predicted for a given structural size. Scaled indices are appealing because they cor- rect for variance in body mass that is unrelated to energy stores, but available evidence indicates that size-corrected measures do not necessarily improve estimates of body condition compared to use of body mass alone (Krebs and Singleton 1993; Green 2001; Schamber ef al. 2009). However, most evaluations of condition indices have focussed on mammals that do not store fat for hibernation or energy reserves, and the poor relationship between the scaled condition index and measured fat content may occur because residuals of these relatively lean species primarily reflect differences in protein or water content rather than fat (Schulte-Hostedde et al. 2001). Scaled indi- ces might be more appropriate for species in which fat content is a greater proportion of body mass, such as hibernators (Schulte-Hostedde et a/. 2001), but the predictive ability of simple versus scaled condition indices for hibernating squirrels is poorly known. Fat storage in hibernating squirrels follows circ- annual cycles of accumulation and depletion (Buck and Barnes 1999), reflecting seasonal changes in the balance between energy acquisition and expenditure (Kenagy ef al. 1989). For an index to be an appropri- ate estimate of body condition, it should explain a high proportion of the variation in fat storage, prefer- ably across multiple stages of the circannual cycle. Adult females are often excluded from condition index validation because of the confounding effect of fetal lean tissue elaboration during gestation (Krebs and Singleton 1993; Schulte-Hostedde et a/. 2005), yet energetic costs associated with hibernation and reproduction deplete fat stores, and therefore affect body condition, in adult females as well as males (Kenagy 1989; Michener and Locklear 1990; Buck and Barnes 1999). In this paper we use measurements from adult female Golden-mantled Ground Squirrels (Callospermophilus lateralis) to evaluate fat stores during four major stages (emergence, reproduction, post-reproduction, and pre-hibernation) in their cir- cannual cycle. Our goal is to assess body mass as a simple index of body condition in each stage, and determine if using a scaled index improves estimates of body condition. Methods We studied Golden-mantled Ground Squirrels over three years (2003-2005) in the northern Sierra Nevada mountains of California. These squirrels are WELLS ET AL.: GROUND SQUIRREL BODY CONDITION 35 locally abundant, medium-sized (200-300 g), and relatively well-known both ecologically and physio- logically (Bartels and Thompson 1993). Our study was conducted in the Plumas National Forest (40.004012°N, 120.810829°W) near Quincy, California, at an elevation of ~2100 m. In this area, adults emerge from hibernation in May and pups are weaned in late July; all squirrels gain weight dur- ing September before immerging into hibernation in October. Gestation in Golden-mantled Ground Squirrels is 28 days (Cameron 1967) and weaning occurs when pups reach 30 days old (Phillips 1981). We divided the active season into four circannual stages, defined broadly to encompass individual vari- ation in circannual timing: emergence, 15 May—15 June (emergence from hibernation through mating and early gestation); reproduction, 16 June—31 July (late gestation through lactation); post-reproduction, 1 August-31 August (after lactation but before late summer fattening becomes pronounced); and pre- hibernation, 1 September—early October (when pre- hibernation fattening occurs). Because we did not determine reproductive status for all females in this study, our sample may have included non- reproductive females. We captured adult female squirrels with Toma- hawk live traps (Model 201, Tomahawk Live Trap Co., Hazelhurst, Wisconsin, USA) baited with rolled oats and black oil sunflower seeds coated with pea- nut butter. Traps were set in the early morning and checked mid-morning. Our methods were conducted according to a protocol approved by the Animal Care and Use Committee of the University of Cali- fornia, Davis, and followed guidelines approved by the American Society of Mammalogists (Sikes et al. 2016). At first capture, squirrels were fitted with a uniquely numbered metal tag (Self-piercing fish tag, Style 1005-1, National Band & Tag Company, Newport, Kentucky, USA) in each ear for perma- nent identification. We attempted to capture all squir- rels monthly, but due to differential trapping success not all squirrels were captured each month. We transported captured squirrels to a laboratory near Quincy, where we anesthetized them with an intra- muscular injection of ketamine hydrochloride (100 mg/ml). We recorded body mass to the nearest 0.1 g using a portable electronic balance and body length (measured as tip of nose to anus) to the nearest 0.1 cm (Pulawa and Florant 2000). We used body length as a measure of structural size (Bachman 1993; Allainé et al. 2000); our measurements of body length showed good repeatability for individuals recaptured in the same stage (Pearson correlation r = 0.83, n = 5). We quantified body fat using an EM-SCAN SA-3000 body composition analyzer (EM-SCAN, Springfield, 36 THE CANADIAN FIELD-NATURALIST Illinois, USA; no longer available from the manufac- turer) to measure total body electrical conductivity. Total body electrical conductivity (TOBEC) is a non- destructive method to analyze the body composition of animals (Scott et a/. 2001) that has been used to obtain estimates of lean and fat mass from free-liv- ing small mammals (Walsberg 1988; Koteja 1996), including ground squirrels (Nunes and Holekamp 1996; Buck and Barnes 1999; Pulawa and Florant 2000). The TOBEC method uses electrical cur- rent, which travels differentially through fat versus lean tissue, to generate measures of electrical resist- ance; resistance measures are then converted to fat mass using species-specific calibration equations (Bachman 1994; Koteja 1996; Walsberg 1998; Scott et al. 2001). EM-SCAN readings are known to vary with ani- mal movement during measurement, differences in gut contents, changes in ambient temperature, and changes in body temperature greater than 4°C (Wals- berg 1988; Scott et al. 2001). To minimize variation due to movement, we placed immobilized squirrels on a plastic sample tray and lightly restrained them with rubber bands to maintain each squirrel in the same position (dorsoventrally, ventral side down, with the tail tucked under the body). To minimize variation due to gut contents, we only trapped squirrels early in the morning (as foraging began) and did not pro- vide food or water until after TOBEC measurement. To minimize variation due to ambient temperature, we performed measurements in a laboratory at a field station. Anesthesia often causes a drop in body tem- perature; throughout our study, however, the mean change in body temperature was —1.6 + 0.3°C (SE), and no individuals lowered their body temperatures more than 4°C. Body composition was calculated as the mean of five replicate measurements; we recorded seven replicate measurements and then discarded the highest and lowest values, though variation in meas- urements was minimal (coefficient of variation = 0.03). We determined lean mass (V,) using the cali- bration curve for Golden-mantled Ground Squirrels: M,, = 18.0 + 0.3M, + 1.2VL, x EM where M, is body mass, L, is body length, and EM is the EM Scan measurement (r? = 0.98; Pulawa and Florant 2000). We calculated fat mass by subtracting lean mass from body mass. We characterized the body composition (lean mass versus fat mass) and condition (fat mass) of adult fe- male ground squirrels during emergence, reproduc- tion, post-reproduction, and pre-hibernation stages. Because female energetic needs shift throughout the active season from expenditure on reproduction to acquisition before hibernation (Kenagy ef al. 1989), Vol. 133 potentially changing the relationship between body mass and fat mass, we considered each circannual stage separately. We assessed fat stores of 23 adult fe- male Golden-mantled Ground Squirrels; seven were measured in a single circannual stage, six were meas- ured in two circannual stages, eight were measured in three circannual stages, and two were measured in all four stages. Sample size varied by stage, and each female was included only once per stage. If females were measured more than once within the same stage, we randomly selected a single measurement from those taken in the same year (” = 5 females), and we considered measures to be independent if taken in different years (n = 2 females; Broussard et al. 2005). We also tried averaging measurements for the same female within a year, but the results were similar whether we averaged or chose measurements at random. We used analysis of variance (ANOVA) with Tukey’s HSD post-hoc tests to test for signifi- cant differences in mean body length and mean body composition among circannual stages. We used linear regression to examine the relationship between body length and mass by each circannual stage. Next, we used bivariate linear regression to evalu- ate the relationship between body mass and fat mass for each circannual stage, and also the relationship between mass-length residuals, calculated from re- gressing body mass on body length, and fat mass. In addition, because percent fat (fat mass/total body mass) is sometimes used as a measure of body con- dition in hibernating squirrels (Barnes 1984; Nunes and Holekamp 1996; Neuhaus 2003) we performed the same regressions for percent fat as we did for fat mass. The use of body mass as a variable in both the TOBEC calibration equation and as a predictor of fat mass may introduce some underlying structure to the data, with the potential to inflate the r? values. While this is unavoidable, we therefore report 7” values asso- ciated with linear regressions for comparison among stages and indexes, and without associated signifi- cance tests (Wasserstein and Lazar 2016). Finally, because our data contained substantial individual and annual variation in percent fat, which may confound relationships between condition indi- ces and percent fat inferred through linear regres- sion, we fitted linear mixed models with individual female identity and year as random effects, and circ- annual stage and condition index specified as fixed effects. Models were estimated with Bayesian infer- ence. We used a Bayesian, mixed-effects approach for two reasons: 1) the hierarchical structure of our data suggested the use of mixed effects models that produce more accurate estimates of all parameters, and 2) Bayesian approaches more accurately parti- tion variance among mixed effect parameters than 2019 likelihood-only approaches (McElreath 2016). We developed four models: two with fat mass (in grams) as the response variable, predicted by either mass or mass-length residuals, and two with percent fat as the response variable, predicted by either mass or mass- length residuals. We included all measurements (” = 61) of the 23 adult females in this analysis. We used a model comparison approach to evalu- ate the ability of each index to predict fat mass and percent fat. Specifically, we used the Watanabe- Akaike Information Criterion (WAIC) to rank mod- els, based on WAIC differences (AWAIC) and Akaike weights. Such values are analogous to other informa- tion criteria, where low AWAIC values indicate pre- ferred model, and high weight indicates increased probability that the model will successfully predict new data (Gelman ef al. 2014; McElreath 2016). All analyses were run in R version 3.5.2 (R Development Core Team 2016); we used the packages RStan (Stan Development Team 2016) and rethinking (McElreath 2016) to fit and compare mixed models, and ggplot2 (Wickham and Chang 2013) to plot figures. Results Lean mass of adult female Golden-mantled Ground Squirrels varied among circannual stages (F353; = 3.52, P = 0.02; Table 1), and was lowest at emergence from hibernation and highest before immergence. Estimated fat mass also varied among circannual stages (F3;,; = 7.35, P < 0.001), and was lowest at emergence from hibernation and highest before immergence. Percent fat varied among circ- annual stages (F3;, = 5.90, P = 0.002), and appeared stable throughout the first three stages before show- ing a sharp increase in the pre-hibernation stage. Additionally, mixed models revealed a generally positive effect of the pre-hibernation stage on fat mass, after controlling for year, individual, and mass or mass-length residual (Table 2). The relationship between body mass and body length was positive during emergence (r? = 0.55, n = 12, P < 0.01), reproduction (7? = 0.41, n = 15, P< 0.01), and post-reproduction stages (r? = 0.33, n = 16, WELLS ET AL.: GROUND SQUIRREL BODY CONDITION ov, P =0.02), but was no longer apparent during the pre- hibernation stage (77= 0.00, n= 12, P=0.98; Figure 1). Body mass explained a very high proportion of the variation (93-96%) in fat mass during the emergence, post-reproduction, and pre-hibernation stages, but a lower proportion (84%) during the reproduction stage (Figure 2). Correcting for structural size, as measured by head and body length, did not improve fit within any stage: the proportion of variation explained by mass-length residuals was less than that for the sim- ple index based on body mass during the emergence, reproduction, and post-reproduction stages (57-70%), and equivalent to that explained by body mass during the pre-hibernation stage (96%). Overall, a similar pattern was evident for the analysis based on percent fat. Body mass explained a moderate to high proportion of the variation in per- cent fat during the emergence (77 = 0.79), post-repro- duction (7? = 0.69), and pre-hibernation stages (7? = 0.91), but a lower proportion during the reproduction stage (7? = 0.56). Correcting for structural size did not markedly improve fit within most stages, though mass-length residuals did explain a significant pro- portion of the variation in percent fat (emergence r? = 0.61, post-reproduction r? = 0.46, pre-hiberna- tion r?= 0.91). Correcting body mass by body length improved model fit only in the reproduction stage (r? = 0.86). While both mass and mass-length residuals showed strong positive effects on fat mass and per- cent fat, WAIC metrics showed a clear preference for the mass models (w; =1, AWAIC=0.0; AWAIC for the second model >69 for fat grams and >20 for percent fat; Table 2). Discussion Our results suggest that body mass is a useful esti- mate of body condition during the critical periods of emergence from hibernation and pre-hibernation fat- tening, and perhaps during the post-reproductive per- iod, supporting the use of body mass as a simple index to predict important components of fitness such as female reproductive success (Rieger 1996) and over- winter survival (Murie and Boag 1984). Body mass TABLE 1. Mean length and body composition (+ 1 SE) of adult female Golden-mantled Ground squirrels (Callospermophilus lateralis) near Quincy, California, from 2003 to 2005, by circannual stage. Emergence Reproduction Post-reproduction Pre-hibernation 15 May-—15 June 16 June—31 July 1-31 August 1 September—1 October n 12 15 16 12 Mean length (cm) 17404 18+ 0.3 1740.3 18+0.3 Mean total mass (g) 158 + 8.5” IIS 3:3 1675.9" 198+ 9.4%! 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More recently 5-minute point counts have been used to better represent popu- lation indices of selected species (Ralph et al. 1995). All birds (males and females) both seen and heard are recorded at point-count stops. Male Rock Ptarmigan during the breeding period (late April to early June) are conspicuous (they can range in colour from white to mottled shades of light brown to almost black [Attu] with white bellies and wings), perch up, and make flights from conspicuous sites while calling as they advertise and defend terri- tories (Holder and Montgomerie 1993; Pelletier and Krebs 1997). Males in late May and early June can be solitary or paired with females, which are drab brown (in flight they have white wings) or mottled and in- conspicuous in many, if not most, situations (some may be on nests). Our objectives were to review the available liter- ature on surveys of Rock Ptarmigan on three islands in the Western Aleutian Islands and develop and im- plement a 5-minute point count protocol to estimate trends in breeding populations of Rock Ptarmigan on Adak, Amchitka, and Attu islands in the Western Aleutian Islands (Figure 1). Study Area We reviewed the published and other available lit- erature on Rock Ptarmigan (no other species of ptar- migan occur on these islands) on Adak (51.883°N, 176.65°W), Amchitka (51.35°-51.65°N, 178.617°— Figure 1. Aleutian Archipelago, Alaska showing Adak, Amchitka, and Attu islands. 2019 179.483°E), and Attu (52.85°N, 173.183°E) in the Western Aleutian Islands, Alaska (Appendix S1). Areas surveyed on the three islands by us and others were similar low elevation sites (i.e., marine and stream terraces) adjacent to rarely-used trails (Am- chitka and Attu) and occasionally used roads (Adak) that tended to follow coastal areas. The islands vary in size from ~300 km? (Amchitka) to 711 km? (Adak) and 894 km? (Attu). Adak is in the Andréanof group while Amchitka is in the Rat Island group and Attu is in the Near Islands. All are bounded by the North Pacific Ocean to the south and west and the Bering Sea to the north and east. The three islands are dis- tant from each other with Amchitka being 301 km southwest of Adak that is 720 km east of Attu. There are no human residents on Amchitka and Attu and the resident population on Adak is variable and <100 people. The geology of the three islands is complex with multiple inactive volcanos and volcanic flows as well as past glacial and marine erosion (Coats 1956; Fraser and Snyder 1959; Powers etal. 1960). Topography var- ies from gently sloping marine terraces to undulating tundra ranging to rugged mountains. We surveyed ptarmigan at an elevation of 10 to 300 m on all three islands. Lower and well-drained sites are occupied by grasses and sedges (Calamagrostis spp., Leymus spp., Carex spp.), and low-growing forbs including Caltha spp., Ranunculus spp., and Lupinus spp. with higher slopes dominated by crowberry (Empetrum spp.), Empetrum-Cladonia tundra, Cladina spp. lichens, and other mosses with some low-growing heather BRAUN ET AL.: ROCK PTARMIGAN IN THE WESTERN ALEUTIAN ISLANDS 51 (Cassiope spp., Phyllodoce spp.), willow (Salix spp.), and Kamchatka Rhododendron (Rhododendron cam- tschaticum Pallas) shrubs (Amundsen and Clebsch 1971; Everett 1971; White et al. 1977). The climate on all three islands is moist marine with frequent high velocity winds, rain, and fog (Gates etal. 1971). Mean daily average temperatures vary seasonally ranging from 0.4°C in January to 11°C in August. Daily (3.9°C in all seasons) and sea- sonal (9.4°C) ranges in temperature are limited (Arm- strong 1971, 1977). Wind speeds are highly variable, and the mean annual precipitation ranges from 83 to 139 cm, depending on the island, with June and July being the months with lowest precipitation (Weather- base 2015). Methods We established and conducted 5-minute point stations (protocol in Table 1) in 2015 following Ralph et al. (1995) at 0.8 km intervals along trails and roads on all three islands (dates in Table 2). All routes were conducted using an all-terrain vehicle. Starting points were at trail junctions or easily rec- ognized local features and were recorded as global positioning system coordinates (on file with Alaska Maritime National Wildlife Refuge, Homer, Alaska, USA). Point-count routes were in areas where at least four stops at 0.8 km intervals could be established. There were four routes on Adak with from six to 17 stops, three routes on Amchitka with from four to 21 stops, and three routes with from four to seven stops on Attu (Table 2). TABLE 1. Protocol for Rock Ptarmigan (Lagopus muta) surveys on Adak, Amchitka, and Attu islands, Western Aleutians, Alaska. 5-minute point counts Count and record all Rock Ptarmigan seen (as male or female) or heard at each point stop. Rarely, the vehicle stopped near a ptarmigan which did not call or flush during the 5-minute count but flushed when the vehicle departed. These birds were included in the count. Use of two counters is best (one front and one back). A central spot should be chosen. Consolidate and record totals at end of each point count before returning to the vehicle. Conduct point-count route prior to 10:00 AM. Try to avoid high winds (>30 km per hr) and heavy rain. Adak (use GPS locations at >0.8 km points) Old Loran Road (10 stops). Finger Cove (six stops). Navfac Creek to past Clam Cove (17 stops). Lake Andrew (six stops). Amchitka (use GPS locations at >0.8 km points) Jones Lake/Engineer Road (17 stops). Charlie-Baker Taxiway south (four stops). Infantry Road (21 stops). Attu (use GPS locations at >0.8 km points) Casco Cove to old airstrips (four stops). Engineer Hill (top towards Peace monument) from Massacre Creek Beach Trail (seven stops). Navytown (two stops) to Quonset Valley (four stops). 52 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Point-count (5-minute) surveys of Rock Ptarmigan (Lagopus muta, RP) on Adak, Amchitka, and Attu islands, Alaska, 2015-2017. Island Routes Date Adak 2015 Finger Bay 29 May 2015 Navfac Creek-Clam Lagoon 29 May 2015 Old Loran Station Road 30 May 2015 Andrew Lake 30 May 2015 Mean Adak 2016 Finger Bay 18 May 2016 Navfac Creek-Clam Lagoon 3 June 2016 Old Loran Station Road 27 May 2016 Andrew Lake 20 May 2016 Mean Adak 2017 Finger Bay 1 June 2017 Navfac Creek-Clam Lagoon 29 May 2017 Old Loran Station Road 30 May 2017 Andrew Lake 3 June 2017 Mean Amchitka Infantry Road 9 June 2015 Jones Lake-Engineer Road 9 June 2015 Charlie to Baker Taxiway 9 June 2015 Mean Attu Old Loran/Old Runways 3 June 2015 Massacre to Top Engineer 4 June 2015 Navytown to Quonset Valley 4 June 2015 Mean n points RP seen/heard Birds/Stop 6 6 1.0 17 12 0.7 10 49 49 6 6 1.0 1:9 3 0.5 17 16 1.4 10 10 21 6 10 17 1.4 6 0 0.0 17 16 0.9 10 16 1.6 a 4 1.3 1.0 21 12 0.6 17 3 0.2 4 0 0.0 0.4 4 0 0.0 7 0 0.0 6 0 0.0 0.0 *High winds did not allow completion or resurveys of three of six routes. Results Rock Ptarmigan were heard or seen on all but one (2017 only) point-count routes on Adak and two of three on Amchitka but none was recorded on any of the three point-count routes on Attu (Table 2). However, one ptarmigan pair was seen and four males were heard prior to establishment of point- count routes but not near any of the point-count stops on Attu. Numbers of ptarmigan per stop recorded on point-count routes were highest (1.9, 1.4, 1.0; 2015— 2017, respectively) on Adak, lower (0.4) on Amchitka, and non-existent (0.0) on Attu. Discussion A literature review of surveys and reports of Rock Ptarmigan on Adak, Amchitka, and Attu Islands re- vealed that Rock Ptarmigan were mentioned but that few surveys occurred over time with the exception of Amchitka with less information for Attu and Adak (Appendix S1). Large populations were documented for Amchitka (White et al. 1977) and Attu (Braun ef al. 2014) over short periods. Overall, the literature suggests populations of Rock Ptarmigan on the three islands were historically low, especially on Attu. Our point counts indicate the Rock Ptarmigan population was highest on Adak (2015-2017), lower in 2015 on Amchitka, and very low on Attu in 2015. Our point-count survey data on Attu in 2015 con- firmed the ongoing decline on this island reported by Braun ef al. (2014) from an intensive survey area con- ducted in 2003-2009. No effort was made to quantify ptarmigan numbers on Attu at higher elevations but ptarmigan were common at lower elevations in 2003-— 2005 (Braun et al. 2014). The areas that we surveyed on all three islands had similar relief (low marine and stream terraces), were highly disturbed in the mid 1940s and 1950s (Amchitka and Attu) to the late 1990s (Adak), but are now well vegetated with low to non-existent re- cent human occupation. The three islands have simi- lar predator assemblages (no ground mammals except rats, but with eagles, falcons, gulls, jaegers, owls, and ravens), but we have no estimate of densities. We have no basis to expect that predators (Gilg et al. 2003; Therrien et al. 2014) affected ptarmigan numbers on the three islands in 2015. We also detected no evi- dence that male aggressive behaviour was a factor at the densities we observed (Mougeot ef al. 2003). The possibility that herbivory (Sinclair et a/. 1988) could affect populations of ptarmigan across islands at substantial distances from each other through plant compounds was considered but was deemed unlikely because of isolation, few deciduous shrubs, and dis- tances involved. We documented three different levels of abun- dance of Rock Ptarmigan on Adak (high), Amchitka 2019 (lower), and Attu (very low) in 2015. The apparent, long-term decline on Attu since 2003 (Braun ef al. 2014) appears to have stabilized from 2009 to 2014 (Braun et al. 2014). We agree with the hypothesis of Sandercock et al. (2005) that animal cycles in Arctic marine and terrestrial environments are most likely affected by latitudinal gradients in the north and alti- tudinal gradients elsewhere. The islands we studied are surrounded by the North Pacific Ocean and the Bering Sea and we worked at or below 300 m, thus the birds on these high latitude islands are mostly af- fected by the marine environment. We further agree that systematic surveys (Tesar et al. 2016) to detect trends in breeding populations (Nichols and Williams 2006) of different populations of Rock Ptarmigan are needed at least at 3-5 year intervals for both theor- etical and practical reasons and should be able to de- tect population changes. It is possible that further translocations, similar to the one from Attu to Agattu in 2003-2005 (Kaler et al. 2010), may be considered to re-establish populations where they were extirpated by introduced Arctic Fox (Vulpes lagopus). Braun et al. (2014) documented the immediate recovery of ptarmigan after removal of Arctic Fox. But before such future translocations can be considered, a better survey protocol was needed to determine population status and trends of ptarmigan on these other islands. Knowing when ptarmigan populations may be ‘high’, especially if they cycle, also would be important so ad- equate numbers can be captured for immediate release on islands currently unoccupied by Rock Ptarmigan. This should reduce costs and improve chances for suc- cess of the transplants. Understanding fluctuations of Rock Ptarmigan populations, if they occur, is also important in the Arctic as the results from studies on islands may have relevance to ‘cycles’ and manage- ment of species of ptarmigan in mainland areas. Point counts may be the most efficient and least expensive method to obtain standardized data (all birds seen and/or heard) for Rock Ptarmigan in areas with road or trail systems because large areas can be surveyed with few personnel. Early counts (May) should provide an opportunity to record more fe- males than counts in early to mid June when females will be nesting. The three islands of Adak, Amchitka, and Attu each have different Rock Ptarmigan sub- species of conservation importance (Pruett ef al. 2010) and their population dynamics deserve further study. There is a continuing need for population data to provide insight into whether cycles exist and their periodicity and synchronicity among islands. Author Contributions Conceptualization: C.E.B., W.PT., and S.M.E.; Funding Acquisition: S.M.E.; Investigation: C.E.B., BRAUN ET AL.: ROCK PTARMIGAN IN THE WESTERN ALEUTIAN ISLANDS 53 W.P.T., S.M.E., and L.M.S.; Writing — Original Draft: C.E.B; Writing — Review & Editing: C.E.B., W.PT., S.M.E., and L.M.S. Acknowledgements We thank the crew of U.S. Fish and Wildlife Ser- vice research ship Tiglax for safe transit along the North Pacific Ocean and the Bering Sea and espe- cially the administrators of the Alaska Maritime National Wildlife Refuge for support of our efforts over a period of years. We acknowledge the help of the reviewers and the Editor for improvements in the manuscript. The findings and conclusions in this arti- cle are those of the author(s) and do not necessar- ily represent the views of the U.S. Fish and Wildlife Service. Literature Cited Amundsen, C.C., and E.E.C. Clebsch. 1971. Dynamics of the terrestrial ecosystem vegetation of Amchitka Island, Alaska. Bioscience 21: 619-623. https://doi.org/ 10.2307/1295734 AOU (American Ornithologists’ Union). 1957. Check- list of North American birds. Fifth Edition. American Ornithologists’ Union, Washington, DC, USA. Armstrong, R.H. 1971. Physical climatology of Amchitka Island, Alaska. Bioscience 21: 607—609. https://do1.org/ 10.2307/1295730 Armstrong, R.H. 1977. Weather and climate. Pages 53— 58 in The Environment of Amchitka Island. Edited by M.L. Merritt and R.G. Fuller. U.S. Energy Research and Development Administration, TID-26712. U.S. De- partment of Energy, Washington, DC, USA. Bart, J., M. Fuller, P. Smith, and L. Dunn. 2011. Use of large-scale, multi-species surveys to monitor Gyr- falcon and ptarmigan populations. Pages 263-271 in Gyrfalcons and Ptarmigan in a Changing World. Edited by R.T. Watson, T.J. Cade, M. Fuller, G. Hunt, and E. Potapov. The Peregrine Fund, Boise, Idaho, USA. Accessed 2 March 2018. http://www.peregrinefund.org/ subsites/conference-gyr/proceedings/. Braun, C.E., W.P. Taylor, and S.M. Ebbert. 2014. Changes in Evermann’s Rock Ptarmigan density on an eastern portion of Attu Island, Alaska, 2003-2009. Northwestern Naturalist 95: 28-34. https://doi.org/10. 1898/NWN13-10.1 Brodsky, L.M., and R.D. Montgomerie. 1987. Asym- metrical contests in defence of Rock Ptarmigan territor- ies. Behavioral Ecology and Sociobiology 21: 267—272. https://doi.org/10.1007/bf00292508 Bystrak, D. 1981. The North American breeding bird sur- vey. Studies in Avian Biology 6: 34—41. Cattadori, I.M., D.T. Haydon, and P.J. Hudson. 2005. Parasites and climate synchronize Red Grouse popula- tions. Nature 433: 737-740. https://do1.org/10.1038/nature 03276 Coats, R.R. 1956. Geology of northern Adak Island, Alaska. U.S. Department of the Interior, Geological Survey Bulletin 1028-C. U.S. Government Printing 54 THE CANADIAN FIELD-NATURALIST Office, Washington, DC, USA. Accessed 2 March 2018. https://pubs.usgs.gov/bul/1028c/report.pdf. Cotter, R.C. 1991. Population attributes and reproduct- ive biology of Rock Ptarmigan (Lagopus mutus) in the central Canadian Arctic. M.Sc. thesis, University of Alberta, Edmonton, Alberta, Canada. Cotter, R.C. 1999. The reproductive ecology of Rock Ptar- migan (Lagopus mutus) in the central Canadian Arctic. Arctic 52: 23-32. https://doi.org/10.14430/arctic 906 Dobson, A.P., and P.J. Hudson. 1992. Regulation and stability of a free-living host—parasite system: Tricho- strongylus tenuis in Red Grouse. II: Population mod- els. Journal of Animal Ecology 61: 487—498. https://doi. org/10.2307/5339 Droege, S. 1990. Survey designs and statistical methods for the estimation of avian population trends. Pages 1—4 in The North American Breeding Bird Survey. Edited by J.R. Sauer and S. Droege. Biological Report 90 (1). U.S. Department of the Interior, Fish and Wildlife Service, Washington, DC, USA. Accessed 2 March 2018. https:// pubs.er.usgs.gov/publication/5230211. Elton, C.S. 1924. Periodic fluctuations in the numbers of animals: their causes and effects. British Journal of Experimental Biology 2: 119-163. Accessed 2 March 2018. https://pdfs.semanticscholar.org/22fe/34 8adeftd9 2761 fa5a5396a7 1c7820f5b8c7. pdf. Everett, K.R. 1971. The structure and origin of the organic soil cover of Amchitka Island, Alaska. Bioscience 21: 618. https://do1.org/10.2307/1295733 Favaron, M., G.C. Scherini, D. Preatoni, G. Tosi, and L.A. Wauters. 2006. Spacing behavior and habitat use of rock ptarmigan (Lagopus mutus) at low density in the Italian Alps. Journal of Ornithology 147: 618-628. https://doi.org/10.1007/s10336-006-0087-z Fraser, G.D., and G.L. Snyder. 1959. Geology of south- ern Adak Island and Kagalaska Island, Alaska. U.S. Department of the Interior, Geological Survey Bulletin 1028-M. U.S. Government Printing Office, Washington, DC, USA. Accessed 2 March 2018. https://pubs.usgs. gov/bul/1028m/report. pdf. Gates, O., H.A. Powers, and R.E. Wilcox. 1971. Geology of the Near Islands, Alaska. U.S. Department of the Interior, Geological Survey Bulletin 1028-U. U.S. Government Printing Office, Washington, DC, USA. Accessed 2 March 2018. https://pubs.er.usgs.gov/publication/b1028U. Gibson, D.D., and B. Kessel. 1997. Inventory of the species and subspecies of Alaska birds. Western Birds 28: 45— 95. Accessed 2 March 2018. https://www.westernfield ornithologists.org/archive/V46/46(2)-p094-p185. pdf. Gilg, O., I. Hanski, and B. Sittler. 2003. Cyclic dynam- ics in a simple vertebrate predator-prey community. Science 302: 866-868. https://doi.org/10.1126/science. 1087509 Holder, K., and R. Montgomerie. 1993. Rock Ptarmigan (Lagopus mutus). In The Birds of North America, No. 51. Edited by A. Poole and F. Gill. Academy of Natural Sciences, Philadelphia, Pennsylvania, USA. Hudson, P.J., D. Newborn, and A.P. Dobson. 1992. Re- gulation and stability of a free-living host—parasite sys- tem: Trichostrongylus tenuis in Red Grouse. I: Monitor- ing and parasite reduction experiments. Journal of Animal Vol. 133 Ecology 61: 477—486. https://doi.org/10.2307/5339 Kaler, R.S.A., S.M. Ebbert, C.E. Braun, and B.K. San- dercock. 2010. Demography of a reintroduced popu- lation of Evermann’s Rock Ptarmigan in the Aleu- tian Islands. Wilson Journal of Ornithology 122: 1-14. https://doi.org/10.1676/08-099.1 Magnusson, K., J. Brynjarsdottir, and O.K. Nielson. 2004. Population cycles in Rock Ptarmigan Lagopus muta: modelling and parameter estimation. RH-19-2004, Raun- visindastofnun Haskolans, Iceland. Montgomerie, R., and K. Holder. 2008. Rock Ptarmigan (Lagopus muta). In The Birds of North America. Edited by A.F. Poole. Cornell Laboratory of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.51 Mougeot, F., S.M. Redpath, F. Leckie, and P.J. Hudson. 2003. The effect of aggressiveness on the population dy- namics of a territorial bird. Nature 421: 737—739. https:// doi.org/10.1038/nature01395 Nichols, J.D., and B.K. Williams. 2006. Monitoring for conservation. Trends in Ecology & Evolution 21: 668— 673. https://do1.org/10.1016/j.tree.2006.08.007 Pedersen, A.O., B.-J. Bardsen, N.G. Yoccoz, N. Lecomte, and E. Fuglei. 2012. Monitoring Svalbard Rock Ptar- migan: distance sampling and occupancy modeling. Journal of Wildlife Management 76: 308-316. https:// doi.org/10.1002/jwmg.276 Pedersen, A.O., O. Overrein, S. Unander, and E. Fuglei. 2005. Svalbard Rock Ptarmigan (Lagopus mutus hyper- boreus): a status report. Rapportserie No. 125, Norwe- gian Polar Institute, Tromso, Norway. Accessed 2 March 2018. http://hdl.handle.net/11250/173200. Pelletier, L., and C.J. Krebs. 1997. Line-transect sam- pling for estimating ptarmigan (Lagopus spp.) density. Canadian Journal of Zoology 75: 1185-1192. https://doi. org/10.1139/z97-141 Powers, H.A., R.R. Coats, and W.H. Nelson. 1960. Geology of and submarine physiology of Amchitka Island, Alaska. U.S. Department of Interior, Geological Survey Bulletin 1028-P. U.S. Government Printing Of- fice. Washington, DC, USA. Accessed 2 March 2018. https://pubs.usgs.gov/bul/1028p/report. pdf. Pruett, C.L., T.N. Turner, C.M. Topp, S.V. Zagrebelny, and K. Winker. 2010. Divergence in an archipelago and its conservation consequences in Aleutian Island Rock Ptarmigan. Conservation Genetics 11: 241-248. https:// doi.org/10.1007/s10592-009-0026-7 Ralph, C.J., S. Droege, and J.R. Sauer. 1995. Managing and monitoring birds using point counts: standards and applications. Pages 161-170 in Monitoring Bird Popu- lations by Point Counts. Edited by C.J. Ralph, J.R. Sauer, and S. Droege. U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station General Technical Report PSW-GTR-149. Albany, California, USA. Accessed 2 March 2018. https://www.fs. fed.us/ psw/publications/documents/gtr-149/pg161_ 168 .pdf. Robbins, C.S., D. Bystrak, and P.H. Geissler. 1986. The Breeding Bird Survey: its first fifteen years, 1965-1979. U.S. Department of the Interior, Fish and Wildlife Ser- vice Resource Publication 157. Accessed 2 March 2018. https://pubs.er.usgs.gov/publication/5230189. 2019 Sandercock, B.K., K. Martin, and S.J. Hannon. 2005. Life history strategies in extreme environments: com- parative demography of Arctic and alpine ptarmigan. Ecology 86: 2176-2186. https://doi.org/10.1890/04-0563 Sinclair, A.R.E., C.J. Krebs, J.N. Smith, and S. Boutin. 1988. Population biology of snowshoe hares. III. Nutri- tion, plant secondary compounds and food limitation. Journal of Animal Ecology 57: 787—806. https://doi. org/10.2307/5093 Taylor, W.P. 2013. Status of upland game within Alaska’s highway system: a comprehensive report focusing on 2007-2011. Wildlife Management Report, ADF&G/ DWC/WMR-2013-1. Alaska Department of Fish and Game, Division of Wildlife Conservation, Palmer, Alas- ka, USA. Accessed 12 August 2015. https://www.adfg. alaska.gov/static/research/programs/smallgame/pdfs/ upland_game_status_2007_2011.pdf. Tesar, C., M.-A. Dubois, and A. Shestakov. 2016. Toward strategic, coherent, policy-relevant Arctic science. Sci- ence 353 (6306): 1368-1370. https://doi.org/10.1126/ science.aai8198 Therrien, J.F., G. Gauthier, E. Korpimaki, and J. Béty. 2014. Predation pressure by avian predators suggests summer limitation of small-mammal populations in the Canadian Arctic. Ecology 95: 56—67. https://doi.org/10. SUPPLEMENTARY MATERIAL: BRAUN ET AL.: ROCK PTARMIGAN IN THE WESTERN ALEUTIAN ISLANDS 55 1890/13-0458.1 Watson, A. 1965. A population study of ptarmigan (Lagopus mutus) in Scotland. Journal of Animal Ecology 34: 135— 172. https://doi.org/10.2307/2373 Watson A., R. Moss, and P. Rothery. 2000. Weather and synchrony in 10-year population cycles of Rock Ptarmi- gan and Red Grouse in Scotland. Ecology 81: 2126— 2136. https://doi.org/10.1890/0012-9658(2000)081[2126: waslyp]2.0.co;2 Weatherbase. 2015. Travel weather averages for Adak, Alaska, USA. Accessed 12 August 2015. http://www. weatherbase.com/adak. Weeden, R.B. 1965. Breeding density, reproductive suc- cess and mortality of Rock Ptarmigan at Eagle Creek, central Alaska from 1960-1964. Transactions of the North American Wildlife Conference 30: 336-348. White, C.M., F.S.L. Williamson, and W.B. Emison. 1977. Avifaunal investigations. Pages 227-260 in The Environment of Amchitka Island. Edited by M.L. Mer- ritt and R.G. Fuller. U.S. Energy Research and Develop- ment Administration, TID-26712. U.S. Department of Energy, Washington, DC, USA. Received 3 May 2017 Accepted 6 February 2018 Appendix S1. Historical review of information and previous studies and surveys for Rock Ptarmigan (Lagopus muta) on three islands (Adak, Amchitka, and Attu) in the Western Aleutian Islands in Alaska. The Canadian Field-Naturalist Note Japanese Chafi-flower, Achyranthes japonica (Amaranthaceae), on the Erie islands, an invasive plant new to Canada JAMES KAMSTRA 560 Scugog Line 4, Port Perry, Ontario LOL 1Z8 Canada; email: jkamstra@powergate.ca Kamstra, J. 2019. Japanese Chaff-flower, Achyranthes japonica (Amaranthaceae), on the Erie islands, an invasive plant new to Canada. Canadian Field-Naturalist 133(1): 56-59. https://doi.org/10.22621/cfn.v133i1.2161 Abstract Japanese Chaff-flower, Achyranthes japonica (Miquel) Nakai (Amaranthaceae) was found growing on two islands in west- ern Lake Erie: East Sister Island and Middle Island. These are the first documented reports for this species in Canada, and these locations are approximately 300 km north of the nearest reported observations in southern Ohio. Japanese Chaff- flower is a non-native plant from Asia, which is highly invasive in the United States and has the potential to become so in Canada. Key words: Japanese Chaff-flower; Achyranthes japonica; East Sister Island; Middle Island; Ontario; non-native invasive plant; range extension During a visit to East Sister Island Provincial Nature Reserve in Lake Erie (Essex County, Ontario) on 27 September 2018, I found a small popula- tion of Japanese Chaff-flower, Achyranthes japoni- ca (Miquel) Nakai (Amaranthaceae) in the shade of moist deciduous forest co-dominated by American Elm (U/mus americana L.) and Common Hackberry (Celtis occidentalis L.; Figure 1). The location was at 41.81230°N, 82.85764°W in the island’s interior, at least 100 m from the Lake Erie shoreline. The popu- lation consisted of 15 plants, up to 50 cm tall. One specimen was collected. Two larger and taller stands of the plant were encountered on East Sister Island at ~41.8117°N, 82.8587°W in a small gap in a similar type of for- est and at 41.8120°N, 82.8582°W. These were ~50 m from the shoreline. Both stands were more robust and dense than those at the first location, each consisting of several dozen plants over 60 cm tall. They were growing on moist, level ground that may receive sea- sonal inundation and were partly shaded by Common Elderberry (Sambucus canadensis L.) and associated with Stinging Nettle (Urtica dioica L.), Common Pokeweed (Phytolacca americana L.), Spotted Jewel- weed (/mpatiens capensis Meerburgh), and Dwarf Clearweed (Pilea pumila (L.) A. Gray). On 5 October 2018, I visited Middle Island, part of Point Pelee National Park, also in Lake Erie and situated ~20 km southeast of East Sister Island. I observed two patches of Japanese Chaff-flower on the west end of that island within 15 m of the shore- line. Five plants were at the base of a limestone shin- gle berm in shade under a moist deciduous forest dominated by Common Hackberry at 41.68366°N, 82.68593°W. There were few other plants in the ground layer. The second larger nearby patch, at 41.68358°N, 82.68620°W, consisted of several dozen individuals. This patch was in semi-shade under for- est dominated by Common Hackberry, associated with Stinging Nettle and Common Pokeweed. The island was surveyed quite comprehensively and no other patches of Japanese Chaff-flower were found. T. Dobbie (pers. comm. 3 December 2018) reported finding and photographing a plant that she did not recognize while conducting a plant survey on Middle Island on 8 June 2018. The plant was nondescript with no fruit or flowers because it was early in the sea- son. She sent the photo to me, as I was now familiar with the species, and I confirmed that it was Japanese Chaff-flower, making that the earliest documentation of the species in Canada. Discussions with leading field botanists, the Canadian Food Inspection Agency (C. Wilson pers. comm. January 2019) and a check of the records and specimens available at the herb- aria of the Canadian Museum of Nature, Agriculture and Agri-Food Canada, the Database of Vascular A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. ©The Ottawa Field-Naturalists’ Club 2019 growing in partial shade. Photo: James Kamstra. Plants of Canada (VASCAN 2019), and the Ontario Ministry of Natural Resources and Foresty revealed no previous reports of this species in Canada. Japanese Chaff-flower is native to Japan, Korea, and China, where it is known for its medicinal prop- erties (Jung et al. 2007). The first North American record of Japanese Chaff-flower was from Martin County in eastern Kentucky, in August 1981 (Medley et al. 1985). Within a decade, the plant had domin- ated the floodplains in the area where it was first dis- covered and, within 15 years, had spread over 500 km along the Ohio River valley (Evans and Taylor 2011). The first documentation in Ohio was from 1992 (Vincent and Cusick 1998). In Flora of North America, Robertson (2003) indicated the plant’s pres- ence in Kentucky, West Virginia, and Ohio. Evans and Taylor (2011) mapped the distribution in the United States at the time showing that, by 2011, it had been confirmed in every county along the Ohio River from West Virginia to Illinois, with isolated popula- tions in Georgia and Tennessee. Japanese Chaff-flower is on the watch list for sev- eral states including Michigan (Michigan Invasive Species 2018) and Wisconsin (Wisconsin DNR 2018), both states where it has not yet been reported. None of the available mapping shows Japanese Chaff- flower to be currently present near Lake Erie. Both oo ae as “er. es Ficure 1. Japanese Chaff-flower (Achyranthes japonica) observed on East Sister Island, 27 September 2018. The plant was KAMSTRA: JAPANESE CHAFF-FLOWER, A NEW INVASIVE IN CANADA Dy EDDMapS (2018) and iNaturalist (2018) show that the nearest reported observations are in the vicin- ity of Cincinnati, Ohio, ~300 km south of the Erie Islands. R. Gardner (pers. comm. 22 January 2019), confirmed that, currently, Japanese Chaff-flower has been recorded only from the southern counties bor- dering the Ohio River in that state. At first glance, Japanese Chaff-flower superficially resembles Lopseed (Phryma leptostachya L.) because of the narrow erect spikes crowded with deflexed fruits and opposite leaf arrangement. However, the flowers of Japanese Chaff-flower have five deflexed tepals, and the ovate-elliptical leaves are glabrous and lack teeth (Robertson 2003). The fruits hang tightly on the spike; each one contains a pair of spiny bracts that adhere to fur and feathers. The seeds are mainly spread by animals or water transport (Evans and Taylor 2011). Japanese Chaff-flower is typical- ly 0.75—1.5 m tall (Robertson 2003), but can reach 3 m (Schwartz er al. 2016). Throughout its introduced range in the United States, it most frequently grows in semi-shaded moist soils, but can also occur in drier and sunny sites (Schwartz et al. 2016) and is, there- fore, capable of colonizing a variety of habitats. The origin of the plants on the two Erie islands can only be speculated. Middle Island is ~18.5 ha in area and located 4.5 km south of Pelee Island, the 58 THE CANADIAN FIELD-NATURALIST nearest land mass. East Sister Island is ~13 ha in area and more remote at 10 km north of North Bass Island, Ohio, the nearest land mass of more than 2.5 ha. Both East Sister and Middle Islands support large nesting colonies of Double-crested Cormorants (Phalacrocorax auritus) numbering in the thousands (McGrath and Murphy 2012) as well as smaller num- bers of nesting Herring Gulls (Larus argentatus), Black-crowned Night-herons (Nycticorax nyctico- rax), Great Blue Herons (Ardea herodius), and Great Egrets (Ardea alba; IBA Canada 2018). Because the seeds of Japanese Chaff-flower can readily attach to fur or feathers, the plants may have been carried to both islands by cormorants or other birds. Cormorant nests were present in trees in the vicinity of Japanese Chaff-flower patches on both islands. Choi et al. (2010) examined nearly 4000 birds for the presence of plant propagules on a remote is- land off Korea. Three species of migratory birds were found to have seeds of Japanese Chaff-flower at- tached to their feathers, including two marsh species: Eurasian Bittern (Botaurus stellaris) and Swinhoe’s Rail (Coturnicops exquisitus). Choi et al. (2010) sug- gest that birds may have been responsible for the spread of Japanese Chaff-flower to offshore islands in Korea where it has become highly invasive. Considering the relatively intense floristic survey of the Lake Erie Islands (Duncan ef a/. 2010), it seems likely that Japanese Chaff-flower is a recent arrival. Given that Japanese Chaff-flower is not known from the south shore of Lake Erie (R. Gardner pers. comm. 22 January 2019), it seems likely that birds were the means for its spread onto East Sister and Middle Islands. The sheer abundance of cormorants, which nest all over both islands, make them a likely vec- tor. Japanese Chaff-flower also spreads by water, but the plant is not known to be present elsewhere along the Lake Erie shore. Furthermore, the plants found on East Sister Island were inland and not along the immediate shoreline. Japanese Chaff-flower has the potential to become an aggressive invasive plant in southern Ontario. Because of the seriousness of this new threat, sev- eral fact sheets have been produced to inform the public about control methods and encourage them to report sightings (Evans and Taylor 2011; Rathfon and Eubank 2013; Schwartz et a/. 2015). The plant has spread rapidly along river systems in the United States and has been identified as a high priority inva- sive in Indiana (Rathfon and Eubank 2013). A single large plant can produce more than 1000 seeds and 94% of the seeds have been shown to be viable (Evans and Taylor 2011). Infestations can attain densities of more than 70 plants/m2, which will shade out all other plants below them (Evans and Taylor 2011). They also Vol. 133 have the ability to invade undisturbed forests that have not been previously impacted (Schwartz et al. 2016). The United States Department of Agriculture (USDA 2014) has evaluated the plant’s weed risk po- tential and, based on its native range, determined that it can survive in hardiness zones 5—10. Consequently, it has the potential to spread to all of Ontario south of the Canadian Shield, as well as parts of Quebec, the Maritimes, and even parts of British Columbia. The Canadian Food Inspection Agency, Canada’s national plant protection authority, is conducting a full risk as- sessment of the Japanese Chaff-flower to determine its invasive potential in the country (C. Wilson pers. comm. January 2019). Control, research, and ideally eradication should be high priorities before the plant gains a strong foothold, although this may not be pos- sible given its spread in the USA. Voucher specimens Canada, Ontario: Essex County, Pelee Township, East Sister Island. 41.81230°N, 82.85764°W. About 15 plants growing in moist soil in deciduous forest co-dominated by American Elm and Common Hack- berry, 27 September 2018, CAN-10091002 (CAN). Canada, Ontario: Essex County, Pelee Township, Middle Island, Point Pelee National Park. 41.68358°N, 82.68620°W. About 35 plants growing in moist soil near shoreline in deciduous forest dominated by Common Hackberry, 5 October 2018, CAN-10091001 (CAN). Acknowledgements Michael J. Oldham of the Natural Heritage In- formation Centre provided literature references and many useful comments and reviewed the draft. Dr. Tony Reznicek from the University of Michigan in- itially identified the plant from photographs and re- viewed the draft. Y. Robert Tymstra accompanied me on field visits and also searched Middle Sister Island for the plant. Tammy Dobbie sent photographs and details about plants that she observed on Middle Island. Richard Gardner, chief botanist with the Ohio Department of Natural Resources, provided informa- tion on the species in Ohio. Claire Wilson, with the Plant Health Risk Assessment Unit of the Canadian Food Inspection Agency, provided comments from the perspective of that agency. Literature Cited Choi, C., H. Nam, and H. Chae. 2010. Exotic seeds on the feathers of migratory birds on a stopover island in Korea. Journal of Ecology and Field Biology 33: 19-22. https://do.org/10.5141/jefb.2010.33.1.019 Duncan, T., L. Brohl, J. Kartesz, M.J. Oldham, and R.J. Stuckey. 2010. Flora of the Erie Islands: a review of floristic, ecological and historical research and con- servation Activities, 1976-2010. Ohio Journal of Sci- 2019 ence 110: 3-12. Accessed December 2018. https://pdfs. semanticscholar.org/4d20/c50bdf257ef3b0ccca0dIcf07 ed86ed03368 pdf. EDDMapS. 2018. Japanese Chaff-flower: Achyranthes Japonica (Migq.) Nakai. Early Detection and Distribution Mapping System, Center for Invasive Species and Eco- system Health, University of Georgia, Tifton, Georgia, USA. Accessed November 2018. http://www.eddmaps. org/distribution/viewmap.cfm?sub=14211. Evans, C., and D.D. Taylor. 2011. New invader profile: Japanese Chaff-flower—Achyranthes japonica. Wild- land Weeds Summer/Fall: 4—6. IBA (Important Bird Areas) Canada. 2018. Pelee Island Archipelago, Western Lake Erie Basin, Ontario. Site Summary ONOI4. Bird Studies Canada, Port Rowan, Ontario, Canada. Accessed November 2018. https://tiny url.com/y5jqa9cw. iNaturalist. 2018. Japanese Chaff Flower. California Aca- demy of Sciences, San Francisco, California, USA. Accessed November 2018. https://inaturalist.ca/observa tions?place_id=97394&taxon_1d=157962. Jung, S.M., S.I. Choi, S.M. Park, and T.R. Heo. 2007. Antimicrobial effect of Achyranthes japonica Nakai extracts against Clostridium difficile. Korean Journal of Food Science and Technology 39: 564-568. McGrath, D.M., and S.D. Murphy. 2012. Double-crested Cormorant (Phalacrocorax auritus) nesting effects on understory composition and diversity on island eco- systems in Lake Erie. Environmental Management 50: 304-314. Medley, M.E., H. Bryan, J. MacGregor, and J.W. Thie- ret. 1985. Achyranthes japonica (Miq.) Nakai (Amaran- thaceae) in Kentucky and West Virginia: new to North America. Sida 11: 92-95. Michigan Invasive Species. 2018. Japanese Chaff Flower. Government of Michigan, Lansing, Michigan, USA. Accessed November 2018. https://www.michigan.gov/ invasives/0,5664,7-324-68002_ 74188-476192--,00. html. Rathfon, R., and E. Eubank. 2013. Japanese Chaff Flower. Invasive plant series fact sheets. Purdue Extension, KAMSTRA: JAPANESE CHAFF-FLOWER, A NEW INVASIVE IN CANADA 59 Lafayette, Indiana, USA. Accessed December 2018. https://www.extension. purdue.edu/extmedia/FNR/ FNR-477-W pdf. Robertson, K.R. 2003. Achyranthes L. Pages 435, 437 in Flora of North America. Volume 4. Edited by Flora of North America Editorial Committee. eFloras.org, New York, New York, USA. Accessed December 2018. http:// www.efloras.org/florataxon.aspx?flora_id=l&taxon_ id=242300552. Schwartz, L.M., D.J. Gibson, and B.G. Young. 2016. Life history of Achyranthes japonica (Amaranthaceae): an invasive species in southern Illinois. Journal of the Torrey Botanical Society 143: 93-102. https://doi. org/10.3159/torrey-d-14-00014 Schwartz, L.M., K.M. Smith, C. Evans, K.L. Gage, D.J. Gibson, and B.G. Young. 2015. Fact sheet: ecol- ogy and control of Japanese Chaff Flower [Achyranthes Japonica (Miq.) Nakai]. Accessed 19 July 2019. https:// bugwoodcloud.org/mura/rtrcwma/assets/File/Chaff_ FactSheet.pdf. USDA (United States Department of Agriculture). 2014. Weed risk assessment for Achyranthes japonica (Miq.) Nakai—Japanese Chaff-flower. Accessed December 2018. https://www.aphis.usda.gov/plant_health/plant_ pest_info/weeds/downloads/wra/Achyranthes%20 Japonica%20WRA. pdf. VASCAN (Vascular Plants of Canada). 2019. Canadensys. Accessed January 2019. http://data.canadensys.net/vas can/search. Vincent, M.A., and A.W. Cusick. 1998. New records of alien species in the Ohio vascular flora. Ohio Journal of Science 98: 10-17. Wisconsin Department of Natural Resources. 2018. Japanese chaff flower (Achyranthes japonica). Wiscon- sin Department of Natural Resources, Madison, Wis- consin, USA. Accessed December 2018. https://dnr.wi. gov/topic/Invasives/fact/JapaneseChaffF lower.html. Received 4 December 2018 Accepted 7 February 2019 The Canadian Field-Naturalist Gray Wolf (Canis lupus) recolonization failure: a Minnesota case study L. Davip MeEcu!*, Forest ISBELL’, JIM KRUEGER?, and JOHN HART* ‘United States Geological Survey, Northern Prairie Wildlife Research Center, 8711 — 37th Street SE, Jamestown, North Dakota 58401-7317 USA *Department of Ecology, Evolution and Behavior, University of Minnesota, St. Paul, Minnesota 55108 USA >Cedar Creek Ecosystems Science Reserve, University of Minnesota, East Bethel, Minnesota 55005 USA ‘United States Department of Agriculture, Wildlife Services, Grand Rapids, Minnesota 55744 USA “Corresponding author: mechx002@umn.edu Mech, L.D., F. Isbell, J. Krueger, and J. Hart. 2019. Gray Wolf (Canis /upus) recolonization failure: a Minnesota case study. Canadian Field-Naturalist 133(1): 60-65. https://doi.org/10.22621/cfn.v13311.2078 Abstract During the past few decades, Gray Wolves (Canis /upus) have recolonized many areas in the United States and Europe. In many other cases, however, although dispersing wolves reached areas with adequate prey, a population failed to recolon- ize. Herein, we provide a case study detailing how a wolf pack attempted for three years to recolonize an area 55 km froma long-established population and within 25 km of Minneapolis and St. Paul, Minnesota, but failed. The pack produced three litters of pups and at one time included 11-19 members, but it preyed on livestock and dogs and, consequently, was lethally removed. The history of this pack’s attempt to recolonize an area long devoid of wolves exemplifies the issues that have pre- vented earlier recolonizations in non-wild lands in Minnesota and elsewhere and that promise to do so well into the future. Key words: Canis lupus, depredation; distribution; Gray Wolf; livestock; recolonization Introduction During the past several decades, Gray Wolves (Canis [upus) have been recolonizing many areas of the world (Boitani 2003; Chapron et al. 2014; Mech 2017). In the contiguous United States, they have recolonized Wisconsin, Michigan, the northwestern USA, and new areas of Minnesota, and are dispersing into adjacent states (Mech 2017). Biologically, wolves are prolific and can survive anywhere with sufficient food. Because they can subsist not only on prey but also on carrion and even garbage, the only constraints on where they recolonize are anthropogenic factors, including vehicle strikes, legal harvest, illegal killing (including poisoning), and legal livestock-depreda- tion control. Humans persecuted wolves throughout much of their original range; thus, those that survived lived primarily in wilderness or areas with low human density. That gave some biologists the impression that wilderness was required for their survival, and early models to predict potential wolf habitat in the Upper Midwest made that assumption (Mladenoff et al. 1995, 1999, 2006), although it was later challenged (Mech 2006a,b). Eventually the models were refined (Mladenoff et al. 2009) to reflect the fact that wolves do not require wilderness (Mech 2015). However, to survive and repopulate a new location for multiple generations, wolves do need to avoid areas and be- haviours that bring them into conflict with human ac- tivities (Erb and Don Carlos 2009; Mech 2017). In Minnesota, wolves have been expanding their range from a wilderness reservoir in the northeast- ern part of the state. Since the early 1970s, they have been gradually recolonizing westward and southward toward semi-wilderness, agricultural areas, and a ma- jor metropolitan area (Fuller et a/. 1992; Erb and Don Carlos 2009; Erb et al. 2017). As their numbers and distribution have increased, so have depredations of livestock and the number of wolves killed for live- stock-depredation control (Mech 1998; Harper eg al. 2005; Ruid et al. 2009). By 1997-1998, the annu- ally estimated Minnesota wolf population of 2445— 2856 had reached the extent of its current distribution (Figure 1) and has since failed to further recolonize the state (Berg and Benson 1999; Erb et al. 2017). Individual maturing male and female wolves have dispersed far and wide from their northern Minnesota reservoir to all parts of the state and have entered nearby states including Wisconsin, Michigan, South Dakota, and North Dakota (Fritts and Mech 1981; Gese and Mech 1991; Merrill and Mech 2000). To recolonize a new area, unrelated males and females ©This work is made available under the Creative Commons CCO 1.0 Universal Public Domain Dedication (CCO 1.0). 2019 See! rf ! I ——_ = _— 5 km = = oe MECH &T AL.: WOLF RECOLONIZATION FAILURE 61 ~, eee * New pack. x Mpls-St. Paul@® Ficure 1. Study area where the Isanti Gray Wolf (Canis /upus) pack attempted to recolonize. Dashed line connects outer- most locations where wolf signs were found and represents the minimum area the pack used from 2014 through 2017. Solid line represents approximate boundary of the Cedar Creek Ecosystem Science Reserve. must find each other in a suitable location, establish a territory there, pair bond, produce pups, and sur- vive for several years. If pets or livestock are avail- able locally, resident wolves often begin preying on them. Such depredations decrease human toler- ance of wolves (Williams et a/. 2002; Karlsson and Sjostrom 2007; Olson et al. 2015), and state and/or federal wolf depredation control agencies often leth- ally remove them or translocate them depending on applicable laws. Thus, wolves are only able to recol- onize areas with low human presence. In Minnesota, wolves have attempted to recolon- ize and establish a breeding population southward ~25 km north of the Minneapolis—St. Paul suburbs, at about 45°43'N (Erb and Don Carlos 2009; Erb et al. 2017). During 1997, a pack or pair was recorded about 45 km west of there near the Sherburne National Wildlife Refuge (Berg and Benson 1999), but by 2004 that pack no longer existed for reasons unknown (Erb and Benson 2004). In 2010-2011, a new pack survived for two years about 25 km south of the current wolf range, but two adults, a yearling, and four pups were then lethally removed for depre- dation control. In 2014, a new pack (the Isanti pack) formed 55 km south-southeast of the current wolf range, and within 25 km of the Minneapolis—St. Paul suburbs in an area with O-10% chance of wolf recolonization according to the latest wolf habitat models, which consider road density and agriculture (Mladenoff er al. 2009). This article details the 3-year attempt by wolves to recolonize that area. Study Area The study area (at about 45°27'N, 93°08'W) com- prises ~80 km? in northern Anoka and southern Isanti counties in east-central Minnesota (Figure 1). Most of the area is rural residential and agricul- tural, interspersed with patches of uninhabited low- land and woodlots, the largest being the University of Minnesota Cedar Creek Ecosystem Science Reserve (CCESR) covering about 21 km”. Roughly 50-60% of the study area is open agricultural fields, and the area is heavily roaded; the most remote location in the area is 1.54 km from the nearest road. Much of the known territory of the Isanti pack fell in Athens township, which had a 2016 population density of 24 people/ km? (Towncharts 2018) and Linwood township with a density of 62 people/km? (calculated from the town- ship area of 84.992 km? and the 2016 total popula- tion of 5284; American Factfinder 2019). Estimated pre-fawning White-tailed Deer (Odocoileus virgin- ianus) density for this area of the state in 2016 was 8.5 deer/km? (D’Angelo et al. 2016), and Wild Turkeys (Meleagris gallopavo) were common. Small herds of cattle are widely scattered throughout the study area. Some 9452 cattle, including calves, occupied Isanti County (1157 km?) in 2012 (USDA 2012). Domestic dogs are common, and some are free-ranging. 62 Methods and Results The first record of wolves having bred in the study area was a trail camera photo of three and possibly four adult-sized wolves during winter 2014—2015. Although a pair of wolves can form at any time, a pair with at least one other adult-sized wolf in win- ter would almost certainly indicate that the pair had established a territory and produced at least one pup, most likely in the previous spring (Mech and Boitani 2003). There was also a report of a Coyote (Canis latrans) trapper catching a wolf in the area in win- ter 2014-2015. During summer 2015, wolves denned on the CCESR within 1.4 km of an occupied residence and produced at least eight pups that were observed and photographed several times. Throughout summer, researchers associated with the reserve frequented areas within 100 m of the den multiple times a week during the course of their previously established research. During autumn 2015, nine wolves were seen twice on CCESR property and, in November 2015, 11 (which could indicate that nine pups were produced). In mid-January 2016, a Coyote trapper captured and released a wolf from a snare just outside the CCESR. In mid-winter 2015-2016, L.D.M. drove the roads throughout the study area and found several places where, between 1 January and 6 February 2016, up to eight wolves had crossed. The greatest distance between locations where wolf tracks, or in one case wolf fur on a barbed-wire fence, were found was 14 km, with the centre of that area being 5.5 km from the 2015 den (Figure 1). By calculating the area enclosed by all the locations where such wolf sign was found, we estimated that the minimum area used by this wolf pack was 80 km7. From August 2015 to April 2016, within the area covered by these wolves, three cattle were killed and THE CANADIAN FIELD-NATURALIST Vol. 133 one wounded, and three dogs were killed by wolves (Table 1). Thus, in April 2016, Wildlife Services, the federal government’s depredation control agency (Ruid et al. 2009), lethally removed three male wolves, weighing 35, 42, and 47 kg. Trail cameras on the CCESR continued to rec- ord wolf presence throughout summer 2016. In June, wolves killed a 91-kg calf, and Wildlife Services leth- ally removed a 36-kg male wolf, a 27-kg yearling female, and a 32-kg breeding female from the study area; sign of additional adult wolves remained. Four pups were captured alive during depredation control in late June and released on site according to United States Fish and Wildlife Service requirements that all young of the year be released before 2 August when wolves are protected by the Endangered Species Act as they were in 2016. One pup was dead in a snare so could not be released. Local residents had reported seeing up to eight pups nearby before this. Because the 2015 den at the CCESR was unused in 2016, and trail cameras on CCESR failed to indicate concentrated wolf use of the CCESR, the 2016 den was very likely off CCESR property. Based on where the five pups were caught in late June 2016, on reports of local residents, and on the nearest remote area, we judged that the 2016 den was about 10 km east-northeast of the 2015 den. During autumn 2016, trail camera photos indica- ted that at least one wolf still used the study area, and, in May 2017, wolves killed another calf in the same area as the 2016 depredations; Wildlife Services leth- ally removed a 32-kg male wolf and a 26-kg, non- breeding female. Since 2015, all but one complaint of wolves attacking livestock or dogs in this area were verified by authorities. As of February 2019, CCESR trail cameras have recorded only a single wolf. TABLE 1. Estimated numbers of Gray Wolves (Canis /upus), verified complaints, and numbers of wolves removed by year for the Isanti pack, Minnesota. 2014 Estimated number of wolves in Isanti pack Adults/yearlings 2 Pups Pal Total oo Number of verified complaints Dog complaints 0 Livestock complaints 0 Total 0 Number of wolves removed Adults/yearlings 0 Pups 0 Total 0 2015 2016 2017 3 1 3 8 8 0 11 19* 3 2 1 0 1 2 i 3 3 1 0 6 2 0 1 0 0 TF 2 *Assumes eight pups were born in early April before three adults were lethally removed later in that month. tIn addition to the seven wolves removed, four wolf pups were captured and released on site according to United States Fish and Wildlife Service guidelines because they were caught before 2 August. 2019 Discussion Although wolves have recolonized much of the northern half of Minnesota as well as many areas of Wisconsin and Michigan over the last few decades, they have failed to recolonize many other adjacent areas with adequate natural prey. These latter areas are those with considerable populations of people and domestic animals. However, it is not for lack of try- ing (Mech 2017). This case history illustrates the details of how and when wolves begin to establish in areas with live- stock and dogs, they may begin treating these domes- tic animals as natural prey. This usually happens soon after the wolves start reproducing, especially when a third age class is present. Domestic animals are easy targets because they can nearly always be found in the same place, unlike most natural prey, which require hunting down. The increase in domes- tic animal depredations with the presence of a third age class or a larger pack (Bradley et al. 2015) may result from reduced natural local food resources and more dependent wolves to feed. Regions similar to our study area were predicted to have probabilities of wolf recolonization of 0O-10% (Mladenoff et al. 1995, 1999, 2006), and our findings explain why. Wolves can and do inhabit these areas (Mech 2006a,b) but tend to persist longer in wilder- ness and wild lands where they conflict much less with human interests (Mech 2017). Given the great variation in land use across large areas, gradients of wolf-recolonization suitability exist; thus, along the frontiers of established wolf populations, wolves will continue to attempt to expand into areas with higher predicted probabilities of recolonization, with varied results. The large body masses of the wolves captured in this study area showed that their lack of success in recolonization and their predation on domestic prey were not because they were desperate for food. All the wolves caught were in excellent condition. Four of the eight were above average for wolves feed- ing on all-natural prey (Mech 2006c), including the 47-kg male that weighed more than all but two of 873 captures of Minnesota wolves on a natural-prey diet (L.D.M. and S.B. Barber-Meyer unpubl. data). Despite living among people and livestock close to the suburbs of Minneapolis and St. Paul, the Isanti wolf pack was able to use small areas away from humans to den and raise their young and, in that way, persist for at least three years. Like so many other wolf attempts to recolonize similar areas of Minnesota and other states, this one nevertheless failed because of the conflict that often results from wolves living close to areas with high densities of people, livestock, and pets. Wolf survival in the long term requires large MECH &£T AL.: WOLF RECOLONIZATION FAILURE 63 areas of extensive wild lands (Young and Goldman 1944; Mech 1970, 2017; Ruid et al. 2009). This case study details why. Author Contributions Writing — Original Draft: L.D.M.; Writing — Re- view & Editing: L.D.M., FL, J.K., and J.H.; Conceptu- alization: L.D.M. and F.I; Investigation: L.D.M., FL, J.K., and J.H.; Methodology: L.D.M., FI, J.K., and J.H.; Formal Analysis: L.D.M.; Funding Acquisition: L.D.M. and FI. Acknowledgements Funding for this project was provided by the Unit- ed States Geological Survey (USGS), the Minneso- ta Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Com- mission on Minnesota Resources, and the United States Department of Agriculture Wildlife Services. We thank Larry Hogie for cooperating with the study, S.B. Barber-Meyer of the USGS for use of un- published data, and Quinn Harrison, University of Minnesota, for critiquing an earlier draft and offering helpful suggestions for improving it. Literature Cited American Factfinder. 2019. Linwood township, Anoka, Minnesota. United States Census, Washington, DC, USA. Accessed 29 May 2019. https://factfinder.census. gov/faces/tableservices/jsf/pages/productview. xhtml? src=CF. Berg, W., and S. Benson. 1999. Updated wolf population estimate for Minnesota 1997-1998. Minnesota Depart- ment of Natural Resources, Grand Rapids, Minnesota, USA. Boitani, L. 2003. Wolf conservation and recovery. Pages 317-340 in Wolves: Behavior, Ecology, and Conserva- tion. Edited by L.D. Mech and L. Boitani. University of Chicago Press, Chicago, Illinois, USA. Bradley, E.H., H.S. Robinson, E.E. Bangs, K. Kunkel, M.D. Jimenez, J.A. Gude, and T. Grimm. 2015. Effects of wolf removal on livestock depredation recur- rence and wolf recovery in Montana, Idaho, and Wyo- ming. Journal of Wildlife Management 79: 1337-1346. https://doi.org/10.1002/jwmg.948 Chapron, G., P. Kaczensky, J.D.C. Linnell, M. von Arx, D. Huber, H. Andren, J.V. Lépez-Bao, M. Adamec, F. Alvares, O. Anders, L. Baléiauskas, V. Balys, P. Bedé, F. Bego, J.C. Blanco, U. Breitenmoser, H. Bre- seth, L. Bufka, R. Bunikyte, P. Ciucci, A. Dutsov, T. Engleder, C. Fuxjager, C. Groff, K. Holmala, B. Hox- ha, Y. Iliopoulos, O. Ionescu, J. Jeremié, K. Jerina, G. Kluth, F. Knauer, I. Kojola, I. Kos, M. Krofel, J. Kubala, S. Kunovac, J. Kusak, M. Kutal, O. Liberg, A. Majié, P. Mannil, R. Manz, E. Marboutin, F. Marucco, D. Melovski, K. Mersini, Y. Mertzanis, R.W. Mystajek, S. Nowak, J. Odden, J. Ozolins, G. Palomero, M. Paunovicé, J. Persson, H. Potoénik, P.-Y. Quenette, G. Rauer, I. Reinhardt, R. Rigg, A. Ryser, 64 THE CANADIAN FIELD-NATURALIST V. Salvatori, T. SkrbinSek, A. Stojanov, J.E. Swenson, L. Szemethy, A. Trajce, E. Tsingarska-Sedefcheva, M. Vata, R. Veeroja, P. Wabakken, M. Wé6lfl, S. Wéolfl, F. Zimmermann, D. Zlatanova, and L. Boitani. 2014. Recovery of large carnivores in Europe’s modern human-dominated landscapes. Science 346: 1517-1519. https://do1.org/10.1126/science.1257553 D’Angelo, G.J., and J.H. Giudice. 2016. Monitoring popu- lation trends of White-tailed Deer in Minnesota. Min- nesota Department of Natural Resources, St. Paul, Minnesota, USA. Accessed 24 May 2019. http://files. dnr.state.mn.us/wildlife/deer/reports/popmodel/pop model_2016.pdf. Erb, J., and S. Benson. 2004. Distribution and abun- dance of wolves in Minnesota, 2003-04. Minnesota Department of Natural Resources, St. Paul, Minnesota, USA. Accessed 11 February 2019. https://www.leg. state.mn.us/docs/2015/other/150681/PFEISref_1/Erb %20and%20Benson%202004. pdf. Erb, J., and M. Don Carlos. 2009. An overview of the legal history and population status of wolves in Minnesota. Pages 49-64 in Recovery of Gray Wolves in the Great Lakes Region of the United States: an Endangered Species Success Story. Edited by S.A.P. Wydeven, T.R. Van Deelen, and E.J. Heske. Springer, New York, New York, USA. https://doi.org/10.1007/ 978-0-387-85952-1_4 Erb, J., C. Humpal, and B. Sampson. 2017. Minnesota wolf population update 2017. Minnesota Department of Natural Resources, St. Paul, Minnesota, USA. Accessed 11 February 2019. https://files.dnr.state.mn.us/wildlife/ wolves/2017/survey-wolf. pdf. Fritts, S.H., and L.D. Mech. 1981. Dynamics, Movements, and Feeding Ecology of a Newly Protected Wolf Popu- lation in Northwestern Minnesota. Wildlife Monographs 80. Wildlife Society, Bethesda, Maryland, USA. Fuller, T.K., W.E. Berg, G.L. Radde, M.S. Lenarz, and G.B. Joselyn. 1992. A history and current estimate of wolf distribution and numbers in Minnesota. Wildlife Society Bulletin 20: 42-55. Gese, E.M., and L.D. Mech. 1991. Dispersal of wolves (Canis lupus) in northeastern Minnesota, 1969-1989. Canadian Journal of Zoology 69: 2946-2955. https:// doi.org/10.1139/z91-415 Harper, E.K., W.J. Paul, and L.D. Mech. 2005. Causes of wolf depredation increase in Minnesota from 1979— 1998. Wildlife Society Bulletin 33: 888-896. Karlsson, J., and M. Sjostrom. 2007. Human attitudes towards wolves, a matter of distance. Biological Con- servation 137: 610—616. https://doi.org/10.1016/j.biocon. 2007.03.023 Mech, L.D. 1970. The Wolf: the Ecology and Behavior of an Endangered Species. Natural History Press, Doubleday Publishing Co., New York, New York, USA. Mech, L.D. 1998. Estimated costs of maintaining a re- covered wolf population in agricultural regions of Min- nesota. Wildlife Society Bulletin 26: 817-822. Mech, L.D. 2006a. Prediction failure of a wolf landscape model. Wildlife Society Bulletin 34: 874-877. https:// doi.org/10.2193/0091-7648(2006)34[874: pfoawl]2.0 .co;2 Vol. 133 Mech, L.D. 2006b. Mladenoff et al. rebut lacks supportive data. Wildlife Society Bulletin 34: 882—883. https://doi. org/10.2193/0091-7648(2006)34[882:mearls]2.0.co;2 Mech, L.D. 2006c. Age-related body mass and repro- ductive measurements of gray wolves in Minnesota. Journal of Mammalogy 87: 80-84. https://doi.org/10. 1644/05-mamm-f-212r1.1 Mech, L.D. 2015. Wisconsin wolf management: a cauldron of controversy. Pages 223-225 in Conflicts in Conser- vation: Navigating Towards Solutions. Edited by S.M. Redpath, R.J. Gutierrez, K.A. Wood, and J.C. Young. Cambridge University Press, Cambridge, United King- dom. Mech, L.D. 2017. Where can wolves live and how can we live with them? Biological Conservation 210: 310-317. https://doi.org/10.1016/}.biocon.2017.04.029 Mech, L.D., and L. Boitani. 2003. Wolf Social Ecology. Pages 1-34 in Wolves: Behavior, Ecology, and Con- servation. Edited by L.D. Mech and L. Boitani. Univer- sity of Chicago Press, Chicago, Illinois, USA. Merrill, S.B., and L.D. Mech. 2000. Details of extensive movements by Minnesota wolves. American Midland Naturalist 144: 428-433. Mladenoff, D.J., M.K. Clayton, S.D. Pratt, T.A. Sickley, and A.P. Wydeven. 2009. Change in unoccupied wolf habitat in the northern Great Lakes Region. Pages 119— 138 in Recovery of Gray Wolves in the Great Lakes Re- gion of the United States: an Endangered Species Suc- cess Story. Edited by A.P. Wydeven, T.R. Van Deelen, and E.J. Heske. Springer, New York, New York, USA. Mladenoff, D.J., M.K. Clayton, T.A. Sickley, and A.P. Wydeven. 2006. L.D. Mech critique of our work lacks scientific validity. Wildlife Society Bulletin 34: 878-881. Mladenoff, D.J., T.A. Sickley, R.G. Haight, and A.P. Wydeven. 1995. A regional landscape analysis and pre- dication of favorable gray wolf habitat in the Northern Great Lakes Region. Conservation Biology 9: 279-294. https://doi.org/10.1046/j.1523-1739.1995.9020279.x Mladenoff, D.J., T.A. Sickley, and A.P. Wydeven. 1999. Predicting pray wolf landscape recolonization: logistic regression models vs new field data. Ecological Appli- cations 9: 37—44. Olson, E.R., J.L. Stenglein, V. Shelley, A.R. Rissman, C. Browne-Nunez, Z. Voyles, A.P. Wydeven, and T. Van Deelen. 2015. Pendulum swings in wolf management led to conflict, illegal kills and a legislated wolf hunt. Conservation Letters 8: 351-360. https://doi.org/10.1111/ conl.12141 Ruid, D.B., W.J. Paul, B.J. Roell, A.P. Wydeven, R.C. Willging, R.L. Jurewicz, and D.H. Lonsway. 2009. Wolf—human conflicts and management in Minnesota, Wisconsin and Michigan. Pages 279-295 in Recovery of Gray Wolves in the Great Lakes Region of the United States: an Endangered Species Success Story. Edited by A.P. Wydeven, T.R. Van Deelen, and E.J. Heske. Springer, New York, New York, USA. https://doi.org/10. 1007/978-0-387-85952-1_ 18 TownCharts.com. 2018. Athens township, Minnesota de- mographics data. TownCharts. Accessed 9 April 2018. http://www.towncharts.com/Minnesota/Demographics/ Athens-township-MN-Demographics-data.html. 2019 MECH &T AL.: WOLF RECOLONIZATION FAILURE 65 USDA (United States Department of Agriculture). 2012. and their reintroduction (1972—2000). Wildlife Society State and county profiles: Isanti County, Minnesota. Bulletin 30: 575-584. USDA, Washington, DC, USA. Accessed 24 May 2019. Young, S.P., and E.A. Goldman. 1944. The Wolves of https://www.nass.usda.gov/Publications/AgCensus/ North America. American Wildlife Institute, Washing- 2012/Online_Resources/County_Profiles/Minnesota/ ton, DC, USA. cp27059. pdf. Williams, C.K., G. Ericsson, and T.A. Heberlein. 2002. Received 13 April 2018 A quantitative summary of attitudes toward wolves Accepted 19 February 2019 The Canadian Field-Naturalist Book Reviews Book Review Editor’s Note: The Canadian Field-Naturalist is a peer-reviewed scientific journal publishing papers on ecology, behaviour, taxonomy, conservation, and other topics relevant to Canadian natural history. In line with this mandate, we review books with a Canadian connection, including those on any species (na- tive or non-native) that inhabits Canada, as well books covering topics of global relevance, including climate change, biodiversity, species extinction, habitat loss, evolution, and field research experiences. Currency Codes: CAD Canadian Dollars, USD United States Dollars, EUR Euros, AUD Australian Dollars, GBP British Pound. CLIMATE CHANGE The Uninhabitable Earth: Life After Warming By David Wallace-Wells. 2019. Tim Duggan Books, Penguin Random House. 320 pages, 27.00 USD, Cloth. “We run carelessly over the precipice after having put something in front of us to The prevent us seeing it”, —Blaise = Uninhabitable Pascal, Pensées (1623-1662) Earth I can summarize this re- Se neo aaa view in a single sentence. David Everyone should be com- Wallace-Wells pelled to read this book to truly appreciate the nature of the threat from climate change. David Wallace-Wells lays out in elegant, yet blunt, language the nature and potential extent of the in- evitable disruption to humanity from climate change. The threat is ubiquitous and inexorable, it is occur- ring now, accelerating much more rapidly than we think, and the outcome is, to quote from his opening sentence, “worse, much worse, than you think” (p. 3). Our response to this horrific scenario has been and continues to be inadequate. One can read all man- ner of apocalyptic claims about climate change and less dire public warnings from the climate scientists themselves who fear to sound alarmist. These proc- lamations are always couched in sugary dribbles of “but there is still hope”. Most of this hope emanates from ignorance and from the belief that the calam- ity is far down the arrow of time and can be allevi- ated by technology. Wallace-Wells puts paid to this dream by noting the rapidity with which the current 1.1°C and causal doubling of atmospheric CO, have occurred. I worked it out; 90% of greenhouse gases have been accumulated since Bob Dylan was born, % 66 and 80% since he sang the “The Times They are a Changin”. In one baby boomer generation, this catas- trophe has occurred. This book starts with one stunning fact after another and never falters through 225 pages of text. It leaves one breathless and raw. In Section 1, “Cascades”, Wallace-Wells describes the cascad- ing effects of climate change, emphasizing the inter- connectedness of our earthly paradise. He reminds the reader of the five great mass extinctions and points out that four of them involved increases in greenhouse gases and warming temperatures of up to 5°C. But these events took thousands or even millions of years to develop, we have only had Dylan’s life- time. Mind you, we have known since the mid-19th century that the simple fact of adding certain gases to the atmosphere would warm the planet, but until recently the idea that we would release hundreds of millions of years of accumulated hothouse carbon in just one century never really sank in. Indeed, for 30% of the population, it is still fake news. Wallace-Wells is Al Gore on steroids and presents an endless mass of facts from every facet of our earthly sphere, and yet we still act as though climate change is something distant, to be fixed by recycling, getting a smaller car, becoming vegetarian, and all manner of single issue fixes that most people resist and resent. Why? Because we can’t easily grasp the big picture: the frailty of permanence, the cumulative impact of eight billion of us and our fossil fuel technologies. And thus, we are doing too little. Section II, “Elements of Chaos”, is a series of chapters on specific disastrous effects of climate change, such as heat, hunger, drowning, wildfire. I 2019 won't try to recount the terrors that Wallace-Wells chronicles, there are far too many. Suffice to say these make depressing and startling reading. Section III, “The Climate Kaleidoscope”, covers a range of issues: storytelling, capitalism, technology, the pol- itics of consumption, history after progress (a very in- teresting read), and ethics at the End of the World. All of this is provocative and suitably disturbing. Finally, section IV discusses the concept of the Anthropic Principle. I will try to paint the context Wallace-Wells presents: how fragile our civilization and ecosystems are, and how inevitable, drastic and long-term cli- mate change will be. Wallace-Wells lists several major misconceptions that we hold about climate change, “myths” that en- courage us to be blasé about the end of the world en- visioned by so many climate scientists. First, he says we believe the “fairy tale” that climate change is slow. It isn’t. By geological or even human timescales, It is advancing at terrifying speed. Check your news stor- ies of 2019; climate change is here. Second, many see climate change as a problem largely confined to the Arctic. Climate change is global of course. In 2019, Karachi recorded the hottest April of any city on earth ever, and in 2019 earth itself experienced the hottest June ever recorded. Third, many see cli- mate change as a problem for the natural world and some species, like Polar Bears, but not for human- ity. This misapprehension arises from our failure to see ourselves as part of nature, and a disruptive yet wholly dependent part at that. Fourth, many see cli- mate change as a matter of slowly rising sea levels relevant only to low-lying coastal areas and remote tropical atolls. A fifth pernicious myth is that burn- ing fossil fuels is a necessary price we pay to foster “economic growth and good paying jobs”. These ne- cessary benefits pay for themselves by creating the technologies needed to repair the problems caused by the resultant climate change, using, for example, car- bon capture, cold fusion, daring experiments spray- ing other gases into the atmosphere or lead powder into the oceans, or building giant reflective umbrel- las. In other words, more of the same hubris that got us here. Throughout the book Wallace-Wells drops ‘fact bombs’ that support my view that we are insane. Example, bitcoin (p. 33) consumes more electricity than is produced by all the solar panels on our planet. The same bitcoin produces as much CO, annually as one million transatlantic flights (p. 179), and nearly 2% of the global total CO, emissions. We deeply embrace the belief that progress is built into our civilization and society. There will be more and better food, time-saving devices, medical life ex- tending treatments, poverty eliminated, and endless entertainment and travel opportunities. This myth is Book REVIEWS 67 readily embraced by the wealthy fractions of soci- ety and the well-off eagerly endorse claims, such as those of the biologist and prolific writer Steven Pin- ker (Pinker 2018), that every measure shows hu- man progress, that wealth, the economy, longev- ity, health, human rights, etc., are improving and that the Cassandras have been repeatedly proven wrong. Malthus, Ehrlich, the Club of Rome, even David Attenborough can hang their heads in shame for doubting the sharp upward thrust of history. We tend to forget that for 95% of human history progress was, to say the least, minimal, that population growth was almost invisible (see Hardin 1995), and economic growth was not a concept, much less a reality. Only in the last 5% of our time on earth have we seen nota- ble changes from the 200 000 years of hunter gather- ers and even that period of “progress” has regularly been blighted by setbacks from wars, diseases, and various genocides and pogroms. So-called progress has been largely over the past 200 years a product of the massive consumption of fossil fuels. A major re- sult has been the staggering increase to eight billion large, warm blooded, consumptive apes. A large por- tion still wallows in poverty and can’t afford to attend a Raptor’s game much less a holiday in space. The glorious irony of climate change is, of course, that we are entirely to blame and that the fabulous wealth and sumptuous lifestyles we have created are the exact reasons we are careening to calam- ity. Several times Wallace-Wells highlights this 1m- portant point. For example, on p. 53, “The graphs that show so much recent progress in the developing world (i.e., decline in poverty and hunger, improve- ment in life expectancy, education gender relations, and more) are, practically speaking, the same graphs that trace the dramatic rise in carbon emissions that has brought the planet to the brink of overall catastro- phe”. Everyone reading this review was/is/will be a major creative participant in climate change. Climate change is occurring now, will acceler- ate in the future, will endure for thousands of mil- lennia, and is entirely a product of the very recent past. Our impact on climate will last not until our grandchildren die off but for millions of years. We need more books like this one to slap us hard, to not sugarcoat reality with false reassurances. If we con- tinue our current insanity, then our civilization will be a tiny blip of an afterthought in the eternity of cli- mate change. “Man is not clever enough to limit his greed to courses that will not destroy the ecosystem.” —Gregory Bateson, Mind and Nature (E.P. Dutton, 1968) Note: I have deliberately omitted the myriad ter- rifying scenarios recounted by Wallace-Wells. Read 68 THE CANADIAN FIELD-NATURALIST the book. Meanwhile, I include Corn and Yalkin (2019), an examination of the great physical and men- tal toll on those scientists conducting research on cli- mate change. Literature Cited Corn, D., and D. Yalkin. 2019. It’s the end of the world as they know it. The distinct burden of being a climate scientist. July 8 2019. Accessed 28 August 2019. https:// www.motherjones.com/environment/2019/07/weight- BOTANY Vol. 133 of-the-world-climate-change-scientist-grief/. Hardin, G. 1995. Living within Limits: Ecology, Econo- mics, and Population Taboos. Oxford University Press, Oxford, United Kingdom. Pinker, S. 2018. Enlightenment Now: the Case for Reason, Science, Humanism, and Progress. Viking, New York, New York, USA. RON BROOKS University of Guelph, ON, Canada Michigan Ferns & Lycophytes: A Guide to Species of the Great Lakes Region By D.D. Palmer. 2018. University Michigan Press. 381 pages, 29.95 USD, Paper. The state of Michigan, USA, enjoys exceptionally rich floristic coverage and is ad- mirably served both by the Field Manual of Michigan Flora by E.G. Voss and A.A. Reznicek (University of Mi- chigan Press, 2012) and the excellent Michigan Flora Online (https://michiganflora -net/home.aspx). The Field Manual is widely seen to serve more than ‘just’ a lo- cal state flora function and to also provide a region- al (Great Lakes) perspective. For decades, how- ever, a curious gap in the state coverage has been with pteridophytes. Although a fern treatment was provided earlier by C. Billington’s Ferns of Michigan (Cranbrook Institute of Science, 1952), and the University of Michigan was home base for American fern guru W.H. (Herb) Wagner for much of the time thereafter, no modern fern treatment existed. The current volume corrects that omission. The book begins with a variety of standard intro- ductory elements for a flora, including a brief sum- mary of fern investigations in the state, a discussion of what makes pteridophytes ‘tick’, and a review of the abundance and distribution (including habitats) of pteridophytes in the state. A map showing the land- scape diversity and/or major vegetation zones of Michigan would have been helpful here for under- standing local distributions, especially for out-of- state readers, but the text does satisfy our basic needs in that regard. Similarly, it would have been useful to have a brief discussion of what makes Michigan’s fern flora special on a regional or even continental scale. We eventually get some of this with discus- sion of endemics, but there are several broad biogeo- graphic and evolutionary themes well represented in MICHIGAN Ferns &-Lycophytes A GUIDE TO SPECIES OF THE GREAT LAKES REGION the Michigan pteridophyte flora that also could have been profitably discussed here. It is quickly evident that Michigan Ferns & Lyco- phytes provides an admirable introduction and re- view of the distribution and identification of pterido- phytes in that state. There are excellent photographic and line drawing illustrations of key identification features, only slightly hampered by the absence of scale bars. The /soetes photo montage (p. 290), for ex- ample, is particularly effective for this tricky group. Individual treatments provide effective, clearly ex- pressed technical descriptions for the taxon in ques- tion with an emphasis on identification. The com- parative feature tables provided for most complex groups such as Botrychium, Dryopteris, Equisetum, Lycopodiella, and Woodsia, are very helpful. The identification keys for each genus are sound and are not overly laden with technical jargon. Distributional information seems to be quite up-to-date and accur- ate, although the unreferenced report of Cystopteris tennesseensis being (disjunct) in northern Ontario (p. 101) is news to us. In some cases, we suspect treatments may be over-simplified. For challenging members of Lyco- podiaceae, for example, it would be great to believe Great Lakes taxa are as straightforward to identify as they are presented to be in Michigan Ferns & Lyco- phytes. More than 30 years of wrestling with them on this side of the border suggests they are often other- wise! Michigan Ferns & Lycophytes prominently claims a secondary objective, professing to share the same regional scope as that of the Field Manual of Michi- gan Flora. Yes, most Great Lakes pteridophyte taxa are found in Michigan, but that is equally true for New York, Ontario, Ohio, etc. To truly be a regional guide, however, also requires that a local treatment explicitly reflect the regional context. Michigan Ferns & Lycophytes falls short in this, particularly regarding 2019 Canadian input. At least seven species are listed (pp. 11 and 238) as occurring in the Great Lakes portion of adjacent Ontario, Minnesota, and Wisconsin but not in Michigan, without further discussion. Another species—Isoetes tuckermanii A. Braun—and at least seven more hybrids known in Ontario from within this region are not even mentioned. The discussion of Dryopteris hybridization (pp. 150-153) omits reference to any of the regionally— indeed, globally—significant cytological research on this genus undertaken by Ontario’s Donald M. Brit- ton. Similarly, the discussion of Botrychium (s. 1.) di- versity also makes no mention of how sites along the Ontario shore of Lake Superior were critical to the taxonomic discoveries and innovations of the Univer- sity of Michigan team studying this group. Simply put, Wagner and Britton made the largest contributions of anyone to our understanding of the pteridophytes of the Great Lakes. Accordingly, the ab- sence of even a single citation from Britton’s volu- minous Great Lakes-relevant literature—not even W.J. Cody and D.M. Britton’s 1989 Ferns and Fern Allies of Canada (Agriculture Canada)—is_ sur- prising, even within just a Michigan ferns context. Together, these various omissions present a signifi- cant credibility problem for Michigan Ferns & Lyco- phytes’ claim to offer a regional perspective on Great Lakes pteridophytes. There are 108 species treated in Michigan Ferns & Lycophytes (121 full treatments including a selection of some additional subspecies, varieties, or hybrids). Taxa within some genera are treated in considerable detail while others receive more basic consideration. The Equisetum treatment, for example, employs 42 pages of text for the treatment of 13 taxa. This in- cludes species-comparable treatments for four sterile hybrids because they are “quite common and often form large clones” (p. 51). The considerably more ecologically, genetically, and biogeographically sig- nificant Dryopteris genus, however, is addressed in Book REVIEWS 69 only 27 pages treating 12 taxa. This treatment in- cludes stand-alone discussions of the two hybrids considered to be most common in the state. Another 16 hybrid combinations are listed as occurring in Michigan but without any supporting documenta- tion or references. Why stop there? Readers should at least have been directed to some pertinent references from Britton’s Ontario literature on Dryopteris hy- brids and/or to James Montgomery’s excellent 1982 North American treatment (Fiddlehead Forum 9: 23— 30). The paucity of supporting references is a prob- lem throughout, in fact, with the References section of the book having a surprisingly low total of fewer than 50 citations. Etymology is discussed for each taxon that re- ceives a stand-alone treatment. There is no harm in that because the origin of names has some popular in- terest. When these cultural/biographical discussions use large amounts of text space that could otherwise be profitably applied to the core identification object- ive of the book, however, they become counter-pro- ductive. The excessively long, biography-like etymo- logical discussion for Huperzia xjosephbeitelii A. Haines (p. 318), for example, is twice the length of the remaining text available for the technical description of this difficult taxon. At its core, Michigan Ferns & Lycophytes pre- sents a valuable tool for the identification of pterido- phytes in Michigan and substantially fills a long-stand- ing need. It also is an asset for the understanding of pteridophyte diversity in a geographically wider area as well. Out-of-state (especially Canadian) readers, however, will need arange of supplementary literature in order to gain the appropriate regional perspective. DANIEL F. BRUNTON Ottawa, ON, Canada MICHAEL J. OLDHAM Peterborough, ON, Canada 70 THE CANADIAN FIELD-NATURALIST Vol. 133 Flora of Florida Volume 6 (Dicotyledons, Convolvulaceae through Paulowniaceae) By R.P. Wunderlin, B.F. Hansen, and A.R. Franck. 2019. University Press of Florida. 372 pages, 70.00 USD, Cloth. The monumental Flora of | Florida project 1s now over half completed. With the publication in 2000 of Vol- ume | and a flurry of addi- tional publication activity in recent years (for reviews of earlier Flora of Florida volumes see The Canadian Field-Naturalist 130: 248— 249, 2016 [Volumes 2 and 3]; 131: 375, 2017 [Volume 4]; and 132: 68, 2018 [Volume 5]), there is but one more volume necessary to achieve complete cover- age of the dicot taxa. The final three volumes treating the monocot species are then to be published. The au- thors’ goal of having all 10 volumes in print by 2020 may not be achieved; putting out four volumes in less than a year and a half seems unlikely from both a pro- duction and marketing perspective. The good news, however, is that their ambitious completion objective may not be far off the mark. A total of 470 species in 19 vascular plant families are treated here. That number of taxa addressed is in- creased by the description of additional subspecies, varieties, and/or named hybrids within particular Species accounts. With the completion of Volume 6 some 2375 species have been described. All told, 62% of the 3834 species identified in Volume 1 as occur- ring or to have once occurred outside of cultivation in the state, have now been described. The names of a substantial number of excluded species that were re- ported in error or unconfirmed are also enumerated. As within those parts of Flora of Florida that pre- ceded Volume 6, effective species identification keys updated from Wunderlin’s Guide to the Vascular Plants of Florida (University of Florida Press, 1998) are placed immediately after each genus description. Al- phabetically arranged species treatments follow, each employing up-to-date nomenclature and commencing with the detailed and comprehensive compilation of synonyms that is a trademark of Flora of Florida. No compilation in the present volume, however, approaches the astonishing 75 synonyms listed in Volume 3 (pp. 220-223) for Crataegus michauxii Persoon! The physical characteristics of each taxon are de- scribed in the text (and keys) with precise but not overly technical terminology. The text is presented in a small but easily readable type on good quality paper within a hard cover binding. That readability is particularly important because the text is un-illus- trated. This is understandable in an already volumin- i Florida ous text but is nonetheless unfortunate, particularly for non-local readers who are likely less familiar with the Florida flora. Readers are encouraged to con- sult the online Atlas of Florida Plants (http://florida. plantatlas.usf.edu) for photos of most taxa, however, and for more detailed range information than is in the brief statements provided here. Habitat and (especially) other life history con- siderations are described sparsely, this presumably also a reflection of space limitations and cost control. That is also unfortunate for readers ‘from away’ who could benefit from comparing ecological character- istics of Floridian plants with those of their own lo- cal populations. As with the other treatments Volume 6 addresses many species that extend northward into southern Canada. That range limit seems to be quite accurately reflected for the most part. The copy examined for this review was weakly bound, the spine panel separating from the side boards after minimal use. This is contrary to my ex- perience with the firmly bound copies examined for reviews of earlier volumes, however, and likely repre- sents an infrequent aberration. A considerable number of taxa in Volume 6 are in families such as Convolvulaceae, Solanaceae, and (especially) Acanthaceae which typically do not con- tribute substantially to floristic diversity in north- ern areas of this continent. Some families with more prominent northern diversity are well represented however, including Plantaginaceae, Lamiaceae, and Lentibulariaceae. Floridian representation of the in- sectivorous genus of bladderworts (Utricularia, Lenti- bulariaceae) for instance, includes four species which occur in eastern Canada. The description of Floridian populations of taxa such as the Eastern Purple Blad- derwort (U. purpurea Walter) suggest striking differ- ences may exist with the Canadian populations of this species—and perhaps with others as well. The Flora of Florida is a grand undertaking that is providing a valuable floristic tool applicable far be- yond the limits of that state. Were Volume 6 a stand- alone analysis, however, I would not recommend it as a high priority acquisition for northern botanists be- cause of its lesser representation of north-related taxa than seen in earlier contributions. Just the same, how- ever, It warrants serious consideration for northern li- braries as a significant component of what will un- doubtedly be a classic of North American botanical literature almost immediately upon its completion. DANIEL F. BRUNTON Ottawa, ON, Canada 2019 Book REVIEWS Al Identification of Trees and Shrubs in Winter using Buds and Twigs By Bernd Schulz. 2018. Royal Botanic Gardens, Kew, distributed by University of Chicago Press. 368 pages, 45.00 GBP, 80.00 USD, Cloth or E-book. Originally published in German as: Geholzbestimmung im Winter: mit Knospen und Zweigen, 2013. As a Canadian naturalist, how will you be spending the upcoming winter? Per- haps a Christmas Bird Count shortly after the landscape transforms into snow and ice, and then as frigid weather lingers toward a seemingly relentless polar vortex, may- be an occasional excursion such as hiking the trails in snowshoes and perhaps a nocturnal oe Hien by the local dam? Many nature lovers by then be- come relatively dormant, and begin to dream about early spring arrival of migrating birds or of Skunk Cabbage bursting through a melting snowpack, only a month or two away. How about hunting for trees and shrubs? Woody plants are always available for study, and are perhaps best observed during the winter months when dis- tinctive twig features are fully developed and easily visible without being obscured by deciduous leaves (in contrast to herbaceous plants which may be with- ered and hidden under snow; e.g., Levine 1995). Besides being a rewarding and easily accessible win- ter activity from cultivated yards to remote wilder- ness, real contemporary conservation questions are awaiting resolution such as occurrences of popula- tions for rare/threatened species: relevant examples from this writer’s current work are establishing the locations of Red Spruce populations in the green- belt of Ottawa, and documenting the presence of new Rock Elm populations in Quebec along the land border of eastern Ontario and Quebec (Vaudreuil- Soulanges; FloraQuebeca 2009). Spotting trees and shrubs is frequently best done in winter, coincident- ally when there may be little else obvious for the nat- uralist to do outdoors. But how does one confirm that plant species of interest have really been found? The ‘fingerprint’ of woody plant species identi- fication is their twigs. All features of woody plants, from growth form to bark, leaves, flowers, and fruit, are useful to observe, but the only identification trait reliably present and virtually invariant among indi- vidual plants of a given species, at any age from sap- ling to centuries-old giant, is the twigs. Trees and shrubs can nearly always be accurately determined to genus via mature twig features (leaf scars, buds, etc.), and usually to species with a trained eye. Bernd Schulz’s /dentification of Trees and Shrubs Identification of Trees and Shrubs in Winter using Buds and Twigs Bernd Schulz in Winter using Buds and Twigs is an excellent re- source to learn the key features for differentiat- ing woody plants from family to genus to species. The book is at its essence a thoroughly modern text- book sorely needed for a curriculum which, one may state with little exaggeration, is no longer actively taught. The book has useful and accessible chapters regarding the history of twig-identification botany, twig structure and terminology, and identification keys, but is at its core a comprehensive species-by- species treatment of the trees and shrubs found in central Europe with detailed descriptions and hun- dreds of technically sound and attractive colour illustrations. A number of high-quality twig iden- tification books relevant to eastern North America were published from early to mid 20th century (e.g., Blakeslee and Jarvis 1911; Trelease 1918; Harlow 1941; Graves 1952; Core and Ammons 1958; Petrides 1958; Symonds 1958, 1963). These classic books are still very much useful and worth studying but suf- fer from being stuck in an historical era before the current “globalization” of cultivated plants, and the corresponding emergence of widespread naturaliza- tion and invasive species as dominant ecological fea- tures. Little on the subject was published from the 1960s until the 2010s. Schulz’s book is therefore a major step forward toward modernization for the subject, with comprehensive descriptions and illus- trations showing the native, cultivated, and natur- alized plants of central Europe in the 21st century; there is much species overlap with the woody plants of eastern North America in modern times. The clas- sic 20th-century books, in contrast, often give at most a light treatment of non-native plants, some of which have only arrived and become dominant in the wild in the last few decades. It is worth noting that the subject matter of Schulz’s book is not just relevant to winter: “winter” twig features (mature buds, etc.) are indeed present from late summer until spring, approximately two thirds of the year from August to April. Despite being an authoritative and high-qual- ity publication, a few features are worth considering which may be drawbacks to some of the intended audience. The book is large and heavy, with a typ- ical ‘textbook form’ factor and so is not a portable handbook to be easily tossed in one’s backpack. The book was written from a central European perspec- tive, and so is a comprehensive treatment of the na- tive, naturalized, and commonly cultivated trees 72 THE CANADIAN FIELD-NATURALIST and shrubs of that region specifically. While there is much overlap of naturalized and commonly culti- vated woody plants of Europe and North America, as noted above, some woody plants of Canada are not in- cluded. Evergreen trees and shrubs are not treated, so one cannot use this book to learn their identification traits: e.g., spruces versus firs or Sheep-laurel ver- sus Bog-laurel. This book only considers twigs, and despite doing this with unprecedented breadth and depth, does not treat other aspects of woody plants such as growth form, leaves, etc. which many of the classic books cover in addition to twigs. Overall, Schulz’s book is highly recommended to anyone interested in temperate zone woody plants and their conservation, and is available at reasonable prices (~50.00—70.00 CAD) from a variety of sellers. Literature Cited Blakeslee, A.F., and C.D. Jarvis. 1911. New England trees in winter. Storrs Agricultural Experiment Station Bul- letin 69: 307-576. https://doi.org/10.5962/bh1.title.32385 Core, E.L., and N.P. Ammons. 1958. Woody Plants in Winter: a Manual of Common Trees and Shrubs in Winter in the Northeastern United States and South- eastern Canada. Boxwood Press, Pacific Grove, Cali- Vol. 133 fornia, USA. FloraQuebeca. 2009. Plantes rares du Québec meridional. Publications Québec, Quebec, Canada. Graves, A.H. 1952. Illustrated Guide to Trees and Shrubs: a Handbook of the Woody Plants of the Northeastern United States and Adjacent Canada. Self-Published, Wallingford, Connecticut, USA. Harlow, W.M. 1941. Twig Key to the Deciduous Woody Plants of Eastern North America. Self-Published, Syra- cuse, New York, USA. Levine, C. 1995. A Guide to Wildflowers in Winter: Herba- ceous Plants of Northeastern North America. Yale Uni- versity Press, New Haven, Connecticut, USA. Petrides, G.A. 1958. A Field Guide to Trees and Shrubs: Field Marks of all Trees, Shrubs, and Woody Vines that Grow Wild in the Northeastern and North-central United States and in Southeastern and South-central Canada. Houghton Mifflin, Boston, Massachusetts, USA. Symonds, G.W.D. 1958. The Tree Identification Book. William MorrowandCo. Inc., New York, New York, USA. Symonds, G.W.D. 1963. The Shrub Identification Book. Wil- liam Morrow and Co. Inc., New York, New York, USA. Trelease, W. 1918. Winter Botany. Self-Published, Urbana, Illinois, USA. OweEN CLARKIN Ottawa, ON, Canada 2019 ENTOMOLOGY Book REVIEWS 73 Field Guide to the Flower Flies of Northeastern North America By Jeffrey H. Skevington, Michelle M. Locke, Andrew D. Young, Kevin Moran, William J. Crins, and Stephen A. Marshall. 2019. Princeton University Press. 512 pages, 3000 images, and 414 maps, 27.95 USD, Flexibound Paper. This is a beautiful book, big enough to include mul- tiple photographs of all the known, and until recently, a few unknown flower flies, yet small enough to be car- ried into the field in a larg- ish pocket or small satchel. The general introduc- tion describes the book’s layout and how to use it most efficiently. Techniques on observing and trapping these flies and how to rec- ord your data are also included. Identification poin- ters, including a reference to an online Key to the Genera of Nearctic Syrphidae (Miranda et al. 2013), give the reader a good start to determining the spe- cies. The book ends with a thorough glossary and several very useful diagrams illustrating much used anatomy. Prior to the species accounts is a two-page spread illustrating the differences among the four subfam- ilies of flower flies. Having never attempted to key out one of these flies to species before, I used this two- page spread, largely with success. One then goes to the colour-coded section of the book for the appro- priate subfamily to find their insect. And herein lies the one issue I have with this guide. Because there is no further key to identify the species, the reader may have to flip through quite a number of pages be- fore finding their fly. The online Key to the Genera of Nearctic Syrphidae, co-authored by four of the six authors of this book as well as two others, should be downloaded and used in conjunction with the guide PRINCETON FIELO GUIDES to identify your specimen to species. The species accounts are well done. There are two per double-page spread, with text on the left and im- ages on the right, allowing for rapid flipping while you're trying to find your fly. There is always a dor- sal view and lateral view, sometimes more than one if the species is dimorphic. Other salient features, such as wing venation, facial structure, or leg characteris- tics are also shown, as needed, and nicely magnified. All the photographs are at least adequate, but most are crisp, often beautiful shots. The text includes a size range, but the silhouette of the fly presented with each species gives a more rapid indication of size. An interesting series of icons that I’ve not seen before tells the reader if the insect can typically be identified by the unaided eye (sur- prisingly many), a hand lens or, ultimately, if a micro- scope will be needed. A map showing records, and usually a range estimate as well, does not restrict it- self to the geographic scope of this book, but includes all of North America. The text includes flight times, abundance, and identification tips; I couldn’t think of anything else that a field guide should have. Overall, this is a very nice book that, in conjunc- tion with the online key, will do its job quite well. Literature Cited Miranda, G.F.G., A.D. Young, M.M. Locke, S.A. Mar- shall, J.H. Skevington, and F.C. Thompson. 2013. Key to the Genera of Nearctic Syrphidae. Canadian Journal of Arthropod Identification 23: 23 August 2013. https://doi.org/10.3752/cjai.2013.23 RANDY LAUFF Department of Biology, St. Francis Xavier University, Antigonish, NS, Canada 74 THE CANADIAN FIELD-NATURALIST ORNITHOLOGY The Handbook of Bird Families Vol. 133 By Jonathan Elphick. 2018. Firefly Books and The Trustees of the Natural History Museum, London. 416 pages, 35.00 CAD, Paper. As most people with more than a slight interest in birds THE HANDBOOUOE understand, the taxonomy Mier ESS of Aves is at present in flux. S\DER KE RISO RCN of modern environmental thinking. Although each chapter is largely its own piece, I found some portions of each to overlap at times in terms of content, often referring to other chapters within the book. This does not detract from the work as a whole, however, but illustrates how a number of distinct but interweaving concepts coalesce to form the modern concept of the environment. I found this book to largely avoid tech- nical language, with references and technical notes appended into a considerable Notes section at the end of the book. This makes the book more access- ible to the non-specialized reader but allows one to dive further into the material referenced if desired. While each chapter is a summary of historical con- text, the authors add to its value and interpretation by providing meaningful original commentary and an- alysis throughout the book. Should you give it a read? Absolutely. Do not let the “history” in the title dissuade you from giving this book a try if you are not a history buff: this book is intended for a broad audience. The authors have followed their own suggestion by writing this work as a means of “framing a problem or concept as a nar- rative or story” (p. 178). I felt like much of the writing is presented in almost a ‘story-like’ narrative. In some parts, I almost felt as if I were listening to the authors discuss the content of the book over coffee. Overall, I found The Environment: A History of the Idea to be an accessible, enjoyable, and very informative read: I learned something new at the turn of nearly every page. Also, this work contains many references to other books (in addition to so much more) and as a result has introduced me to a number of titles which are now on my future reading list. I highly recom- mend The Environment: A History of the Idea to any- one with an interest in any aspect of environmental study. This would likely include most readers of The Canadian Field-Naturalist. Literature Cited Vogt, W. 1948. Road to Survival. Kessinger Publishing, LLC, Whitefish, Montana, USA. SEAN M. HARTZELL Bloomsburg University of Pennsylvania, Bloomsburg, PA, USA 2019 Book REVIEWS Ty. The Great Himalayan National Park: The Struggle to Save the Western Himalayas By Sanjeeva Pandey and Anthony J. Gaston. 2019. Niyogi Books. 364 pages, 284 colour photos, and 15 maps, 54.00 CAD, Cloth. The Himalayas: surely this is one of the most evocative names in world geography, for who has not seen inspir- ing images of the world’s grandest peaks and heard epic tales of scaling their summits? This book is set in the Himalayas but to ap- preciate its significance you eam must lower your eyes a few degrees, focussing not on the snow-shrouded crags but on the green slopes below, on the meadows and forests. The biological richness of the Himalayas lies at these elevations, not in the awe-inspiring realm of rock, ice, and snow. The mountains’ richness rests on three axes: first and most conspicuously, a ver- tical dimension, driven by the steep climate gradi- ent tied to elevation that generates profoundly differ- ent habitats for a diverse suite of species. Secondly, there is a sharp north-south axis because the height of the Himalayan range makes it a barrier between two major biogeographic realms. Thus, you can have Himalayan Brown Bears (Ursus arctos isabellinus) and Ibex (Capra sibirica sakeen) of the Palearctic realm living just over the ridge from Asiatic Black Bears (Ursus tibetanus) and Himalayan Serow (Capricornis thar) of the Oriental realm. Finally, there is an important west to east increase in rainfall. All three of these considerations make the area se- lected for the Great Himalayan National Park a geo- graphic crossroads meriting special protection, es- pecially given the threat of fragmenting ecological continuity. Imagine a species inhabiting forests on the south face of the Himalayas at elevations be- tween 2000-3000 m; their geographic range is es- sentially a long narrow ribbon, easily severed by de- forestation. Indeed, throughout the Himalayas forests at these intermediate elevations are markedly under- represented in protected area systems. These per- Spectives are introduced in the book’s opening chap- ter and prominent throughout, along with the other main rationale for siting a park here: the area’s low levels of human settlement and concomitant disturb- ance from logging and livestock grazing. The next chapter, “Trekking”, describes in lovely prose and photos what a visit to the park is like and it has inspired me to try to organize a return. (Full disclosure: I participated in two of the early surveys [spring and fall of 1980] organized by Tony Gaston that became a foundation for park creation but have A UNESCO WORLD HERITA oo sete As a wre sg THE GREAT NATIONAL PARK ‘The Scruggle to'Save the Western Himalayas SANJEEVA_PANDEY. ANTHONY | GASTON not been back since.) Unfortunately, as a ‘details per- son’ I found it frustrating not to be able to find most of the spots described or photographed on the maps; even the trekking routes are not mapped. In “Development of GHNP”, the book returns to the 1980s and the creation of the park. Some of this chapter is down in the weeds of the requisite steps that are specific to India’s governance but of wider interest is how park development was advanced with the local population. Thoughtful park design miti- gated some potential conflicts, 1.e., by delineating a core area that contained just three villages (120 in- habitants) and then along the western boundary cre- ating a 230 km? Ecodevelopment Zone that held 160 villages with 14 000 people. Traditional rights to col- lect herbs for personal use became a problem when this morphed into commercial sales on which a thou- sand households were dependent, and inevitably these people became very agitated when this com- merce was terminated. The key to recovery came in the formation of 95 Women’s Saving and Credit Groups (a large number to accommodate both dif- ferent villages and different social strata) that earned income from vermicomposting and other activities. Further insights into park-people relations are cov- ered in “People and the GHNP”, which has a pot- pourri of sections including religious traditions (sacred trees and groves), education (from 1989 to 2002 the only school in the park was in a cave), species targetted by herbalists, and the creation of a support organization, Friends of GHNP, with the Western Tragopan (7ragopan melanocephalus) as its mascot. The Friends group took the lead in applying for UNESCO World Heritage Site status for the park and adjoining protected areas, collectively an area of 2854 km7?, thus garnering formal recognition of the global significance of this place. The next three chapters focus on the park’s nat- ural history, the first a seasonal chronicle beautifully illustrated with flower photos, followed by two over- views of the park’s birds and mammals with spe- cial attention on charismatic species like the pheas- ants and carnivores. In the final chapter, “Future of Biodiversity in the Western Himalayas”, the auth- ors ruminate on the on-going process of manag- ing the park in a way that will further the interests of the local people while maintaining its unique bio- logical heritage. This challenge will unfold in the face of ever-changing threats, such as tourism pressures from the millions of Indians who seek cool, pleasant playgrounds to escape Delhi and other cities. 78 THE CANADIAN FIELD-NATURALIST In summary, this book shares many attributes— such as wonderful photographs and writing—with other volumes that have been written to celebrate the natural wonders of the world’s special places. It dif- fers from most analogous books in the depth with which it tells the story of all the hard work that under- lies protecting such places—from foundational sci- ence to pure politics. I particularly enjoyed the distil- lation of this that is captured nicely in the Foreword and Afterword, written by Gaston and Pandey re- spectively, in which they give personal accounts of ZOOLOGY Return of the Wolf: Conflict and Coexistence Vol. 133 the decades of work they have devoted to this unique place. All in all, it is a tale that should be of inter- est to anyone concerned with protected area creation and management, especially in places where the live- lihoods of local people are directly and tightly tied to natural resources. MaLco”M L. HUNTER, JR. Department of Wildlife, Fisheries, and Conservation Biology, University of Maine, Orono, ME, USA By Paula Wild. 2018. Douglas and McIntyre. 272 pages, 32.95 CAD, 29.95 USD, Cloth. Return of the Wolf by Paula Wild was an easy, enjoy- able read about the recov- ery and return of wolves (Canis spp.) throughout the world, focussing much of her time in her home coun- try of Canada studying F Gray Wolves (Canis lupus). The purpose of the book is to give the real picture of wolves, as neither saint nor sinner nor good versus bad, but rather just another animal (albeit a predator that often conflicts with humans where the two are sympatric) trying to survive in an increasingly hu- man-dominated landscape. Wild provides an histor- ical summary of wolves in both the Old World as well as the New World, given that they were once common throughout North America and Eurasia. Chapters 2 and 3 describe how bounties and organized hunts had drastically reduced their numbers worldwide by the 1900s. However, the wolf is now recovering from near extermination in many areas, especially parts of the United States’ lower 48 states, and is becom- ing more common as evidenced by their recovery in the Yellowstone area (p. 149) and upper mid-west. Throughout the book, Wild expresses amazement at the dualities of hatred and love that she encountered when ascribing emotions about wolves in her mis- sion to set the record straight by letting us appreci- ate the real animal. I have over 50 books on wolves— not even counting the 25+ I have on their close cousin the Coyote, Canis latrans—and | feel like this book is an appropriate summary of all those books. It pro- vides much of the historical background of, say, Barry Lopez’s 1978 Of Wolves and Men (Scribner, 2004), but also discusses modern happenings such RETURN of the WOLF Paula Wild as Yellowstone wolf recovery (see Way 2017) and marine-food-eating coastal wolves discussed in Ian McAllister’s book The Last Wild Wolves: Ghost of the Great Bear Rainforest (University of California Press, 2007). Curiously, the title of Wild’s book is strangely similar to two other books I own, including The Return of the Wolf (NorthWord Press, 1999) by Steve Grooms, which presents the wolf’s comeback in Canada and the United States, albeit 25 years ago now, and The Return of the Wolf: Reflections on the Future of Wolves in the Northeast (University Press of New England, 2000), edited by John Elder, that discusses the implications and potential of wolves re- turning to the northeast United States. Return of the Wolf is a mixture of natural hist- ory, native peoples’ stories, and conversations with scientists and conservationists. We learn how soci- ety’s attitudes affect the population dynamics, be- haviour, and conservation of wolves on the modern landscape, a setting where more and more people ap- preciate having nature around even if it challenges us both financially and safety-wise. Wild notes that the fate of wolves remains uncertain and she ques- tions how humans will adapt to wolves. She is opti- mistic that we will, noting that “I want to hear the wolves but I don’t want them to come too close. For their safety, not mine” (p. 241). Accordingly, the first Appendix item is a unique 2.5 page “Wolf Safety Checklist” (p. 243-245), one that you might think is more in tune with living in bear country. However, Wild spends much of the second half of her book dis- cussing ‘the myth’ that wolves are not dangerous and documents that healthy (1.e., not rabid) wolves are in- creasingly confronting, and sometimes even killing, humans in North America (Chapter 9). She also de- scribes some first-hand accounts of highly habituated wolves living on Vancouver Island, British Columbia (Chapter 10). In fact, the ecology of those animals 2019 living in human-dominated areas reminds me of the wolf’s smaller cousin, Coyote (e.g., Way 2014), in many respects. Of course, it is important to keep in mind that even with increasing boldness of some wolves, the chance of one harming us is still astro- nomically small compared to potential dangers from our everyday activities. Given my interest in studying Eastern Coyotes/ coywolves (Way 2014), I was fascinated with Wild’s discussion of this animal (pp. 91-96) and her deci- sion—due to their unique genetic background—to call them coywolves. While describing the rapid evo- lution of the coywolf, Wild also discusses the other lesser known wolves, Eastern Wolf (Canis lycaon) and Red Wolf (Canis rufus), which are possibly the same species living on opposite ends of their native eastern North American range. Wild circles back to Eastern Wolf a few times when also discussing recent aggressive encounters people have had with wolves, some of those with Eastern Wolves in Algonquin Provincial Park. Overall, this is an easy-to-read, well researched, timely book. While perhaps not having the exciting Book REVIEWS 79 flair of a book written by a biologist(s) in their study area, it provides a great up-to-date account of the hap- penings of wolves worldwide, with a North American focus. Whether you are new to the world of wolves, or a veteran, I recommend adding this book to your li- brary. The nice 16-page colour plate section as well as many black and white photos adds greatly to the read. Hopefully, it will provide food for thought and create compassion for a creature that has been maligned for far too long. Literature Cited Way, J.G. 2014. Suburban Howls: Tracking the Eastern Coyote in Urban Massachusetts. Revised Edition (edited and e-book). Dog Ear Publishing, Indianapolis, Indiana, USA. Way, J.G. 2017. [Book Review] American Wolf: A True Story of Survival and Obsession in the West. Canadian Field-Naturalist 131: 375-376. https://doi.org/10.22621/ cfn.v13114.2091 JONATHAN (JON) WAY Eastern Coyote/Coywolf Research, Osterville, MA, USA 80 THE CANADIAN FIELD-NATURALIST NEw TITLES Prepared by Barry Cottam Vol. 133 Please note: Only books marked ¢ or * have been received from publishers. All other titles are listed as books of potential interest to subscribers. Please send notice of new books—or copies for review—to the Book Review Editor. tAvailable for review *Assigned Currency Codes: CAD Canadian Dollars, AUD Australian Dollars, USD United States Dollars, EUR Euros, GBP British Pound. BIOLOGY Fires of Life: Endothermy in Birds and Mammals. By Barry Gordon Lovegrove. Foreword by Roger S. Seymour. 2019. Yale University Press. 384 pages, 40.00 USD, Cloth. The Evolutionary Biology of Species. Oxford Series in Ecology and Evolution. By Timothy G. Barra- clough. 2019. Oxford University Press. 288 pages, 90.00 USD, Cloth, 45.95 USD, Paper. Also available as an E-book. The Tangled Tree: A Radical New History of Life. By David Quammen. 2018. Simon & Schuster. 30.00 USD, Cloth, 18.00 USD, Paper, 11.99 USD, E-book. The Ethnobotany of Eden: Rethinking the Jungle Medicine Narrative. By Robert A. Voeks. 2018. Uni- versity of Chicago Press. 328 pages, 45.00 USD, Cloth, 10.00—45.00 USD, E-book. Evolution in the Dark: Darwin’s Loss Without Se- lection. By Horst Wilkens and Ulrike Strecker. 2017. Springer International Publishing. 226 pages, 179.99 USD, Cloth or Paper, 139.00 USD, E-book. BOTANY Carnivorous Plants. By Dan Torre. 2019. Reaktion Books. 240 pages, 27.00 USD, Cloth. Essentials of Developmental Plant Anatomy. By Taylor A. Steeves and Vipen K. Sawhney. 2017. Ox- ford University Press. 184 pages, 74.00 USD, Cloth. Also available as an E-book. Flora Unveiled: The Discovery and Denial of Sex in Plants. By Lincoln Taiz and Lee Taiz. 2017. Oxford University Press. 520 pages, 76.95 USD, Cloth. Great Trees of New Brunswick. Second Edition. By David Palmer and Tracy Glynn. Photographs by Arielle DeMerchant. 2019. Goose Lane Editions. 264 pages, 27.95 CAD, Paper. Primrose. By Elizabeth Lawson. 2019. Reaktion Books. 256 pages, 27.00 USD, Cloth. CLIMATE CHANGE Climate Change and Rocky Mountain Ecosystems. Advances in Global Change Research. By Jessica E. Halofsky and David L. Peterson. 2019. Springer In- ternational Publishing. 253 pages, 149.99 USD, Cloth or Paper, 109.00 USD, E-book. Effects of Climate Change on Birds. Second Edi- tion. Edited by Peter O. Dunn and Anders Pape Mol- ler. 2019. Oxford University Press. 288 pages, 100.00 USD, Cloth. Also available as an E-book. In Search of the Canary Tree: The Story of a Sci- entist, a Cypress, and a Changing World. By Lau- ren E. Oakes. 2018. Basic Books. 288 pages, 16.99 USD / 21.99 CAD, Paper. *No One is Too Small to Make a Difference. By Greta Thunberg. 2019. Penguin Books. 80 pages, 9.99 CAD, Paper. CONSERVATION & ECOLOGY Ecological Forecasting. By Michael C. Dietze. 2017. Princeton University Press. 288 pages, 65.00 USD, Cloth. Also available as an E-book. Effective Conservation Science: Data Not Dogma. Edited by Peter Kareiva, Michelle Marvier, and Brian Silliman. 2017. Oxford University Press. 384 pages, 100.00 CAD, Cloth, 49.95 CAD, Paper. Also avail- able as an E-book. Freshwater Ecology and Conservation: Approaches and Techniques. Edited by Jocelyne Hughes. 2019. Oxford University Press. 464 pages, 90.00 USD, Cloth, 45.95 USD, Paper. Also available as an E-book. Hierarchy: Perspectives for Ecological Complex- ity. By T.F.H. Allen and Thomas B. Starr. 2017. Uni- versity of Chicago Press. 352 pages, 125.00 USD, Cloth, 47.50 USD, Paper, 10.00—47.50 USD, E-book. Time in Ecology: A Theoretical Framework. By Eric Post. 2019. Princeton University Press. 248 pages, 105.00 USD, Cloth, 40.00 USD, Paper. Also available as an E-book. The North American Model of Wildlife Conser- vation. Edited by Shane P. Mahoney and Valerius Geist. 2019. Johns Hopkins University Press. 184 pages, 74.95 USD, Cloth or E-Book. 2019 Renewable Energy and Wildlife Conservation. Edi- ted by Christopher E. Moorman, Steven M. Grodsky, and Susan P. Rupp. 2019. Johns Hopkins University Press. 280 pages, 74.95 USD, Cloth or E-book. Human-Wildlife Interactions: Turning Conflict into Coexistence. Conservation Biology Series. Edi- ted by Beatrice Frank, Jenny A. Glikman, and Silvio Marchini. 2019. Cambridge University Press. 476 pages, 89.99 USD, Cloth, 44.99 USD, Paper, 36.00 USD, E-book. The Terrestrial Protected Areas of Madagascar: Their History, Description, and Biota. Edited by Steven M. Goodman, Marie Jeanne Raherilalao, and Sébastien Wohlhauser. 2019. University of Chicago Press. Three volumes, 1716 pages, 180.00 USD, Cloth. ENTOMOLOGY Papillons de Nuit et Chenilles du Québec et des Maritimes. Guides Nature Quintin. By Michel Le- boeuf and Stephane Le Tirant. 2018. Editions Michel Quintin. 336 pages, 34,95 CAD, Couverture souple, 19,99 CAD, PDF. The Dark Side of the Hive: The Evolution of the Imperfect Honeybee. By Robin Moritz and Robin Crewe. 2018. Oxford University Press. 208 pages, 80.00 CAD, Cloth. Also available as an E-book. Saproxylic Insects: Diversity, Ecology and Conser- vation. Zoological Monographs, Vol. 1. Edited by Michael D. Ulyshen. 2018. Springer International Publishing. 1003 pages, 279.00 USD, Cloth or Paper, 219.00 USD, E-book. Ecomorphology of Cyclorrhaphan Larvae (Dip- tera). Zoological Monographs, Vol. 4. By Graham Rotheray. 2019. Springer International Publishing. 295 pages, 179.00 USD, Cloth, 139.00 USD, E-book. Buzz, Sting, Bite: Why We Need Insects. By Anne Sverdrup-Thygeson. 2019. Simon & Schuster. 256 pages, 26.00 USD, Cloth, 13.99 USD, E-book. Beneficial Insects. By David V. Alford. 2019. CRC Press. 384 pages and 385 colour illustrations, 99.95 USD, Cloth. Also available as an E-book. Cerambycidae of the World: Biology and Pest Management. Contemporary Topics in Entomology Series. Edited by Qiao Wang. 2017. CRC Press. 628 pages, 175.00 USD, Cloth. Also available as an E-book. +The Mosquito: A Human History of Our Dead- liest Predator. By Timothy C. Winegard. 2019. Allen Lane Canada. 496 pages, 32.95 CAD, Cloth. The Last Butterflies: A Scientist’s Quest to Save a Rare and Vanishing Creature. By Nick Haddad. 2019. Princeton University Press. 256 pages, 24.95 New TITLES 81 USD, Cloth. Also available as an E-book. Raising Butterflies in the Garden. By Brenda Dziedzic. 2019. Firefly Books. 336 pages, 24.95 CAD, Paper. Honey From the Earth: Beekeeping and Honey Hunting on Six Continents. By Eric Tourneret and Sylla de Saint Pierre. Edited by Dr. Leo Sharashkin. 2018. Deep Snow Press. 352 pages and 300+ colour photos, 64.95 USD, Cloth. Published in French in 2015. HERPETOLOGY The Rise of Reptiles: 320 Million Years of Evolu- tion. By Hans-Dieter Sues. 2019. Johns Hopkins Uni- versity Press. 400 pages and 356 illustrations, 84.95 USD, Cloth or E-book. The Field Herping Guide: Finding Amphibians and Reptiles in the Wild. Wormsloe Foundation Nature Book Series. By Mike Pingleton and Joshua Holbrook. 2019. University of Georgia Press. 264 pages, 26.95 USD, Paper. Reptiles of Costa Rica: A Field Guide. Zona Tropi- cal Publications Series. By Twan Leenders. 2019. Cornell University Press. 640 pages, 35.00 USD, Paper. ICHTHYOLOGY The Future of Bluefin Tunas: Ecology, Fisheries Management, and Conservation. Edited by Barbara A. Block. 2019. Johns Hopkins University Press. 360 pages, 124.95 USD, Cloth or E-book. Deep-Sea Fishes: Biology, Diversity, Ecology and Fisheries. By Imants G. Priede. 2017. Cambridge University Press. 492 pages 89.99 USD, Cloth. ORNITHOLOGY Birds in Winter: Surviving the Most Challenging Season. By Roger F. Pasquier. Illustrated by Margaret La Farge. 2019. Princeton University Press. 304 pages, 29.95 USD, Cloth or E-book. Birds of Prey: Biology and Conservation in the XXI Century. By Jose Hernan Sarasola and Juan Manuel Grande. 2018. Springer International Pub- lishing. 530 pages, 219.99 USD, Cloth or Paper, 169.00 USD, E-book. *The Flying Zoo: Birds, Parasites, and the World They Share. By Michael Stock. 2019. The University of Alberta Press. 296 pages, 29.99 CAD, Paper or PDF. Pelican. By Barbara Allen. 2019. Reaktion Books. 208 pages and 80 colour plates, 19.95 USD, Paper. 82 THE CANADIAN FIELD-NATURALIST Thinking Like a Parrot: Perspectives from the Wild. By Alan B. Bond and Judy Diamond. 2019. University of Chicago Press. 184 pages, 35.00 USD, Cloth, 10.00—35.00 USD, E-book. Urban Aviary: A Modern Guide to City Birds. By Stephen Moss. Illustrated by Marc Martin. 2019. White Lion Publishing. 160 pages, 26.00 CAD, Cloth. ZOOLOGY A Bat’s End. The Christmas Island Pipistrelle and Extinction in Australia. By John Woinarski. 2018. CSIRO Publishing. 266 pages, 59.99 AUD, Paper. Also available as an E-book. Field Guide to the Bats of the Amazon. By A. Lopez- Baucells, R. Rocha, P. Bobrowiec, E. Bernard, J. Pal- meirim, and C.F. J. Meyer. 2018. Pelagic Publishing. 175 pages, 48.46 CAD, Paper. Wildlife on Roads: A Handbook. By Kari E. Gun- sen and Frederick W. Schueler. 2019. Eco-Kare Inter- national. 232 pages and 140 colour photographs, 29.95 CAD, Paper. The North Atlantic Right Whale: Disappearing Giants. By Scott Kraus and Kenneth Mallory. 2019. Fitzhenry & Whiteside. 120 pages, 24.95 USD, Paper. +A Field Guide to Marine Life of the Outer Coasts of the Salish Sea and Beyond. By Rick M. Harbo. 2019. Harbour Publishing. 8-fold pamphlet, 7.95 CAD, Paper. tA Field Guide to Marine Life of the Protected Waters of the Salish Sea. By Rick M. Harbo. 2019. Harbour Publishing. 8-fold pamphlet, 7.95 CAD, Paper. Handbook of the Mammals of the World - Volume 9. Bats. Edited by Don E. Wilson and Russell A. Mittermeier. Illustrations by Toni Llobet. 2019. Lynx Edicions in association with Conservation Interna- tional and IUCN. 160.00 EUR, Cloth. +Mammal Tracks and Sign: A Guide to North American Species. Second Edition. By Mark Elbroch with contributions by Casey McFarland. 2019. Globe Pequot/Stackpole Books. 680 pages, 49.95 USD, Paper, 47.50 USD, E-book. A Manual of the Mammalia: An Homage to Law- lor’s “Handbook to the Orders and Families of Living Mammals”. By Douglas A. Kelt and James L. Patton. 2019. 544 pages, 60.00 USD, Cloth or E-book. Canids of the World: Wolves, Wild Dogs, Foxes, Jackals, Coyotes, and Their Relatives. Princeton Field Guides. 2018. Princeton University Press. 336 pages, 29.95 USD, Paper. *The Rise of Wolf 8: Witnessing the Triumph of Vol. 133 Yellowstone’s Underdog. By Rick McIntyre. Fore- word by Robert Redford. 2019. Greystone Books. 304 pages, 34.95 CAD, Cloth. Carnivores of the World. Second Edition. Prince- ton Field Guides. By Luke Hunter. Illustrated by Priscilla Barrett. 2019. Princeton University Press. 256 pages, 29.95 USD, Paper. The Truth About Animals: Stoned Sloths, Love- lorn Hippos, and Other Tales from the Wild Side of Wildlife. By Lucy Cooke. 2018. Perseus Books. 352 pages, 28.00 USD / 36.50 CAD, Cloth, 16.99 USD / 22.49 CAD, Paper, 11.99 USD / 14.99 CAD, E-book. Cats in Australia: Companion and Killer. By John Woinarski, Sarah Legge, and Chris Dickman. 2019. CSIRO Publishing. 344 pages, 59.99 AUS, Paper. Marine Mammals: Adaptations for an Aquatic Life. By Randall W. Davis. 2019. Springer Interna- tional Publishing. 192 pages, 99.99 USD, Cloth. Saving the Dammed: Why We Need Beaver- Modified Ecosystems. By Ellen Wohl. 2019. Oxford University Press. 176 pages, 35.00 USD, Cloth. Also available as an E-book. Guide to Venomous and Medically Important In- vertebrates. By David Bowles, James Swaby, and Harold Harlan. 2018. CSIRO Publishing. 240 pages, 59.99 AUD, Paper. Also available as an E-book. OTHER *To Speak for the Trees: My Life’s Journey from Ancient Celtic Wisdom to a Healing Vision of the Forest. By Diana Beresford-Kroeger. 2019. Random House Canada. 289 pages, 32.00 CAD, Paper. Wild Sea: A History of the Southern Ocean. By Joy McCann. 2019. University of Chicago Press. 256 pages, 28.00 USD, Cloth, 18.00 USD, E-book. Good Enough: The Tolerance for Mediocrity in Nature and Society. By Daniel S. Milo. 2019. Har- vard UP. 320 pages, 28.95 USD, Cloth. An Alfred Russel Wallace Companion. Edited by Charles H. Smith, James Costa, and David A. Collard. 2019. University of Chicago Press. 416 pages, 60.00 USD, Cloth or E-book. Community-Based Control of Invasive Species. Edited by Paul Martin, Theodore Alter, Don Hine, and Tanya Howard. 2019. CSIRO Publishing. 288 pages, 99.99 AUS, Cloth. Wildlife and Wind Farms - Conflicts and Solutions. Volume 4, Offshore: Monitoring and Mitigation. Edited by Martin Perrow. 2019. Pelagic Publishing. 340 pages, 80.43 CAD, Paper. Collecting Experiments: Making Big Data Biol- 2019 ogy. By Bruno J. Strasser. 2019. University of Chicago Press. 392 pages, 135 USD, Cloth, 45.00 USD, Paper. Also available as an E-book. Wilding: Returning Nature to Our Farm. By Isa- bella Tree. Introduction by Eric Schlosser. 2019. Pica- dor. 392 pages, 19.95 USD, Paper. Also available as an E-book. Naturally Curious. A Photographic Field Guide and Month-By-Month Journey Through the Fields, Woods, and Marshes of New England. Second Edition. By Mary Holland. 2019. Trafalgar Square Books. 496 pages, 32.95 USD, Paper. Wading Right In: Discovering the Nature of Wet- lands. By Catherine Owen Koning and Sharon M. Ashworth. 2019. University of Chicago Press. 264 pages, 30.00 USD, Paper, 90.00 USD, Cloth, 10.00— 35.00 USD, E-book. Inside Science: Stories from the Field in Human and Animal Science. By Robert E. Kohler. 2019. University of Chicago Press. 264 pages, 35.00 USD, Cloth, 10.00—35.00 USD, E-book. Quantitative Analyses in Wildlife Science. Edited by Leonard A. Brennan, Andrew N. Tri, and Bruce G. Marcot. 2019. Johns Hopkins University Press. 344 pages, 74.95 USD, Cloth or E-book. Ocean Recovery: A Sustainable Future for Global Fisheries? By Ray Hilborn and Ulrike Hilborn. 2019. Oxford University Press. 208 pages, 37.95 USD, Cloth. Also available as an E-book. Treed: Walking in Canada’s Urban Forests. By Ariel Gordon. 2019. Wolsak & Wynn. 296 pages, New TITLES 83 20.00 CAD, Paper. Rates of Evolution: A Quantitative Synthesis. By Philip D. Gingerich. 2019. Cambridge University Press. 396 pages, 84.99 USD, Cloth. Collecting Evolution: The Galapagos Expedition that Vindicated Darwin. By Matthew J. James. 2017. Oxford University Press. 304 pages, 34.95 USD, Cloth. Also available as an E-book. Forest Landscape Restoration: Integrated Ap- proaches to Support Effective Implementation. The Earthscan Forest Library. Edited by Stephanie Mansourian and John Parrotta. 2018. CRC Press. 266 pages, 150.00 USD, Cloth. Also available as an E-book. The Routledge Handbook of the Polar Regions. Routledge International Handbooks. Edited by Mark Nuttall, Torben R. Christensen, and Martin J. Siegert. 2018. Routledge, Taylor & Francis Group. 530 pages, 260.00 USD, Cloth. Also available as an E-book. Everyday Creatures. A Naturalist on the Surpris- ing Beauty of Ordinary Life in Wild Places. By G.J. Kenagy. 2018. Dockside Sailing Press. 220 pages, 15.95 USD, Paper, 203 pages, 5.99 USD, E-book. For the Birds: American Ornithologist Margaret Morse Nice. By Marilyn Bailey Ogilvie. 2018. Uni- versity of Oklahoma Press. 320 pages, 39.95 USD, Cloth, 34.95 USD, E-book. Slime: How Algae Created Us, Plague Us, and Just Might Save Us. By Ruth Kassinger. 2019. Houghton Mifflin Harcourt. 320 pages, 26.00 USD, Cloth, 16.99 USD, Paper. The Canadian Field-Naturalist News and Comment Upcoming Meetings and Workshops Canadian Herpetological Society Annual Conference The Canadian Herpetological Society Annual Con- Quebec. Registration is currently open. More infor- ference to be held 20—23 September 2019 at the mation is available at http://canadianherpetology.ca/ Redpath Museum in McGill University, Montreal, conf/index.html#2019. Wildlife Society and American Fisheries Society Joint Annual Conference The first Wildlife Society and American Fisheries vention Center, Reno, Nevada. Registration is cur- Society Joint Annual Conference to be held 29 rently open. More information is available at https:// September-—3 October 2019 at the Reno-Sparks Con- — afstws2019.org/. Student Conference on Conservation Science-New York The 10th annual Student Conference on Conservation More information is available at https://www.amnh. Science-New York to be held 2—4 October 2019 at org/research/center-for-biodiversity-conservation/ the American Museum of Natural History, New convening-and-connecting/student-conference-on- York, New York. Registration is currently open. conservation-science-new-york-sccs-ny. Association of Field Ornithologists and the Wilson Ornithological Society Joint Meeting The joint meeting of the Association of Field Orni- is currently open. More information is available at thologists and the Wilson Ornithological Society, http://www.cvent.com/events/2019-afo-wos-joint- hosted by the New Jersey Audubon’s Cape May Bird meeting/event-summary-fc644f2542184eba9fe3b- Observatory, to be held 27-30 October 2019 at the 1d37928e0fd.aspx. Grand Hotel, Cape May, New Jersey. Registration Entomological Society of Ontario The annual general meeting of the Entomology Soci- More information is available at https://www.entsoc ety of Ontario to be held 1-3 November 2019 at the _ ont.ca/. Bark Lake Conference Centre, Haliburton, Ontario. Entomology 2019 The annual meeting of the Entomological Society mation is available at https://www.entsoc.org/events/ of America to be held 17-20 November 2019 at the annual-meeting. America’s Center Convention Complex, St. Louis, Missouri. Registration is currently open. More infor- 84 2019 NEWS AND COMMENT 85 James Fletcher Award for The Canadian Field-Naturalist Volume 132 The James Fletcher Award is awarded to the au- thors of the best paper published in a volume of The Canadian Field-Naturalist (CFN), and first started with Volume 130. The award is in honour of James Fletcher, founder of the Ottawa Field-Naturalists’ Club (OFNC) and the first editor of CFN’s earli- est iteration, Zransactions of the Ottawa Field- Naturalists’ Club. A subcommittee of the OFNC Pub- lications Committee sifted through all papers in Volume 132 of CFN, and came up with a list of the top five papers. From these top five, the committee selected the top paper, which for this volume was ac- tually a tie between two papers, both of which re- ceive the James Fletcher Award. The awards for Vol- ume 132 of CFN go to: Joseph J. Bowden, Kyle M. Knysh, Gergin A. Bla- goev, Robb Bennett, Mark A. Arsenault, Caleb F. Harding, Robert W. Harding, and Rosemary Curley. The spiders of Prince Edward Island: experts and citi- zen scientists collaborate for faunistics. Canadian Field- Naturalist 132(4): 330-349. https://doi.org/10.22621/cfn. v13214.2017 — This paper presents the first comprehensive list of spider species on Prince Edward Island, in- creasing the known list from 44 to 198 species, which is a huge accomplishment. The paper also used a unique collaboration between ex- perts and citizen scientists. And to: Richard Troy McMullin, Katherine Drotos, David Ireland, and Hanna Dorval. Diversity and conser- vation status of lichens and allied fungi in the Greater Toronto Area: results from four years of the Ontario BioBlitz. Canadian Field-Naturalist 132(4): 394—406. https://doi.org/10.22621/cfn.v13214.1997 — This paper presents data on a concentrated ef- fort to collect observations of lichen species during the Ontario BioBlitz in the Greater Toronto Area over four years, and demon- strates the utility of rapid assessments for mon- itoring the diversity of lichens. This paper was based on a collaboration between experts and citizen scientists. Congratulations to Joseph Bowden and co-authors and to Troy McMullin and co-authors for writing these excellent papers. The runners up for this award are: Chris R.J. Hay, R. Greg Thorn, and Clinton R. Jacobs. Taxonomic survey of Agaricomycetes (Fung: Basidiomycota) in Ontario tallgrass prairies deter- mined by fruiting body and soil rDNA sampling. Canadian Field-Naturalist 132(4): 407—424. https:// doi.org/10.22621/cfn.v13214.2027 — This paper used genetic techniques and tradi- tional survey methods to describe the diversity of Agaricomycetes in Ontario tallgrass prairies. Sue Carstairs, Marc Dupuis-Desormeaux, and Christina M. Davy. Revisiting the hypothesis of sex-biased turtle road mortality. Canadian Field- Naturalist 132(3): 289-295. https://doi.org/10.22621/ cfn.v13213.1908 — This paper tested a long-standing hypothesis that female turtles are more at risk of road mor- tality than males using a unique dataset col- lected at the Kawartha Turtle Trauma Centre, and found evidence to refute this hypothesis: both sexes were just as likely to be struck on roadways in three of four turtle species. Rebekah Neufeld, Cary Hamel, and Chris Friesen. Manitoba’s endangered alvars: an initial description of their extent and status. Canadian Field-Naturalist 132(3): 238-253. https://doi.org/10.22621/cfn.v13213. 1865 — This paper describes endangered alvars in Manitoba, describing their extent, plant com- munities, and land use. Congratulations to these finalists. We would also like to show our appreciation to all authors who chose to share their interesting and valuable field- based studies with the readers of Volume 132 of The Canadian Field-Naturalist. WILLIAM D. HALLIDAY AND JEFFERY M. SAARELA OFNC Publications Committee Diana Beresford-Kroeger: a new book, a life’s work Classical botanist, medical biochemist, and revo- lutionizer of how we look at forests has—by the time you read this—launched her seventh book. The event is scheduled for 24 September 2019 in Toronto, and we will have a review of it in our next issue of The Canadian Field-Naturalist. In the meantime, here is a brief note on Diana, the book, and selected as- pects of her work. I spoke with her by telephone— she in Ottawa, I in rural Prince Edward Island—on 13 August 2019 and this account is based in part on our conversation. The title of her new book, 7o Speak for the Trees: 86 THE CANADIAN FIELD-NATURALIST My Life’s Journey from Ancient Celtic Wisdom to a Healing Vision of the Forest (Penguin Random House, 2019), captures the essence of her work, which is a unique combination of almost-lost beliefs in the forest-enabled human connection with nature and the eruption of recent research into the wholeness of land, sky, and waters through the forests. The ca- pacity to hold in the mind—and body and spirit—the knowledge of both modern science and indigenous systems built on that wholeness is rare. Fortunately for the planet, it is becoming less rare, and Diana’s lifework is a major stream contributing to this ‘new renaissance’. She’s particularly encouraged by the range of engagement, from community efforts to improve the health of cities and neighbourhoods to the activism of youth world-wide to combat climate change. Diana informed me of the 2017 passing by the New Zealand Parliament of an Act that gives the Whanganui River the status and rights of a person, a precedent-setting major step forward that settled years of litigation and provided funding for improv- ing the health of the river. Diana’s previous books celebrate and urge pres- ervation of the world’s forests, especially the boreal forest ringing the north, the last, relatively intact great forest on Earth. The message is simple: the for- ests are key to the survival of life on the planet, in- cluding human life. And it isn’t just the capacity to survive but the quality of life in all its aspects. With their concept of “forest bathing”, the Japanese have been aware of this for over 1000 years; through her writings—including a chapter in the forthcoming International Handbook of Forest Therapy—and other media, Diana is helping bring this practice to North America. Her research in biochemistry has en- abled her both to understand the chemical communi- cations among trees and the impact of that communi- cation on land and waters and how that biochemistry can be harnessed for human health: mental, physical, and spiritual. She is also an accomplished medical re- searcher with some 300 articles to her name. Perhaps the most easily accessible summary of her work can be found in the inspiring documentary Call of the Forest: The Forgotten Wisdom of Trees, which can be viewed on the TVO website (https://www.tvo.org/ video/documentaries/call-of-the-forest-the-forgotten- wisdom-of-trees). In the making of this film, Diana travelled to various corners of the world—Tokyo and Hokkaido Island, Ireland’s Raheen Oak Wood, the redwoods of California, the Avatar Grove on Vancouver Island, British Columbia, and UNESCO world site Pimachiowin Aki on the eastern shore of Lake Winnipeg—to meet and share experiences with people engaged in restoration of forests and research demonstrating their value is highest and most essen- Vol. 133 tial when left intact. A second key theme of her work is science-based wisdom for living: her 2013 volume The Sweetness of a Simple Life: Tips for Healthier, Happier, and Kinder Living Gleaned from the Wisdom and Science of Nature (Random House Canada), contains 60 short chapters organized around “Health and Food”, “Home and Garden’, and “The Larger World”. The book concludes with a description of her Bioplan, a constant third theme in all her work. The foundation of the Bioplan is simply stated— everyone plants a tree a year for six years—and po- tentially very effective. She knows not everyone can or will do that, but the science is getting behind it; she cited “The global tree restoration potential” in Science (365: 76-79 https://doi.org/10.1126/science. aax0848), as a recent example. Diana consults with many schools and other institutions on how to fol- low her Bioplan. She also has her own forest and garden, a 65-hectare plot she calls Carriglaith near Merrickville, just outside Ottawa, where she and her husband, Christian Kroeger, work hard to preserve rare species of trees. She goes to phenomenal efforts to find these species. Awards have flowed in. In 2010, she was elected a Fellow by Wings WorldQuest, an international or- ganization “dedicated to recognizing and supporting visionary women” and the following year, the Utne Reader named her one of their 25 Visionaries for 2011. The Royal Canadian Geographical Society elected her to the College of Fellows in 2013 and named her one of 25 women explorers of Canada in 2016. Diana received an honorary doctorate from Carleton University on 11 June 2019. The citation notes that Call of the Forest “was nominated for the Rob Stewart Award for Best Science or Nature Documentary Program at the 2018 Canadian Screen Awards. Her peer reviewed work, Arboretum America: A Philosophy of the Forest, won the prestigious National Arbor Day Foundation Media Award for an exceptional educational work on trees and forests” (https://newsroom.carleton.ca/2019/diana-beresford- kroeger-receives-honorary-degree-from-carleton- university/). She has been profiled in many newspa- pers, the New York Times, Winnipeg Free Press, and Ottawa Citizen, to name only a few; interviewed on Baltimore NPR’s Marc Steiner Show, by the CBC, and other radio stations; in October 2018, Andrew Nikiforuk published a two-part article on her work in The Tyee, an independent, online, British Columbian news magazine. As well, Diana has written over 300 scientific and ‘popular’ accounts of her research for various journals and magazines. (See the Media sec- tion of http://calloftheforest.ca/about-diana/ for a par- tial list.) She gives guest lectures and masterclasses 2019 and is a science advisor for the Archangel Ancient Tree Archive, which works to preserve the Earth’s oldest trees through its Champion Tree Project (https://www.ancienttreearchive.org/). How she does all this while managing to live off-line is a wonder— she’s not sure how she does it either, but noted that not having the internet is certainly helpful in getting things done! A final comment: in addition to inspiring people all over the world, she was recently fictionalized as scientist Patricia Westerford in Richard Powers’ near- NEWS AND COMMENT 87 overwhelming novel about the life and times of trees, The Overstory, soon to be reviewed in The Canadian Field-Naturalist. Space won’t allow recounting her story of how she heard about this, but it became yet another means of getting the message out. Clearly, Diana Beresford-Kroeger’s work, like all works of nature, will live on in many forms. BARRY COTTAM Book Review Editor — The Canadian Field-Naturalist The Canadian Field-Naturalist Minutes of the 140" Annual Business Meeting (ABM) of the Ottawa Field-Naturalists’ Club, 8 January 2019 Place and time: Chairperson: Diane Lepage, President Neatby Building, Carling Avenue, Ottawa, Ontario, 7:00 pm Fifty-three attendees spent the first half-hour reviewing minutes of the previous ABM, the financial state- ments, Treasurer’s Report, and annual reports of Ottawa Field-Naturalists’ Club (OFNC) committees for 2017— 2018. The meeting was called to order at 7:30 pm. During the meeting, relevant documents were projected on a screen for the audience’s information. 1. Minutes of the Previous Annual Business Meeting (ABM) It was moved by Lynn Ovenden, seconded by Julia Cipriani, that the minutes of the 139" ABM be ac- cepted as distributed and published in The Canadian Field-Naturalist (CFN). Carried 2. Business Arising from the Minutes Nil. . Communications Relating to the Annual Business Meeting Nil. Treasurer’s Report by Ann Mackenzie Ann presented a simplified statement of OFNC’s major sources of revenue (membership, donations, interest) and the Club’s donations and expenses (net of associated revenues) from various club activities. Similar to last year, there was a significant shortfall as we continued to use the proceeds of Violetta Czasak’s bequest to further the objectives of the club. How long we can continue to run large deficits before the be- quest funds are depleted will depend on interest rates as well as our level of deficit spending. Subscription fees for The Canadian Field-Natu- ralist increased last year but that increased revenue is deferred until the coming year when the relevant issues are published. Ann also described some of the electronic technol- ogies and programs that are enabling her and other volunteers to process OFNC financial processes more efficiently, maybe even remotely. The Club increas- ingly makes payments by direct deposit instead of cheque, and accepts payment by direct bank transfer and credit card. In response to a question, members were told the budget for 2018-2019 was approved by the Board of Directors at its October 2018 meeting and attached 4. 88 to the minutes, available at http://ofnc.ca/wp-content/ uploads/2018/11/OF NC-Board-Minutes-2018-10-15. pdf. Moved by Ann MacKenzie and seconded by Henry Steger, that the financial statements be accepted as fair representation of the financial position of the Club as of September 30, 2018. Carried 5. Nomination of the Accounting Firm Moved by Ann MacKenzie and seconded by Jeff Skevington, that the accounting firm of Welch LLP be contracted to conduct a review of the OFNC’s ac- counts for the fiscal year ending September 30, 2019. Carried 6. Committee Annual Reports A few highlights from each report were presented by the chair or a representative of each committee. There was a request to make the reports available on the website. Moved by Lynn Ovenden and seconded by Ann MacKenzie, that the committee reports be accepted as distributed. Carried 7. Highlights from 2018 a) Christmas Bird Count (Bob Cermak) The 100" Ottawa-Gatineau Christmas Bird Count (CBC), a joint effort of the Ottawa Field-Naturalists’ Club (OFNC) and the Club des ornithologues de POutaouais (COO), was held 16 December 2018. The CBC, worldwide since the first in 1900, is the longest running citizen science survey in the world. Data col- lected by observers are vital for conservationists. It informs strategies to protect birds and their habitat, and helps identify environmental issues with impli- cations for people as well. b) Safe Wings Ottawa (Anouk Hoedeman) Anouk provided an update on Safe Wings’ pro- 2019 gress in promoting awareness of bird-building colli- sions and effective solutions. She discussed the im- portance of monitoring buildings and collecting data and specimens to convince building owners of the se- riousness of the problems, and the challenges volun- teers face due to widespread misconceptions about preventing collisions and the need to rescue stunned birds. She spoke of the program’s unexpected expan- sion into providing rescue and rehabilitation not only for window collision victims, but for birds requiring care for other reasons as well, due to the high demand for such services. c) Mer Bleue walk with new Canadians (Jakob Mueller) At the request of the Ottawa Community Immi- grant Services Organization, a nature walk at Mer Bleue was offered to about 30 new Canadians in Sep- tember. Several members with experience leading walks participated in the event and everyone enjoyed the outing. 8. Nominations for Board of Directors positions Fenja Brodo presented the slate of candidates no- minated to the Board of Directors for 2019: EXECUTIVE COMMITTEE Diane Lepage President Jakob Mueller 1 Vice President and Chair, Events Committee Elizabeth Moore Recording Secretary Ann MacKenzie Treasurer DIRECTORS Fenja Brodo Past President Robert Cermak Chair, Birds Committee Owen Clarkin Chair, Conservation Committee Representative, Fletcher Wildlife Committee Member-at-Large Edward Farnworth Catherine Hessian MINUTES OF THE 140™ ANNUAL BUSINESS MEETING 89 Anouk Hoedeman — Chair, Safe Wings Ottawa Diane Kitching Representative, Macoun Field Club Bev McBride Member-at-Large Gordon Robertson Chair, Education and Publicity Jeff Saarela Chair, Publications Henry Steger Chair, Membership Ken Young Chair, Finance Eleanor Zurbrigg Chair, Awards EX OFFICIO: Annie Bélair, Editor of Trail & Landscape Dwayne Lepitzki, Editor of The Canadian Field- Naturalist Moved by Fenja Brodo and seconded by Barry Cot- tam that this slate of nominees be accepted as mem- bers of the Board of Directors of the OFNC for 2019. Carried Fenja acknowledged Lynn Ovenden’s departure from the Board after serving as the Recording Secre- tary. She warmly welcomed Elizabeth Moore and Bev McBride to the Board. Committee chairs will be approved by the Board of Directors at the January 2019 meeting. 9. New Business and General Discussion Nil. 10. Presentation: Small Creatures, Big Impact Jakob Mueller described the seven salamander species that live in eastern Ontario and the important role of salamanders in the deciduous and mixed for- ests of eastern North America. 11. Adjournment Moved by Lynn Ovenden, seconded by Eleanor Zurbrigg, that the meeting be adjourned. Carried LYNN OVENDEN Recording Secretary 90 THE CANADIAN FIELD-NATURALIST Vol. 133 Annual Reports of OFNC Committees for October 2017— September 2018 Awards Committee The Awards Committee manages the process to annually recognize those Ottawa Field-Naturalists’ Club (OFNC) members and other qualified persons who, by virtue of their efforts and talents, are de- serving of special recognition. In late 2017, nomina- tions were received and evaluated (see awards crite- ria at https://ofne.ca/about-ofnc/awards), resulting in nominees for four awards being recommended to the Board of Directors for approval. Biographies were written for each award recipient for inclusion in the Club’s publications and posting on the website. The awards were presented at the annual Awards Night in April 2018. The recipients’ names, type of award and rationale for recognition follow below. ¢ Annie Bélair—Member of the Year for rede- signing Trail & Landscape as a communica- tion venue for local members. ¢ Julia Cipriani—George McGee Service Award for dedicated service on Events Committee and other areas of Club activity. ¢ David Seburn—Conservation Award for a Member for advocacy efforts influencing the decision to ban hunting of Snapping Turtles in Ontario. ¢ The Teachers of Regina Street Alternative School—Mary Stuart Education Award for con- necting their students with nature at Mud Lake. President Diane Lepage selected Greg Lutick and Adrienne Jex as the recipients of the 2017 President’s Prize for improving service of refreshments at month- ly meetings. The Awards Committee thanks Mark Brenchley for helping with awards certificate design and printing. Awards Committee: Irwin Brodo, Julia Cipriani, Christine Hanrahan, Karen McLachlan Hamilton ELEANOR ZURBRIGG, Chair Birds Committee Birds Committee (11 members, one ex officio, and one observer), Bird Records Sub-committee (11 mem- bers), and Bird Feeders Sub-committee (one mem- ber and five volunteers) coordinated OFNC bird re- lated activities and directed and encouraged interest in birds within and outside the OFNC area. Two committee members administered OFNC’s Facebook group (2161 members October 2018) which provides a place for club members and prospective members to discuss ideas and exchange information related to all aspects of natural history, club outings, and club initiatives. A committee member provided weekly provincial (Ontbirds) reports of OFNC area (Ottawa-Gatineau) bird sightings which with photos by local photogra- phers was also provided on the OFNC Facebook and the OFNC website. This same member authored a four-part, four issue, Trail & Landscape article “How to find 250 Bird Species in the OFNC Study Area in a Single Year”. Our committee participated in the Ottawa River Watershed Study and the creation of the Lac Des- chénes-Ottawa River Important Bird Area (IBA). We liaised with the Innis Point Bird Observatory and rec- ommended financial support for their Osprey web cam and bander-in-charge. We coordinated updates to the Department of National Defence’s (DND’s) Shirleys Bay causeway access list which currently contains 385 OFNC members. Birds Committee collaborated with other OFNC committees throughout the year. Our committee mem- bers led field trips and participated in Conservation Committee activities such as supporting the Friends of the Carp Hills with their survey and planning for the conservation and public use of Torbolton Ridge land owned by the City of Ottawa. We provided a significant role in OFNC’s efforts to introduce new Canadians to OFNC area natural history by partner- ing with the Ottawa Community Immigrant Services Organization to organize and help lead a field trip at the Mer Bleue boardwalk and area. Birds Committee and the Club des ornithologues de L’Outaouais organized the 2017 Christmas Bird Count with 133 field observers and 28 feeder watch- ers finding 60 bird species. The 2018 Seedathon “Big Day” event was held in early fall to raise funds for purchasing seeds for the five OFNC bird feeders. One hundred and thirty bird species were found and $790 (as of 1 August) was raised. Although the OFNC Falcon Watch is no longer re- quired because the Heron Road nest site is in a safe location, several members keep an informal watch on the nest. Four more Peregrines successfully fledged from this location this year. Bos CERMAK, Chair Conservation Committee A major accomplishment of the OFNC Conser- vation Committee in 2018 was successfully document- ing Red Spruce (Picea rubens) at many sites in eastern Ontario. This tree species is considered a key compo- nent of climax forests of eastern Canada and yet was apparently largely “missed” in the forest surveys of 2019 eastern Ontario to date. Members of the committee and other interested individuals collaborated to: * Confirm the presence of Red Spruce is much more abundant and widespread south of the Mer Bleue bog (centred at approximately Anderson Road/Leitrim Road/Highway 417) than was previously known. ¢ Confirm the presence of Red Spruce at several locations outside of the Ottawa district where it was apparently not previously documented: Hawkesbury, Alfred, near Alfred Bog, Rose Lake, and Vennachar Junction-Denbigh. ¢ This project was the topic of the October OFNC monthly meeting, and will be sum- marized in a forthcoming article currently in preparation. Records are being uploaded to iNaturalist as time permits. We anticipate that we will continue to work on this project at a more relaxed pace in 2019 to further resolve uncertainties regarding Red Spruce in Ontario. It was another busy year for OFNC-Conservation regarding public outreach and active conservation work in the great outdoors. Members of our commit- tee lent their expertise to lead numerous conservation events, including many guided public Nature tours, public lectures, bio-inventories, and attendance/con- tribution at conservation-related meetings. Our com- mittee members continued to enthusiastically doc- ument wildlife near Ottawa and further afield with thousands of observations entered into iNaturalist and other conservation databases. As in previous years, we met regularly to dis- cuss and plan actions regarding species of conser- vation concern (both threatened indigenous species, and emerging potentially invasive exotics), habitat conservation/restoration projects, and engaging both naturalists and the general public alike in conserving Canada’s natural wildlife heritage. We were pleased to collaborate on conservation work with a number of external organizations and several other OFNC com- mittees throughout the year. We were pleased to welcome Greg Lutick to the committee this year. OwEN J. CLARKIN, Chair Education and Publicity Committee Storyboards for display at eight sites around the Fletcher Wildlife Garden (FWG) continue to be de- veloped. We now have 24 bilingual stories, eight per season. They are now all laminated so that they can be reused for years. The previous ones on photopaper were unusable after one season. We held a second Open House at the Resource Centre in collaboration with Jane’s Walks. Several members hosted about 100 visitors with drinks, cook- ANNUAL REPORTS OF OFNC COMMITTEES 91 ies, and tours of the garden. This was approximately double from the previous year. This year we had two applications for sponsorship to the Youth Summit of Ontario Nature. Both candi- dates were judged worthy to attend and have done so. Lucy Patterson and Kathy Conlan were again judges at the annual Ottawa Regional Science Fair. They presented three OFNC awards ($100 each) and |-year club memberships to the winning students. We hosted numerous group tours of the FWG this year including three tours by home schoolers, seven tours by Scouts/Beavers/Brownies, and two school groups. We also developed seasonal Visual/Audio Scavenger Hunts for use with these groups and one for a tour of Mer Bleue for new immigrants. A flyer was also made for Petrie Island but was never used due to closing of its causeway until 2019. There were also several presentations at a seniors’ home, to the DND, and to the Canadian Parks and Wilderness Society (CPAWS). Bug Day was again a success this year with hun- dreds of attendees visiting our microscope tables to view insect specimens provide by Fenja Brodo. A callout for volunteers yielded 16 enthusiastic OFNC members who assisted with identifying insects and observing specimens under the microscopes. Finally, we had one new member confirmed by the Board, Alexandra Brett, with three more possible in the near future. Thanks are extended for the contribu- tions of Mark Brenchley who has stepped down from the committee. GORDON ROBERTSON, Chair Events Committee In 2018, the Events Committee (consisting of Julia Cipriani, Owen Clarkin, Hume Douglas, Bev McBride, Elizabeth Robson-Gordon, and Jakob Mueller) planned or organized field trips, workshops, and presentations for monthly meetings, as well as a Bioblitz-style spe- cies inventory. The committee is grateful for the help it received from leaders, volunteers, and experts too numerous to list. Nine of the club’s ten monthly meetings included presentations (the exception was the Annual Business Meeting). Topics ranged from faraway locations (Uganda, Iceland, and Mars), to conservation chal- lenges (freshwater ecosystems, cod stocks, turtles, and Red Spruce), ecology (the importance and role of night), and profiles of taxonomic groups (lichen). The December meeting had 135 people in atten- dance, the highest for a monthly meeting since the venue moved to the Neatby building several years ago. Of the 48 trips or workshops, primary focusses included birds or birding (19), botany (11), herpetol- ogy (three), insects (two), and mushrooms (one), with 92 THE CANADIAN FIELD-NATURALIST the remaining 12 trips devoted to general natural his- tory and miscellaneous topics. The annual Members’ Photography Night continued, but was postponed to April and held in a new venue due to some logistical challenges in January. In addition, the committee lent their expertise to the coordination of a species inven- tory for the Nature Conservancy of Canada’s prop- erty that forms part of the Kenauk Nature reserve north of Montebello, Quebec. Over two days in July, OFNC members and associated scientists recorded 454 species at the property and revealed opportuni- ties for further study. The Events Committee continues to face the pe- rennial yet pressing challenge of recruiting and devel- oping new leaders for field trips. If you have ideas for events or are willing to share your interests or exper- tise, please do not hesitate to get in touch. JAKOB MUELLER, Chair Finance Committee This report covers financial matters during fis- cal year 2018, which extended from 1 October 2017 through 30 September 2018. Many of these matters go directly to the Board of Directors for resolution. However, they are mentioned here to give OFNC members a sense of the financial issues that occur. The budget for FY2019 was prepared by the Finance Committee, based on input from the Board of Directors and analysis by the Finance Committee. It was presented to the Board in September 2018, and approved at the October 2018 Board meeting. The budget forecasts revenues of $146 000, expenses of $195 775, for a deficit of $49 775. During the 2018 fiscal year, the Board dealt with several financial issues that were not foreseen when the budget for FY2018 was prepared. These were: ¢ A request by the Innis Point Bird Observatory for $4500, to pay a Head Bander. This was ap- proved; ¢ A request by a local bird sanctuary to support its new educational programs. This was de- clined; ¢ A request by a group sponsoring a clean-up at Mud Lake, for $400 for a barbecue lunch for volunteers. This was declined. A financial issue which comes up from time to time is whether a particular type of volunteer work should be recognized by granting an honorarium. During FY2018, this arose at the Board in the context of Facebook administrators. The Board decided not to grant honoraria for this activity. The Treasurer continued her work to improve our systems for bookkeeping, donations, and payments. KEN YOUNG, Chair Vol. 133 Fletcher Wildlife Garden Volunteers The FWG had a productive season in spite of peri- ods of low rainfall and high temperatures. Volunteers working through the summer were joined by several corporate work groups, and students from Carleton University and University of Ottawa to carry out work throughout the property. Habitat preservation and re- habilitation took up the majority of our time, with good progress made on combating a growing list of invasive and noxious plants. Agriculture and Agri- food Canada (AAFC) was particularly helpful in this respect. In 2018, our volunteers put in over 5000 hours of time maintaining and improving the property. Our annual native plant sale was a great success. The sale is a source of revenue, but most important, it promotes the use of native plants in local gardens. In 2018, we also donated native plants to at least 10 school, community, church, and other demonstration pollinator gardens. Wildlife This summer saw an unexpected rise in the num- ber of Monarchs and other butterflies at the FWG. Several volunteers collected and reared eggs and cat- erpillars and released over 150 mature Monarchs. The release events were witnessed by many volun- teers and visitors to the FWG, creating opportuni- ties to chat about conservation issues. In conjunction with Wild Pollinator Partners, we also held two work- shops in April to make our highly successful Mason Bee boxes and another in fall to look at the pupae in the boxes (128) before storing them for winter. Jessica Forrest of the University of Ottawa gave a workshop on bees and was on hand at the latter workshop to identify wasp and bee pupae. Three Snapping Turtle nests were identified at the FWG this year, and we had help from the Canadian Wildlife Federation in rescu- ing eggs from a washed-out nest from the Arboretum. Pond work The south bank of the pond is now covered with a variety of native plants, much used by Giant and Black Swallowtail butterflies and Goldfinches. Two benches were installed along the trail to allow visitors obser- vation points. Erosion damage to the path around the pond was repaired early in the season as a safety mea- sure. In October, a contractor was hired to install a drainage trench to prevent future problems. We con- tinued to remove Flowering Rush from the pond. Backyard Garden The Back Yard Garden (BYG) continues to be a popular area, and the hard work of our Friday morn- ing volunteer group is much appreciated by visitors. Many photographers and artists visited the BYG to take advantage of the colourful variety of flowers and 2019 birds. Last fall, several trees were removed to main- tain favourable sun/shade conditions of the garden. The Gully The area between the west end of the pond and Prince of Wales Drive was a focus of much work this summer. Thanks to students from the Algonquin College horticulture program and the initiative and hard work of FWG volunteers, masses of Dog- strangling Vine and Comfrey were replaced with na- tive plants, such as Swamp Milkweed and Joe-Pye Weed. The transformation of this part of the FWG was inspiring to all involved and rewarding to those of us who saw the many Monarchs and other insects there this summer. The Ravine Intensive removal of invasive species on the north side of the ravine in 2017 was followed by planting of Bush Honeysuckle this summer. The addition of a bench overlooking the ravine has created a popular vantage point for photographers and birders. Tours/Visitors During this summer, the FWG provided tours to a number of youth groups. A series of questionnaires about plants, animals, birds, and insects that can be found at the FWG were used for the first time this summer by young visitors to look for as they walk along the trails. Storyboards have been placed around the property with seasonally appropriate photos to emphasize the diversity of flora and fauna at the FWG. Visitors to the Resource Centre now enjoy new stand up displays that include actual wildlife specimens. TED FARNWORTH, Committee Member Macoun Club Macoun Field Club activities take place almost ev- ery Saturday from September through June, except for public holidays. During 2017-2018, Committee members organized and supervised 17 indoor meet- ings, at which the children and young people (ages eight to 17) gathered for presentations on mainly zo- ological and ecological topics. Committee members CANADIAN USA Individual 385 (399) 7 Family 314 (316) 0 Student 23 (24) 0 Trail & Landscape 2 (2) 0 Honorary 24 (25) 0 Life 40 (41) 3 Other 24 (24) 0 Macoun Club If (19) TOTAL 827 (848) 10 ANNUAL REPORTS OF OFNC COMMITTEES 93 also planned and led 18 field trips, mostly to either the Club’s long-term nature-study area in Ottawa’s western greenbelt, or to Gerry Lee’s wild lands in the Pakenham Hills of Lanark County (from 1967-2002 known to Macoun Club members as Mary Stuart’s property). Part of one field-trip involved a joint outing with the OFNC, at Brewer Park Pond, and one special astronomy trip was held at night. Additionally, there was one overnight camping trip to Gerry Lee’s land. Committee and Macoun Club members hosted the annual nature quiz at the OFNC Awards Night. The Macoun Club quietly celebrated its 70th year. Committee members maintained an up-to-date, il- lustrated record of all events on the Club’s website, now hosted by the OFNC, and produced the monthly newsletter and issue No. 72 of the Club’s annual mag- azine, The Little Bear. RoserT E. LEE, Chair Membership Committee The distribution of Club membership for 2018 on 30 September 2018 is shown in the table below, with the corresponding numbers shown in brackets for 30 September 2017. “Other” represent mostly affiliate organizations that receive complimentary copies of the Club’s publications. There was a slight decrease in membership in 2017. Members within 50 km of Ottawa comprised 713 of the Canadian membership of 827. Families of children in the Macoun Club are given complimentary membership to encourage in- terest in the Club in the longer term. HENRY STEGER, Chair Publications Committee The Publications Committee manages publication of the Club’s scientific journal The Canadian Field- Naturalist (CFN), the Club’s regional publication Trail & Landscape, and Special Publications. Com- mittee members were Annie Beélair (Editor, Trail & Landscape), Dan Brunton, Carolyn Callaghan, Paul Catling, Barry Cottam (Book Review Editor, CFN), William Halliday (Online Journal Manager, CFN), OTHER TOTAL (6) 0 (0) 392 (405) (0) 1 (0) 315 (316) (0) 0 (0) 23 (24) (0) 0 (0) 2 (2) (0) 0 (1) 24 (26) (3) 1 (1) 44 (44) (0) 1 (1) 25 (25) 15 (19) (9) 3 (3) 840 (860) 94 THE CANADIAN FIELD-NATURALIST Karen McLachlan Hamilton, Diane Kitching, Dwayne Lepitzki (Editor-in-Chief, CFN), Amanda Martin (Assistant Editor, CFN), Jeff Saarela (Chair), David Seburn, Ken Young, and Eleanor Zurbrigg. Frank Pope resigned from the committee in December 2017, after many years of committee participation. Trail & Landscape Four issues of Trail & Landscape were published: 51(4) (October-December 2017), 52(1) (January— March 2018), 52(2) (April—June 2018) and 52(3) (July— September 2018). Feedback from Club members on the new, all-colour format of Trail & Landscape (launched with vol. 51(3)) continued to be extremely positive and encouraging. The Canadian Field-Naturalist Three issues of The Canadian Field-Naturalist were published: 131(2), 131(3), and 131(4). In 131(3) a new type of content was introduced in The Canadian Field-Naturalist. “Great Canadian Field-Naturalists” recognizes individuals who made significant contri- butions to our knowledge of the natural history of Canada. Criteria for recognition were established, and Great Canadian Field-Naturalist tributes were published for James Fletcher and John Macoun. Also this year, the James Fletcher Award was established. The award recognizes the best paper published in CFN ina particular volume. The first award, for CFN Volume 130 (2016), was announced in 131(3). Francis Cook, who served as Editor of CFN for more than 30 years, was appointed to the Order of Canada in 2018. Subscription fees for institutional subscribers were increased, based ona survey of fees for journals simi- lar to CFN, and the fact that our production costs have increased without subscription increase for a long time. Revised author guidelines for preparing manu- scripts for CFN were finalized and published on- line at http:/Awww.canadianfieldnaturalist.ca/public/ journals/1/CFNAuthorInstructions.pdf. The committee completed the consolidation pro- cess of back issues of The Canadian Field-Naturalist. A small team convened in spring 2018 and moved the back issues, which were stored in the Red Barn on the Farm during the autumn and winter of 2017-2018, to the Fletcher Interpretive Centre, and then sorted and organized the material. Given limited demand for hard copy back issues of the journal, only a small sub- set of the available back issues was retained; the ma- terial is now being stored by Annie Belair. The sur- plus material was recycled. Ottawa Field-Naturalists’ Club Research Grants 2018 was the fourth year of the Ottawa Field- Naturalists’ Club Research Grants program. Research grants support field-based research activities that re- Vol. 133 flect and promote the Club’s objectives within east- ern Ontario and/or western Quebec, focussed partic- ularly upon the Club’s study area. A total of $15 000 is available each year to fund research proposals. The application deadline was 15 January 2018. A subcom- mittee convened and chaired by Dr. Tony Gaston re- viewed all proposals and submitted funding rec- ommendations to the Board of Directors. A list of recipients of 2018 Research Grants was published in Trail & Landscape 52(3). JEFFERY M. SAARELA, Chair Safe Wings Committee 2017-2018 was Safe Wings Ottawa’s first year as a standing committee of the OFNC, after several years as a program of the Birds Committee. This commit- tee works to address the problem of bird-window col- lisions through research, education, and rescue. Between 1 October 2017 and 30 September 2018, Safe Wings volunteers: ¢ regularly monitored 75+ buildings during spring and fall migration ¢ documented +2300 window collisions ¢ rescued 493 live birds following window col- lisions * answered approximately 2500 phone calls ¢ admitted 610 birds with various types of inju- ries to our own facility * provided rescue assistance and transported hundreds of injured birds to Safe Wings and to the Wild Bird Care Centre. In addition, we produced a brochure on win- dow collisions, and completed a draft Ottawa Bird Strategy. Our outreach efforts led to the adoption of bird-friendly design guidelines by Public Services and Procurement Canada (PSPC), the National Capi- tal Commission, and the City of Ottawa (currently in development), as well as Carleton University and the University of Ottawa. Environment and Climate Change Canada also began working on several initia- tives to address collisions. In 2017-2018, Safe Wings volunteers’ workload expanded significantly due to several factors: public awareness of bird-building collisions continued to in- crease; the City of Ottawa began referring calls about all injured birds to Safe Wings; the Ottawa Valley Wild Bird Care Centre further reduced its own phone hours and increased its reliance on Safe Wings to pro- vide preliminary care to injured birds; and several high-profile, non-collision bird rescues drew media attention, namely the Great Horned Owl family on Petrie Island, a Wild Turkey in downtown Ottawa, the Bluesfest Killdeer (although we did not directly inter- vene), and a Northern Flicker found hung by its neck. Safe Wings also generated media coverage for 2019 its building monitoring program, its annual bird dis- play at City Hall, the National Art Centre’s not-bird- friendly renovation, the National Gallery of Canada’s failure to address collisions, PSPC’s efforts to retro- fit the C.D. Howe building, and Snowy Owls being hit by cars. ANOUK HOEDEMAN, Chair Treasurer’s Annual Report, 2018 The Financial statements for the year ended 30 Sep- tember 2018 have been prepared by our accounting firm, Welch LLP based on a review they conducted of our financial records. We do not get our books audited. There are no major changes to report from last year to this year because many of the trends remain. We are continuing to run deficits as we use the pro- ceeds of the Czasak bequest to further the objectives of the club. During the year we were able to provide a $10 000 donation towards the publication of The Birds of Nunavut by Anthony Gaston, continuing the OFNC’s long history of support for the natural his- tory of the Arctic. For the third year we gave $5000 to the Ottawa-Carleton District School Board to en- able more children to spend time at their Outdoor Education Centres. Support was also provided for a bird bander at Innis Point. Both the Invasive Plant Council (annual meeting) and the Entomological Society (Bug Day) also received support. In the interests of good governance your Board re- viewed the length of time we could run substantial deficits before the bequest funds were depleted. How long this takes will depend on interest rates as well as our level of deficit spending. An analysis of these two variables was carried out thanks to a tool developed by Catherine Hessian. If we continue with our cur- rent level of a $80 000 difference between our inter- est income and budget deficit, then we would be back to our pre-bequest investment level of $500 000 in 19 years. Using this tool the Board can more prudently consider any significant expenditures that might be proposed in the future. For example, if we donated $100 000 for the purchase of property then we would have used up the bequest in 17 years. Last year we reported that we would be closing several funds as part of streamlining of our finan- cial systems and record keeping. This will be the last year that the Seedathon Fund, Anne Hanes Memorial Fund, and the De Kiriline Lawrence Fund will be ANNUAL REPORTS OF OFNC COMMITTEES 95 shown in the statements. These funds have been net- ted to zero as they are incorporated into the General Fund of the club. The two Macoun funds have been merged into one. As a result of increased subscription fees for the CFN journal there was a significant increase in rev- enue from that source. Most of this was deferred un- til next year because the issues it relates to were not yet published. The apparent increased revenue show- ing this year reflects deferral rates between the last couple of years. We are increasingly using technologies and pro- grams to enhance the efficiency of our financial pro- cesses. This past year we started making payments us- ing direct deposits rather than cheques. Increasingly payments we receive are similarly direct bank trans- fers. We also did a trial to accept credit card pay- ments at the Fletcher Wildlife Garden plant sale. That worked so well we decided that we would promote it for 2019. This has been my second full year as Treasurer. I am pleased that we were able to provide the Board with the financial statements for their consideration at their December meeting prior to presenting them at the Annual Business Meeting in January. It is my in- tention to continue that practice into the future. In undertaking my tasks I have been greatly as- sisted by the club financial/administrative team of Ken Young (Chair Finance, CFN invoices, CFN-by- issue reports), Catherine Hessian (Investments, mail and bank deposits), Tanja Schueler (Paypal deposits), Bob Berquist (Donation Receipts), Eleanor Zurbrigg (CFN Subscriptions), and Henry Steger (Member- ship). Professional advice and services were provided by Katryna Coltess (bookkeeping), Sue Anderson (In- vestments), and Eric Leibmann (Accounting Review). Motions: Moved that the financial statements be accepted as a fair representation of the financial position of the Club as of September 30, 2018. Moved that the accounting firm of Welch LLP be contracted to conduct a review of the OFNC’s ac- counts for the fiscal year ending September 30, 2019. ANN MacKENZIE, Treasurer Approved final statements available online at: https://www.canadianfieldnaturalist.ca/index. php/ cfn/article/view/2337/2217. The Canadian Field-Naturalist The Ottawa Field-Naturalists’ Club Awards for 2018, presented February 2019 ELEANOR ZURBRIGG, IRWIN BRopDO, JULIA CIPRIANI, CHRISTINE HANRAHAN, and KAREN McLACHLAN HAMILTON On 23 February 2019 members and friends of the Ottawa Field-Naturalists’ Club (OFNC) gathered for the Club’s Awards Night at St. Basil’s Church in Ottawa to celebrate the presentation of awards for achieve- ments in the previous year. Awards are given to mem- bers or non-members who have distinguished them- selves by accomplishments in the field of natural Member of the Year: Gregory Zbitnew This award is given in recognition of the member judged to have contributed the most to the Club in the previous year. In 1977, two local birders, Paul Matthews and Richard Poulin, wrote a series of articles for Trail & Landscape, on how to find 200 species a year in the Ottawa area. Fast forward to 2017 when a sug- gestion was made to update those articles for Trail & Landscape. Gregory was asked if he was willing to do this and fortunately for us, he agreed. The result is a comprehensive survey entitled: “How to find 250 bird species in the OFNC study area in a single year”. It is no surprise that a dominant interest for OFNC members is birding. These articles have proven ex- ceptionally valuable for all who are keen on birds whether serious birders or casual observers. Even those whose interests may lie elsewhere will find much to enjoy in this excellent series. A nod should also be given to the outstanding photos by Jacques Bouvier accompanying the articles. The articles were published in 2018, Volume 52, one per issue starting with Number 1, and culmin- ating with issue Number 4. Along the way, Gregory gave us informative, well-researched, valuable, and eloquent advice and suggestions on how, when, and where to look for birds. A preamble in the first article talks about “Laying the Foundation” and covers how to find news of re- cent observations (for example, by checking eBird), and what comprises the OFNC study area. It includes definitions of “common”, “uncommon”, “rare”, and other useful and important points. Every article focusses on the birds that can be rea- 96 history and conservation or by extraordinary activ- ity within the Club. Four Club awards were presented for 2018, for: (1) a four-part series on birds in Trail & Landscape, (2) long time service managing the Club’s membership program, (3) establishing and expanding Safe Wings Ottawa, and (4) nature-based education in Ottawa. As well, a President’s Prize was presented. sonably found in the four seasons covered by each of the different issues. The best locations are high- lighted, but not ignored are many other less-vis- ited, excellent spots in which to search for birds. Suggestions as to where some of the seasonal special- ties might be found prove especially useful. For ex- ample, in issue Number 1, a section called “Chasing Winter Birds” provides tips on where one might see winter finches, Bohemian Waxwings, some of the winter raptors, and other groups of birds more likely to be seen in winter. Each article also gives a list of “Important Target Birds” to look for during the particular season. These lists include the uncommon to rare species that may be difficult to find but should be found with some ef- fort. As Gregory notes, it “excludes the very rare spe- cies” that cannot reliably be predicted. In the last issue, Number 4, Gregory provides two excellent summaries in the form of tables. In his own words, Table 1 shows the “appropriate activ- ities/places for birding each week” through the year. Table 2 is a list of all the target species “with their usual habitat and the usual time that you can expect to see them”. Therefore, at a glance, it is easy to find a bounty of relevant information to help plan bird- ing activities. The sheer amount of work that Gregory put into these four lengthy articles is staggering. It is clearly a labour of love, and reveals a solid and deep know- ledge of the world of birds. A wealth of information is contained within this significant work. These articles will become an essential tool for anyone interested in birding, for many years to come. Indeed, they are al- 2019 ready proving very popular. In addition to preparing the above articles, Gregory also maintains and distributes the OFNC weekly online birding report. And as if all that is not enough, Gregory also leads birding trips for the George McGee Service Award: Henry Steger This award is given in recognition of a member who has contributed significantly to the smooth run- ning of the Club over several years. It was very fortuitous in late summer 2001 that Henry Steger was retiring soon, and after seeing an article in the Ottawa Citizen, dropped by and took an interest in the Fletcher Wildlife Garden (FWG) and joined the OFNC. He met David Hobden, then Chair of the Garden’s Management Committee, at an OFNC monthly meeting in early 2002 and offered to volunteer. Henry’s background is in chemistry and mineral processing. His interest in botany came about by ser- endipity in 1981 and he has pursued it since. So upon his retirement, the FWG seemed a perfect way to fol- low this interest and he joined the FWG Management Committee in 2002. Apart from his role on the Management Committee, serving as Chair in 2013, he developed a new database for the bird and wild- life sightings at the FWG (computerized almost 2800 hand-written entries recorded between April 1990 and December 2017). He also developed a database for the FWG library. He participated in planning the renewal strategy for the Amphibian Pond and, more recently, in the replanting and weeding around the Amphibian Pond. He continues to help at the annual native plant sales and provides expertise on native species. Henry continues to be a member of this com- mittee, which would make it 17 years and counting. Henry joined the OFNC Board of Directors as a member at large in 2004 but also was a support- ive voice for the FWG at Board meetings. In 2006, when the Membership Committee was about to lose its Chair, Henry graciously accepted the position. In doing so, he inherited a database system that was in desperate need of an update. Undaunted, Henry took up the challenge and developed a new system CLUB AWARDS 97 OFNC throughout the Ottawa area. For all these reasons we are more than proud to present Gregory Zbitnew with the OFNC Member of the Year award for 2018. (Prepared by Christine Hanrahan) in Microsoft Access that was tailored to the needs of the Club. As Chair, Henry maintains and updates the database on an ongoing basis due to membership changes, mostly renewals and non-renewals, but has also updated the program itself as new features were needed, for example, going to membership renewal by email. He prints the Trail & Landscape (T&L) mailing labels quarterly, manages the emails to mem- bers giving them the latest information on upcom- ing monthly events, and in 2017 created and now up- dates the list of Club members who want access to the Shirleys Bay causeway. Henry is also responsible for the “Welcome New Members” seen in every issue of T&L and its “Golden Anniversary Membership List” that appears annually in the second issue. Throughout the years, Henry has written several articles for T&L. In 2010 and then in 2018 he wrote about his struggles with aster seeds collected on a mature plant in one season not growing into that same plant. Then in 2016 his article about the Tubercled Orchid was an intriguing account of his 16 year hunt for this elusive species. In 2015 when both OFNC Vice-President pos- itions were left vacant, Henry agreed to fill one posi- ton until others could be found. He served as OFNC Vice-President in 2015-2016. Henry has an analytical mind, is keenly observ- ant and is willing to express his opinions. So, when Henry speaks at Board meetings, you can be sure that it was well thought out and important enough to him to contribute to the conversation. His Jack-of-all- trades skillset was honed throughout his career, mak- ing him willing and able to tackle the varied tasks ne- cessary to the Club. That day in 2001 may have been fortuitous, but it has been a real boon for the OFNC. Congratulations Henry for a well-deserved award. (Prepared by Karen McLachlan Hamilton) Conservation Award—Member: Anouk Hoedeman This award recognizes an outstanding contribu- tion by a member in the cause of natural history con- servation in the Ottawa Valley, with particular em- phasis on activities within the Ottawa District. For 2018 we are recognizing the initiative and commitment of Anouk Hoedeman for establishing and expanding Safe Wings Ottawa, a program fo- cussed on bird collision research, education, preven- tion, rescue, and short-term care. Anouk is well known for her work with the OFNC Birds Committee, including the Falcon Watch. She launched an Ottawa chapter of the Toronto-based FLAP (Fatal Light Attraction Program) in 2014, which later evolved into Safe Wings Ottawa, a separ- 98 THE CANADIAN FIELD-NATURALIST ate organization that operates as a committee of the OFNC. Safe Wings estimates that 250 000 birds collide with glass every year in Ottawa. Anouk and her group document about 2000 to 3000 of these—the rest go unnoticed or unreported. Many dead birds are eaten by scavengers or discarded, while many injured ones are taken by predators or fly away to die elsewhere. Safe Wings has a core team of about 15 dedicated volunteers who monitor buildings to collect data and dead bodies, rescue birds, and do what is necessary to support one another and the work they do. Anouk and her fellow volunteers have worked hard to build key relationships with groundskeepers, security guards, and maintenance staff at various buildings downtown and in other areas of the city. As they search every nook and cranny that could shelter a bird, they also educate passersby and encourage them to rescue in- jured birds and report window collisions. Dozens of other Safe Wings volunteers offer their support by transporting birds or helping with other tasks. Safe Wings volunteers also conduct outreach ac- tivities, make presentations to various groups on pre- venting bird collisions, and provide advice on de- signing bird-friendly buildings and on retrofitting existing buildings. Anouk’s personal efforts have re- sulted in the National Capital Commission, Public Services and Procurement Canada, the University of Ottawa, and Carleton University adopting bird- friendly design approaches. In addition, she initiated Mary Stuart Education Award: Bill McMullen This award is given to a member, non-member, or organization, in recognition of outstanding achieve- ments in the field of natural history education in the Ottawa Region. This year’s Mary Stuart Education Award goes to Bill McMullen, a teacher at the Trilltum Public School in Orléans on the eastern edge of Ottawa. Bill grew up in a rural community in the Kawartha Lakes region of central Ontario. There, he was able to spend his childhood years exploring the local forests and fields developing a strong interest in the natural world. As a primary school teacher, he found that he could pass on that passion to his students. On field trips and in the classroom, Bill teaches his young students to be observant of the world around them, and to explore even the small nooks and cran- nies in search of stories that nature can tell us and to experience the “quiet” of the woods. He teaches them the importance of the environment and the interrela- tionships between humans, plants, and animals and encourages them to feel they can be a steward of the Earth. Although Bill is an English and math teacher, Vol. 133 the City of Ottawa’s development of bird-friendly de- sign guidelines, which are expected to come into ef- fect this year. As a result of its reputation, Safe Wings now re- ceives thousands of calls every year for rescue sup- port from individuals and organizations throughout eastern Ontario and western Quebec, including the City of Ottawa, the Ottawa Valley Wild Bird Care Centre, and even the Cornwall SPCA, to name a few. Because there 1s such a demand for help, especially outside the Wild Bird Care Centre’s operating hours, Anouk obtained federal and provincial rehabilitation permits so she could provide medication, fluids, and other life-saving treatment to injured and sick birds. In 2018, Anouk cared for 742 patients on the third floor of her home, which has been transformed into a short-term rehabilitation centre set up with a variety of cages to accommodate and keep birds safe, warm, and fed until they can be released or transferred to the Wild Bird Care Centre. All of this care also requires keeping the space clean, organized, and stocked with supplies ready for injured birds which may arrive at any time. It all makes for unpredictable demands on her time, and a steep learning curve to determine how to handle different species ranging from humming- birds to raptors. We are pleased to recognize the commitment and work of Anouk Hoedeman with this Conservation Award. (Prepared by Julia Cipriani) his knowledge of the natural world gives him the tools he needs. For example, he educates his charges by setting up orienteering courses and taking them skiing, snowshoeing, and hiking. He has even taught children astronomy using his personal telescope. Bill is also a talented and dedicated nature pho- tographer, spending many hours in the field, es- pecially at the MacSkimming Outdoor Education Centre, photographing plants, mushrooms, and wild- life. He often makes videos to show phenomena that occur after school hours or over a long period of time at the MacSkimming Centre (such as a Monarch Butterfly emerging from a chrysalis, or wildlife that passes by a video cam set up on a trail over a period of three months). He incorporates his photos and vid- eos into his own lessons as often as he can and offers them to other teachers at his school to use with their own classes. He also generously shares these vid- eos online. Bill has donated many of his photos and videos to the MacSkimming Centre for their educa- tion programs and also gives students instruction on nature photography. He has also led evening walks 2019 with school children to see and hear owls, resulting in experiences some youngsters will cherish their en- tire lives. By sharing his experience and experiences with his students, Bill McMullen is an example of the kind of teacher who makes a difference. His students are better equipped to understand the conservation issues President’s Prize: Ann MacKenzie This award is given at the President’s discretion in recognition of a member for unusual support of the Club and its aims. The Ottawa Field-Naturalists’ Club is blessed to have members with an interest in natural history but with expertise that serves the Club in areas of man- agement and administration. Ann Mackenzie is such a person. She has served the Club very well in the role of President (2012-2013), Past President (2014—2016), and Treasurer (since 2017). I thank her for this. However, I am recognizing Ann for her on-going work on the Club’s financial sustainability and for her dedication to promote financial professionalism and accountability within the Club. If one reviews how the Club’s financial operations have changed, there is a constant theme. That is, Ann has striven to enhance the Club’s awareness of its financial responsibilities and execution. Today’s Club financial procedures and policies have been modernized and in some areas revolutionized. Before Ann became President, the Club was run- ning unsustainable deficits. Yet there was no attempt to address this issue. Ann dared to suggest an increase in annual fees for the first time in almost 18 years. The practice of a separate fee for paper copies of The Canadian Field-Naturalist was implemented. And she fully supported the project to put The Canadian Field-Naturalist online to reduce costs. In November 2015, Ann responded to an an- nouncement of changes to the Ontario Not-for-Profit Corporations Act with a review and update of the long-obsolete Articles and By-Laws of the Club’s Constitution. This also included a call for all Club CLUB AWARDS 99 of the day, hopefully someday translating that appre- ciation into action for the benefit of the natural world. The Mary Stuart Education Award is a fitting tribute to his vision, skill, energy, and dedication. (Prepared by Irwin Brodo based on input from staff at the Ottawa-Carleton District School Board) committees to update their Terms of Reference. The job is not over. The proclamation of that Act has been delayed. The receipt of a large bequest from Violetta Czasak led to Ann’s recognizing future potential problems with regard to financial management and transparency. She subsequently developed an OFNC Investment Guidelines Policy and initiated the de- velopment of Club policy to administer bequests. The latter has supported local community projects in nat- ural history and scientific research projects and has brought increased public recognition for the Club. Each January the Board of Directors has first-time members. Ann, and probably only she, recognized the need for a guide to help these members become familiar with their roles and expectations. She pub- lished the March 2018 Directors’ Handbook that all Directors have access to online. Lastly, Ann has changed how the office of the Treasurer operates. In the past, the Treasurer did it all by his/herself. Within the last year, she brought in “new” volunteers to assist with investment man- agement and the receipt of monies from membership and donations. Hopefully they will stay in the Club and make other contributions in future. It must also be mentioned that Ann has brought in online pay- ment by the Treasurer of financial claims from Club members and the automated printing of income tax receipts for donations. I wish to award Ann the President’s Prize for 2018, with very best wishes and congratulations. (Prepared by Henry Steger and Diane Lepage, President) THE CANADIAN FIELD-NATURALIST Volume 133, Number 1 Characteristics of Wolverine (Gulo gulo) dens in the lowland boreal forest of north-central Alberta MICHAEL E. JOKINEN, SHEVENELL M. WEBB, DOUGLAS L. MANZER, and ROBERT B. ANDERSON Wolf (Canis sp.) attacks life-like deer decoy: insight into how wolves hunt deer? THOMAS D. GABLE and DANIEL P. GABLE Birds of Mansel Island, northern Hudson Bay ANTHONY J. GASTON Behaviour of a porcupine (Erethizon dorsatum) swimming across a small boreal stream THOMAS S. JUNG More Mountain Chickadees (Poecile gambeli) sing atypical songs 1n urban than in rural areas STEFANIE E. LAZERTE, KRISTEN L.D. MARINI, HANS SLABBEKOORN, MATTHEW W. REUDINK, and KEN A. OTTER Body mass as an estimate of female body condition in a hibernating small mammal CAITLIN P. WELLS, JAMES A. WILSON, DOUGLAS A. KELT, and Dirk H. VAN VUREN Desiccation of herpetofauna on roadway exclusion fencing SEAN P. BoYLe, RACHEL DILLON, JACQUELINE D. LirzGus, and DavID LESBARRERES Monitoring Rock Ptarmigan (Lagopus muta) populations in the Western Aleutian Islands, Alaska CLAIT E. BRAUN, WILLIAM P. TAYLOR, STEVEN M. EBBERT, and Lisa M. SPITLER Japanese Chafl-flower, Achyranthes japonica (Amaranthaceae), on the Erie islands, an invasive plant new to Canada JAMES KAMSTRA Gray Wolf (Canis lupus) recolonization failure: a Minnesota case study L. DaviD MECH, FOREST ISBELL, JIM KRUEGER, and JOHN HART 2019 1 16 20 25 28 34 43 49 56 60 (continued inside back cover) ISSN 0008-3550 TABLE OF CONTENTS (concluded) Volume 133, Number 1 Book Reviews CLIMATE CHANGE: The Uninhabitable Earth: Life After Warming Botany: Michigan Ferns & Lycophytes: A Guide to Species of the Great Lakes Region—Flora of Florida Volume 6 (Dicotyledons, Convolvulaceae through Paulowniaceae)—Identification of Trees and Shrubs in Winter using Buds and Twigs ENTOMOLOGY: Field Guide to the Flower Flies of Northeastern North America ORNITHOLOGY: The Handbook of Bird Families OTHER: The Environment: A History of the Idea—The Great Himalayan National Park: The Struggle to Save the Western Himalayas ZooLocy: Return of the Wolf: Conflict and Coexistence NEw TITLES News and Comment Upcoming Meetings and Workshops Canadian Herpetological Society Annual Conference—Wildlife Society and American Fisheries Society Joint Annual Conference—Student Conference on Conservation Science-New York—Association of Field Ornithologists and the Wilson Ornithological Society Joint Meeting—Entomological Society of Ontario—Entomology 2019 James Fletcher Award for The Canadian Field-Naturalist Volume 132 Diana Beresford-Kroeger: a new book, a life’s work Minutes of the 140" Annual Business Meeting (ABM) of the Ottawa Field- Naturalists’ Club, 8 January 2019 Annual Reports of OFNC Committees for October 2017—September 2018 Club Awards Mailing date of the previous issue 132(4): 31 July 2019 2019 66 68 73 74 76 78 80 84 85 85 88 90 96 The CANADIAN FIELD-NATURALIST A JOURNAL OF FIELD BIOLOGY AND ECOLOGY Promoting the study and conservation of northern biodiversity since 1880 ef Volume 133, Number2 ¢ April-June 2019 Ottawa Field-Naturalists’ Club Club des naturalistes d’Ottawa The Ottawa Field-Naturalists’ Club FOUNDED 1863 (CURRENT INCORPORATION 1879) Patron Her Excellency the Right Honourable Julie Payette, C.C., C.M.M., C.O.M., C.Q., C.D. Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encour- age Investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Honorary Members Ronald E. Bedford Michael D. Cadman J. Bruce Falls Robert E. Lee Allan H. Reddoch Charles D. Bird Paul M. Catling Peter W. Hall John Mcneill Joyce M. Reddoch Fenja Brodo Francis R. Cook Christine Hanrahan Theodore Mosquin Dan Strickland Irwin M. Brodo Bruce Di Labio C. Stuart Houston Robert W. Nero John B. Theberge Daniel F. Brunton Anthony J. Erskine Ross A. Layberry E. Franklin Pope Sheila Thomson 2019 Board of Directors President: Diane Lepage Annie Bélair Edward Farnworth Dwayne Lepitzki Henry Steger 1st Vice-President: Jakob Mueller Fenja Brodo Catherine Hessian —_ Bev McBride Ken Young Recording Secretary: Elizabeth Moore Robert Cermak Anouk Hoedeman Gordon Robertson —__ Eleanor Zurbrigg Treasurer: Ann Mackenzie Owen Clarkin Diane Kitching Jeff Saarela To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofnc.ca. For information on club activities, go to www.ofnc.ca. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this jour- nal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Website: www.canadianfieldnaturalist.ca/index.php/cfn Editor-in-Chief: Dr. Dwayne Lepitzki Assistant Editor: Dr. Amanda Martin Copy Editors: Sandra Garland and Dr. John Wilmshurst Layout: Robert Forsyth Book Review Editor: Dr. Barry Cottam Online Journal Manager: Dr. Bill Halliday Subscription Manager: Eleanor Zurbrigg Author Charges Manager: Ken Young Associate Editors: Dr. Ron Brooks Dr. Jennifer R. Foote Dr. Donald F. McAlpine _ Dr. Jeffery M. Saarela Dr. Carolyn Callaghan Dr. Graham Forbes Dr. Garth Mowat David C. Seburn Dr. Paul M. Catling Thomas S. Jung Dr. Marty Obbard Dr. Jeffrey H. Skevington Dr. Frangois Chapleau Chair, Publications Committee: Dr. Jeffery M. Saarela All manuscripts intended for publication—except Book Reviews—should be submitted through the online submission system at the CFN website: http://www.canadianfieldnaturalist.ca/index.php/cfn/user. Click the “New Submission” link on the right side of the webpage and follow the prompts. 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Subscriptions and Membership: Subscription rates for individuals are $40 (online only), $50 (print only), or $60 (print + online). libraries and other institutions may subscribe for $120 (online only or print only) or $180 (print + online). All foreign print subscrib- ers and members (including USA) must add $10 to cover postage. The Ottawa Field-Naturalists’ Club annual membership fee of $40 (individual), $45 (family), or $20 (student) includes an online subscription to The Canadian Field-Naturalist. Members can receive printed issues of CFN for an additional $30 per volume (four issues). For further details, see http://ofnc.ca/membership-and-donations. The club’s regional journal, 7rail & Landscape, covers the Ottawa District and regional Club events and field trips. It is mailed to all club members. It is available to libraries at $40 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be sent to subscriptions@canadianfieldnaturalist.ca or mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2 Canada. Canada Post Publications Mail Agreement number 40012317. Return postage guaranteed. The Thomas H. Manning fund, a special fund of the OFNC, established in 2000 from the bequest of northern biologist Thomas H. Manning (1911-1998), provides financial assistance for the publication of papers in the CFN by independent (non-institutional) authors, with particular priority given to those addressing Arctic and boreal issues. Qualifying authors should make their application for assistance from the fund at the time of their initial submission. Cover: Collared Pika (Ochotona collaris) in the Ogilvie Mountains at km 158 of Dempster Highway (65.078786°N, 138.360606°W), north of Tombstone Territorial Park, northern Yukon. Aerial and ground surveys were conducted in July 2018 to document oc- currence and general habitat suitability of this cold-adapted, at-risk Beringian species. See article in this issue by Syd Cannings et al., pages 130-135. Photo: Jeffrey H. Skevington, 11 July 2018. The Canadian Field-Naturalist Roadkill of Eastern Newts (Notophthalmus viridescens) in a protected area in Quebec Davip C. SEBURN!*, ELENA KREUZBERG?, and LEAH VIAU2 9 9 'Seburn Ecological Services, 2710 Clarenda Street, Ottawa, Ontario K2B 7S5 Canada ?Canadian Parks and Wilderness Society, Ottawa Valley Chapter, 15 rue Taschereau, suite 240, Gatineau, Quebec J8Y 2V6 Canada “Corresponding author: davidseburn@sympatico.ca Seburn, D.C., E. Kreuzberg, and L. Viau. 2019. Roadkill of Eastern Newts (Notophthalmus viridescens) in a protected area in Quebec. Canadian Field-Naturalist 133(2): 101-104. https://doi.org/10.22621/cfn.v13312.2219 Abstract Roadkill is a threat to populations of many amphibian species, including Eastern Newt (Notophthalmus viridescens), which is widespread in eastern Canada and the northeastern United States. Little is known about the level of road mortality experi- enced by dispersing Eastern Newts in Canada. During extensive road surveys from May to October 2016 and 2017, 279 dead Eastern Newt efts were found on roads in Gatineau Park, Quebec. We found 107 dead Eastern Newts along a 425 m section of road in 2016, but only 30 dead individuals at the same location in 2017. Thus, although the amount of roadkill can vary annually, it may pose a significant threat to the species in some areas. Key words: Eastern Newt; Notophthalmus viridescens; road ecology; salamander; hotspot; Gatineau Park Introduction Roadkill is a threat for many species of amphib- ians (Ashley and Robinson 1996; Andrews ef¢ al. 2008), and they are often the vertebrate group most commonly killed on roads (Ashley and Robinson 1996; Smith and Dodd 2003; Glista et al. 2008). Traffic mortality can have a negative effect on am- phibian populations (Fahrig et a/. 1995; Rytwinski and Fahrig 2012). Although roadkill numbers are often greatest for frogs and toads, roadkill can also be a threat for salamanders (Clevenger et a/. 2001; Gibbs and Shriver 2005; Pagnucco ef al. 2012). Sala- manders may remain immobile on roads when a vehicle approaches, increasing their risk of being run over (Mazerolle 2004). Mortality of >10% of the adult population of Spotted Salamanders (Ambystoma maculata) can lead to population decline and possibly extirpation (Gibbs and Shriver 2005). Here we report on substantial roadkill of Eastern Newt (Notophthalmus viridescens), a species wide- spread in eastern Canada and the northeastern United States (Cook 1984). Eastern Newt is a pond breeding salamander with a complex and variable life cycle. The typical life cycle of Eastern Newt includes four distinct stages: egg, aquatic larva, terrestrial juven- ile or eft, and adult (Gill 1978). After larvae trans- form, the juvenile eft stage disperses from the breed- ing ponds in late summer and early fall (Gill 1978). The eft stage can last for three or more years (Healy 1974), after which maturing efts return to breeding ponds (Hurlbert 1969). Adults may remain in aqua- tic habitats or hibernate on land (Hurlbert 1969; Gill 1978). The eft stage is the primary growth and disper- sal stage in the life cycle, whereas emigrating adults are believed to return to the same breeding ponds re- peatedly (Gill 1978). It is unclear how far efts can dis- perse, but they have been documented to move up to 80 m in a single night (Roe and Grayson 2008) and take up to a year to migrate 800 m (Healy 1974, 1975). The multi-year maturation and dispersal per- iod allows Eastern Newts to colonize newly formed and distant wetlands (Semlitsch 2008) and represents a vulnerable life history stage (Gill 1978). When mi- gration or dispersal routes cross roads, mortality can occur. Methods Road surveys were conducted by car in Gatineau Park, Quebec (45.5°N, 75.8°W) along Meech Lake Road, Gatineau Parkway, Dunlop Road, and Fortune Lake Parkway for a total of ~20 km. Gatineau Park covers an area of 36 131 ha and is managed by the National Capital Commission (NCC 2005). Most of the roads within the park date back to the 1950s or earlier (NCC 2005), but upgrades have occurred over the last several decades. All roads surveyed have A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 101 ©The Ottawa Field-Naturalists’ Club 102 existed for several decades and were paved, two-lane roads. Roadside habitats varied along the roads, but typically the shoulder was narrow with adjacent for- ests coming within a few metres of the road. The car was driven at ~30-—35 km/h and two sur- veyors participated in each survey, scanning both the road surface and road shoulder. All dead animals were removed from the road or road shoulder to pre- vent double counting in a subsequent survey. Surveys were conducted in the morning before scavengers could remove all the roadkill from the previous day or night. Surveys were typically conducted twice a week. Surveys were not conducted during heavy rainfall, but were conducted during light to moderate rain. In addition to driving surveys, walking surveys were conducted at eight locations in the park along the survey route. Walking survey locations were not randomly selected, but based on proximity to desig- nated parking areas in the park to ensure safe parking as well as safety for walking along the roads. Each walking transect was ~90 m in each direction from the stopping point, but extended if high numbers of roadkill were observed. All vertebrate species were recorded, but only results for Eastern Newt are pre- sented here. Weather data from the closest weather station, Chelsea, Quebec, were obtained to compare rainfall patterns with observed roadkill. On dates where no weather data were available from the Chelsea station, data were obtained from the Ottawa International Airport station, ~25 km to the southeast. Results and Discussion We conducted 37 surveys from 12 May until 3 October 2016 and 32 surveys from 15 May to 16 October 2017, for a total of 69 surveys. We found 150 dead Eastern Newts in 2016, and 129 in 2017, for a total of 279. All of the dead Eastern Newts were ju- Number of Eastern Newts N Oo Jun Jul THE CANADIAN FIELD-NATURALIST Vol. 133 veniles (efts). Dead Eastern Newts were found on the road from 18 July to 3 October in 2016, and from 15 May to 8 October in 2017 (Figure 1). The peak mor- tality events were 36 Eastern Newts on 28 July 2016 and 30 Eastern Newts on 5 June 2017 (Figure 1). High counts of dead Eastern Newts likely coincide with peak dispersal events, which usually occur in late summer following rains (Gill 1978; Paton et al. 2000; Leclair et al. 2005). The peak mortality event on 28 July 2016 does not appear to be connected to rainfall as there was no rain on 28 July or the two pre- vious days, although there was 5.2 mm of rain on 25 July (Government of Canada 2019). Rainfall data are missing for the first five days of June 2017, but only 1.0 mm of rainfall was recorded on 4 June 2017, at the Ottawa International Airport. No rainfall was re- corded on 3 June and only 0.2 mm on 2 June 2017. Over 90% of Eastern Newts were found during the walking surveys. We found 107 of the 150 (71.3%) dead Eastern Newts in 2016 along a 425-m section of Meech Lake Road. Smaller numbers of dead Eastern Newts were found at other walking survey locations. The main mortality site was associated with a small creek and numerous wetlands. Habitat at the main mortality site was not obviously different from that at other locations in the park, as creeks, wetlands, and forest are widespread, and the park contains 50 lakes and “several hundred ponds” (NCC 2005). A peak of 30 dead Eastern Newts was found in this mortal- ity “hotspot” on 28 July. Secondary peaks occurred on 12 August (15 Eastern Newts) and 18 August (18 Eastern Newts). Fewer dead Eastern Newts were found in the main hotspot in 2017: only 30 (25.4%) of the 129 individ- uals found that year. A peak of 11 dead Eastern Newts was found in the hotspot on 29 June; no more than five mortalities were found on other survey days. There are many possible reasons why fewer dead Eastern Aug Sep Oct Date FiGurE 1. Number of Eastern Newt (Notophthalmus viridescens) efts found dead on roads in Gatineau Park, Quebec, dur- ing road surveys from 12 May to 3 October 2016 and from 15 May to 16 October 2017. 2019 Newts were found in 2017: survey timing not match- ing peak dispersal events, fewer dispersing efts, or weather conditions limiting dispersal. It does not appear that rainfall is responsible for fewer Eastern Newts being found dead at the main hotspot in 2017, as 2017 was a rainier spring and summer, with 344.6 mm from May to August, compared with only 185.4 mm during the same period in 2016 (Government of Canada 2019). Over the two years, 137 dead Eastern Newts were found at the main hotspot. Likely many other Eastern Newts were killed there, but not observed, given their small size, obliteration by vehicles, and removal by scavengers. Unlike many other salamanders, Eastern Newts are active during the day (Petranka 1998) when motorists are more likely to travel park roads. Other researchers have reported significant num- bers of dead Eastern Newts on roads. Mitchell (2000) found 182 dead Eastern Newts along a 250-m stretch of road on 4 October 1991 in Virginia. An additional 24 dead Eastern Newts were found at the same site over the course of three subsequent surveys in 1992 (Mitchell 2000). The large number of mortalities in 1991 may have been a chance event, or the result of a large production of efts that year (Mitchell 2000). The results from Virginia are similar to our results from Quebec, e., substantially different numbers of ob- served roadkill at the same site in different years. Such results emphasize the fact that one year of road sur- veys will not always provide a complete picture of the overall rate of roadkill. Individuals in our study were not measured, but were assumed to be recently meta- morphosed juveniles dispersing from the natal wetland. The population-level effect of roadkill on sala- manders is not well known. Our results indicate that many Eastern Newts can be killed on roads in some areas. Although efts likely have a relatively high rate of natural mortality, it is unclear what effect the addi- tive mortality of roadkill has on populations. Further research on this topic is warranted. Acknowledgements This study was undertaken by the Canadian Parks and Wilderness Society (CPAWS) Ottawa Valley as part of a larger project on road ecology in eastern Ontario and western Quebec. We thank the numer- ous volunteers who assisted with the road surveys. CPAWS Ottawa Valley is grateful for financial sup- port from the North American Partnership for En- vironmental Community in Action, the Clean Tech Internship Program of Colleges and Canadian Insti- tutes, and the Community Environmental Grant Pro- gram of the City of Ottawa. A permit to conduct re- search in Gatineau Park was issued by the National Capital Commission. SEBURN ET AL.. ROADKILL OF EASTERN NEWTS 103 Literature Cited Andrews, Kk.M., J.W. Gibbons, and D.M. Jochimsen. 2008. Ecological effects of roads on amphibians and rep- tiles: a literature review. Urban Herpetology 3: 121-143. Ashley, E.P., and J.T. Robinson. 1996. Road mortality of amphibians, reptiles and other wildlife on the Long Point Causeway, Lake Erie, Ontario. Canadian Field- Naturalist 110: 403—412. Accessed 30 July 2019. https:// biodiversitylibrary.org/page/34343309, Clevenger, A.P., M. McIvor, D. MclIvor, B. Chruszcz, and K. Gunson. 2001. Tiger Salamander, Ambystoma tigri- num, movements and mortality on the Trans-Canada Highway in southwestern Alberta. Canadian Field-Na- turalist 115: 199-204. Accessed 30 July 2019. https:// biodiversitylibrary.org/page/34995273. Cook, F.R. 1984. An Introduction to Canadian Amphibians and Reptiles. National Museum of Natural Sciences, Ottawa, Ontario, Canada. Fahrig, L., J.H. Pedlar, S.E. Pope, P.D. Taylor, and J.F. Wegner. 1995. Effect of road traffic on amphibian den- sity. Biological Conservation 73: 177-182. https://doi. org/10.1016/0006-3207(94)00102-V Gibbs, J.P., and W.G. Shriver. 2005. Can road mortality limit populations of pool-breeding amphibians? Wet- lands Ecology and Management 13: 281-289. https://do1 .org/10.1007/s11273-004-7522-9 Gill, D.E. 1978. The metapopulation ecology of the Red- spotted Newt, Notophthalmus viridescens (Rafinesque). Ecological Monographs 48: 145-166. https://doi.org/10. 2307/2937297 Glista, D.J., T.L. DeVault, and J.A. DeWoody. 2008. Vertebrate road mortality predominantly impacts am- phibians. Herpetological Conservation and Biology 3: 77-87. Government of Canada. 2019. Historical data: Chelsea and Ottawa MacDonald-Cartier Int’] A. Environment and Natural Resources, Weather, Climate and Hazard, Ottawa, Ontario, Canada. Accessed 25 March 2019. http://climate.weather.gc.ca/historical_data/search_ historic_data_e.html. Healy, W.R. 1974. Population consequences of alternative life histories in Notophthalmus v. viridescens. Copeia 1974: 221-229. https://doi.org/10.2307/1443027 Healy, W.R. 1975. Terrestrial activity and home range in efts of Notophthalmus viridescens. American Midland Naturalist 93: 131-138. https://doi.org/10.2307/2424111 Hurlbert, S.H. 1969. The breeding migrations and inter- habitat wandering of the vermilion-spotted newt Noto- phthalmus viridescens (Rafinesque). Ecological Mono- graphs 39: 465—488. https://doi.org/10.2307/1942356 Leclair, R., M.H. Leclair, and M. Levasseur. 2005. Size and age of migrating eastern red efts (Notophthalmus viridescens) from the Laurentian Shield, Quebec. Jour- nal of Herpetology 39: 51-57. https:/doi.org/10.1670/ 0022-1511(2005)039[0051:saaome]2.0.co;2 Mazerolle, M.J. 2004. Amphibian road mortality in re- sponse to nightly variations in traffic intensity. Herpe- tologica 60: 45-53. https://doi.org/10.1655/02-109 Mitchell, J.C. 2000. Mass mortality of red-spotted newts (Notophthalmus viridescens viridescens Rafinesque) on acentral Virginia road. Banisteria 15: 45—47. 104 NCC (National Capital Commission). 2005. Gatineau Park master plan. National Capital Commission, Ottawa. Ac- cessed 16 March 2019. http://ncc-ccn.gce.ca/our-plans/ gatineau-park-master-plan. Pagnucco, K.S., C.A. Paszkowski, and G.J. Scrimgeour. 2012. Characterizing movement patterns and spatio- temporal use of under-road tunnels by Long-toed Sala- manders in Waterton Lakes National Park, Canada. Co- peia 2012: 331-340. https://doi.org/10.1643/CE-10-128 Paton, P., S. Stevens, and L. Longo. 2000. Seasonal phe- nology of amphibian breeding and recruitment at a pond in Rhode Island. Northeastern Naturalist 7: 255-269. https://doi.org/10.2307/3858358 Petranka, J.W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington, DC, USA. Roe, A.W., and K.L. Grayson. 2008. Terrestrial move- THE CANADIAN FIELD-NATURALIST Vol. 133 ments and habitat use of juvenile and emigrating adult Eastern Red-spotted newts, Notophthalmus virides- cens. Journal of Herpetology 42: 22-30. https://doi.org/ 10.1670/07-040.1 Rytwinski, T., and L. Fahrig. 2012. Do species life history traits explain population responses to roads? A meta- analysis. Biological Conservation 147: 87—98. https:// doi.org/10.1016/j.biocon.2011.11.023 Semlitsch, R.D. 2008. Differentiating migration and dis- persal for pond-breeding amphibians. Journal of Wild- life Management 72: 260-267. https://doi.org/10.2193/ 2007-082 Smith, L.L., and C.K. Dodd, Jr. 2003. Wildlife mortal- ity on US highway 441 across Paynes Prairie, Alachua County, Florida. Florida Scientist 66: 128-140. Received 1 February 2019 Accepted 30 July 2019 The Canadian Field-Naturalist First records of Finescale Dace (Chrosomus neogaeus) in Newfoundland and Labrador, Canada DONALD G. KEEFE!*, ROBERT C. PERRY', and GREGORY R. MCCRACKEN’ 'Department of Fisheries and Land Resources, Forestry and Wildlife Research Division, P.O. Box 2007, Corner Brook, Newfoundland and Labrador A2H 7S1 Canada *Department of Biology, Dalhousie University, 1355 Oxford Street, Halifax, Nova Scotia B3H 4R2 Canada “Corresponding author: donkeefe@gov.nl.ca Keefe, D.G., R.C. Perry, and G.R. McCracken. 2019. First records of Finescale Dace (Chrosomus neogaeus) in Newfoundland and Labrador, Canada. Canadian Field-Naturalist 133(2): 105-112. http://doi.org/10.22621/cfn.v133i2.1991 Abstract The island of Newfoundland has no official record of cyprinid fishes. Here, we report the discovery of a minnow, Finescale Dace (Chromous neogaeus) from four ponds located in a first order tributary of the Exploits River, in central Newfoundland. This finding represents the first record of the species in the province. The location where the species was found is in a local- ized, central portion of insular Newfoundland, therefore, the most parsimonious explanation for this new record is that it was an illegal, intentional introduction. Such introductions in other provinces have occurred by anglers who felt it would serve as a forage fish for other species. The consequences of this introduction to native species are unknown; however, the dace’s local abundance, foraging behaviour, and reproductive capacity are discussed in terms of the interspecific competi- tion with native species. Key words: Finescale Dace; Chrosomus; Goldfish; intentional introduction; interspecific competition; exotic species Introduction The province of Newfoundland and Labrador has a very low rate of introduction for exotic fish (van Zyll de Jong et al. 2004). There are 28 native freshwater fish species in Newfoundland and Labrador; 15 spe- cies can be found in insular Newfoundland with an additional 13 species occurring in Labrador (van Zyll de Jong et al. 2004; Table 1). For the island portion of the province there have been five attempts at species introductions. These introductions were all salmonids including Brown Trout (Sa/mo trutta), Rainbow Trout (Onchorhynchus mykiss), Lake Whitefish (Corego- nus clupeaformis), Lake Trout (Salvelinus namay- cush), and Pink Salmon (Oncorhynchus gorbuscha, Table 1). However, of these five introductions only the first two succeeded. These species were introduced by government agencies intending to use stocking programs to enhance freshwater fisheries (Scott and Crossman 1964; Hustins 2007). The only other account of a species introduction documented for the province is Goldfish (Carassius auratus). This species has been reported 1n three loca- tions, but as yet, has not been substantiated by a gov- ernment agency. The three alleged locations include Mundy Pond (47.55152°N, 52.73891°W) centrally lo- cated in the city of St. John’s, a small pond near the town of Heart’s Delight (47.78691°N, 53.46752°W), and Janes Pond (47.16617°N, 55.16229°W) located in the town of Marystown. All of these locations are in proximity to urban areas, suggesting the likelihood of intentional release of household aquaria pets. In La- brador, there are no records of exotic freshwater fish. On 4 June 2010 the provincial department of Environment and Conservation received a request to identify an unusual fish that had allegedly been angled by a sport fisher in central Newfoundland. The examination of a photograph suggested the unknown species was a minnow from Cyprinidae family. The fish was reportedly angled from a pond on a tributary of the Exploits River. A field trip was subsequently planned to verify the report or determine the extent of its presence. Herein we report on the occurrence of an exotic invasive minnow species existing in four ponds of a first order tributary of the Exploits River watershed, central Newfoundland. Methods Sample collection To verify the species present and the extent of their distribution two site visits occurred. The first occurred 4-16 June 2012 and the second 9-17 June 2014. During the latter trip the number of sites 105 ©The Ottawa Field-Naturalists’ Club 106 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE I. A list of native and introduced freshwater fish species of Newfoundland and Labrador, Canada (van Zyll de Jong et al. 2004). Common name American Eel Fourspine Stickleback Threespine Stickleback Blackspotted Stickleback Ninespine Stickleback Atlantic Salmon Brook Trout Arctic Char Lake Trout Lake Whitefish Round Whitefish Rainbow Smelt Sea Lamprey Alewife American Shad Atlantic Tomcod Banded Killifish Mummichog Northern Pike Lake Chub Longnose Dace Northern Pearl Dace Longnose Sucker White Sucker Burbot Mottled Sculpin Slimy Sculpin Logperch Rainbow Trout* Brown Trout* Goldfish* *Introduced species. Scientific name Anguilla rostrata Apeltes quadracus Gasterosteus aculeatus Gasterosteus wheatlandi Pungitius pungitius Salmo salar Salvelinus fontinalis Salvelinus alpinus Salvelinus namaycush Coregonus clupeaformis Prosopium cylindraceum Osmerus mordax Petromyzon marinus Alosa pseudoharengus Alosa sapidissima Microgadus tomcod Fundulus diaphanus Fundulus heteroclitus Esox lucius Couesius plumbeus Rhinichthys cataractae Margariscus nachtriebi Catostomus catostomus Catostomus commersonii Lota lota Cottus bairdii Cottus cognatus Percina caprodes Onchorhynchus mykiss Salmo trutta Carassius auratus auratus searched was expanded. Initial samples were taken from four headwater lakes located on the south side of the Exploits River near the town of Bishop’s Falls. The lakes are located on a first order tributary (Jumpers Brook 49.02667°N, 55.40216°W: Tributary 1; Figure 1). Jumpers Brook flows into the main stem of the Exploits River (49.02857°N, 55.40164°W), en- tering the Exploits ~8 km upstream from the Atlantic Ocean (in Exploits Bay). Three of the sampled ponds (Mina 1, Mina 2, and Mina 3) were in close proxim- ity (<1 km) to each other and were both small (sur- face area <0.10 km?) and shallow (<2 m depth). The fourth lake, Mina 4 was larger (surface area = 0.35 km?) and deeper (>4 m depth). Mina 4 was ~2 km upstream from Mina 3. The substrate in each pond was mud with scattered rubble and cobble with emer- gent grasses surrounding the shorelines. The water was dark in colour, the product of humic conditions. During the 2012 sampling, surface temperatures ranged from 8°C to 10°C. In 2014 the temperatures were between 11°C and 20°C. Newfoundland Labrador + + + = + +} + —- + + + 1 + + + + — + — + — + + + + + + + + + + | + — + ‘a, — + - + — 1 - + _ + — + — + — + — + — + + sty + —_ + i During the first sampling event, in 2012, both fyke (FYKE-001-06, 10 mm mesh, Shandong, China) and gill nets (Miller Nets, Memphis, Tennessee, USA) were used. Gill nets were composed of nylon monofila- ment and each set consisted of two stretch mesh sizes measuring 1.91 cm and 2.54 cm, with the smallest mesh being set closest to the shoreline. Three gill nets were set in Mina 1, two in Mina 2 and 3, and four sets were placed in Mina 4. All gill nets were set perpen- dicular to the shoreline and were set overnight, for an average of 12 h. Fyke (trap) net exterior netting con- sisted of 1.91 cm stretch mesh size with front hoop of 1 m through to four cylindrical interior hoops with a tied trap end. These nets were set with a 9.8 m mesh lead extending from the front hoop perpendicular to the shoreline. This lead net also consisted of 1.91 cm stretch mesh size. Three small fyke nets were set in the littoral area of Mina 2. All captured native species were retained for measurement and stomach examinations in the laboratory. A subset of unidentified minnow spe- cies were retained and preserved in 95% ethanol. 49°3'N Mina 1 / 49°0'N KEEFE &T AL.: FINESCALE DACE IN NEWFOUNDLAND a Electrofishing site Tributary 1 -—Tributary 2 /Mina 2—@ ~~__Electrofishing @ Mina 3 |_ _' Watershed boundary oO." 4 || —— Road 48°57'N Kilometres 55°27'W 55°24'W FicurE 1. The four ponds (Mina 1, 2, 3, and 4) on Tributary 1 from which Finescale Dace (Chrosomus neogaeus) was sampled, central Newfoundland. Also shown are the locations of Tributary 2, Ponds A and B, and the locations of the elec- trofishing stations. In 2014, sampling was repeated on each of the four ponds originally sampled in 2012. However, instead of gill and fyke nets, minnow pots were used as they have proven to be very effective when sampling dace (He and Lodge 1990). The minnow pots (Gees Feets G-40 Minnow trap, Tackle Factory, Fillmore New York, USA) were constructed of galvanized steel wire with length 42 cm, diameter 23 cm, and square mesh size of 6 mm. Thirty-nine overnight pot sets were placed in ponds Mina 1, 2, 3, and 4 (Table 2). A 107 108 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Year and location of sampling, gear type, and numbers set (n). Frequency of fish sampled per species for all ponds 4—16 June 2012 and 9-17 June 2014 in central Newfoundland. All sets were overnight. Brook Trout Year Location Gear Type (”) (Salvelinus fontinalis) 2012 Mina 1 Gill (3) 29 Mina 2 Gill (2) 10 Mina 2 Fyke (3) 7 Mina 3 Gill (2) — Mina 4 Gill (4) 9 2014 Mina 1 Pot (16) 2 Mina 2 Pot (9) — Mina 3 Pot (6) — Mina 4 Pot (8) — Pond A Pot (4) — Pond B Pot (3) 6 sub-sample of 20 unidentified minnow species were retained, preserved in 95% ethanol, and sent to the Department of Biology at Dalhousie University for genetic-based identification. The sub-sample was a random selection of 10 male and 10 female fish. To determine if the dace were isolated to just the four ponds or were present in other areas of Jumpers Brook, the lower (1.5 km) and upper (1 km) reaches of Tributary | (Figure 1) were sampled by electrofish- ing. Two small ponds located in another first order tributary were also sampled (Tributary 2; Figure 1). In Tributary 2, four pots were set in Pond A and three pots were placed in Pond B. Morphometric and subsequent genetic analysis External morphological measures were recorded for 200 dace chosen from the 2014 collection (Table 3). With the exception of fork length, the morpho- logical characters measured followed those described Threespine Finescale Dace Atlantic Salmon Stickleback (Chrosomus (Salmo salar) (Gasterosteus neogaeus) aculeatus) 2 1 D) 10 9 l l 1 3 = — - 3 m.' 26 391 = 19 24 _ 43 2 7 15 110 == 29 = by Hubbs and Lagler (1958) and Doeringsfeld et al. (2004) which included the predorsal length, pec- toral fin position, head length, maximum body width, maximum body depth, caudal peduncle depth, inter- orbital width, gape width, and the upper jaw length. Species identification using the morphometric data was attempted but proved difficult for several reasons. Given the morphological variability among the sam- pled dace it was determined that it could be one, or a combination of, three possible species, each sharing very similar physical characteristics. The three spe- cies included Northern Redbelly Dace (Chrosomus eos (Cope, 1862)), Finescale Dace (Chrosomus neo- gaeus (Cope, 1867)), or their hybrid Chrosomus eos- neogaeus. Subsequently, genetic analysis was used for determining species. Nevertheless, this did not re- move all the difficulty in identification. Standard DNA barcoding/sequencing methods using the mitochondrial COl gene would not be suf- TABLE 3. Variation in morphological characters measured from 200 Chrosomus sp. collected 9-17 June 2014 from ponds in the Jumpers Brook watershed, central Newfoundland. Measurements as described by Hubbs and Lagler (1958) and Doeringsfeld et a/. (2004): fork length (FL), predorsal length (PDL), pectoral fin position (PFP), head length (HDL), max- imum body width (BYW), maximum body depth (BYD), caudal peduncle depth (CPD), interorbital width (IOW), gape width (GPW), upper jaw length (UJL). Character Range 95% Cl Mean SE FL 39.00—93.00 64.7-68.7 66.69 0.998 TBM 0.71-10.23 3.64—4.26 3.95 0.157 PDL 18.00—48.10 32.80-34.86 33.83 0.520 PFP 6.30—20.70 13.67-14.50 14.08 0.210 HDL 7.90—21.23 13.81-14.65 14.23 0.214 BY W 4.40-15.10 8.28— 8.92 8.59 0.162 BYD 7.50—20.80 13.35-14.25 13.80 0.228 CPD 0.30-3.60 1.27-1.46 Les? 0.046 IOW 2.50—8.50 5.58-5.97 5.78 0.098 GPW 3.90-13.20 8.01—-8.55 8.28 0.136 WIE 1.70—8.90 4.64-4.95 4.79 0.079 2019 ficient for all necessary species identifications in this case. Hybrid individuals (C. eos-neogaeus) share mitochondrial DNA with their maternal parent, most commonly C. neogaeus (Binet and Angers 2005) and these standard identification techniques would not allow us to distinguish these two complexes. As a result, a method of species identification based on nuclear DNA, in addition to mitochondrial DNA, was required. Caudal fin tissues were digested with proteinase K (Bio Basic Inc., Markham, Ontario, Canada) for ~8 h at 50°C. DNA was extracted from these tissue di- gests following a glassmilk approach modified from Elphinstone et a/. (2003) for use with a Multiprobe II liquid handling system (PerkinElmer, Waltham, Massachusetts, USA). The resulting DNA was exam- ined for quality and quantity by means of gel electro- phoresis using a 1% agarose gel. To perform the species identification, we followed the recommendations found in Binet and Angers (2005) in which the GH and the PEG1/MEST nuclear loci were amplified using polymerase chain reaction (PCR). The resulting PCR products were visualized on a 2% agarose gel and examined for band combina- tion and size to detect the presence of C. neogaeus, C. eos, or their hybrid. Five complete Chrosomus sp. from the 2012 col- lection were archived at the Royal Ontario Museum (accession number: ROM 104124). Additionally, 20 caudal fin tissues, used for genetics-based identifi- cation, were transferred to The Rooms Corporation of Newfoundland and Labrador, Provincial Museum Division (accession number: NFM T-2018-63). All specimens were preserved in 95% ethanol. KEEFE ET AL.: FINESCALE DACE IN NEWFOUNDLAND 109 Results Species account In 2012, native fish species captured included Brook Trout (Sa/velinus fontinalis), Atlantic Salmon (Salmo salar), and Threespine Stickleback (Gastero- steus aculeatus, Table 2). In addition, 14 dace were also sampled from three of the four ponds surveyed (Mina 1, 2, and 4; Table 2). Stomach examinations re- vealed that one Brook Trout, from Mina 1, contained approximately 20 partially digested young-of-the- year dace (Figure 2a). In 2014, 527 dace were sampled from all four lakes, for an average capture of 13.5 fish per pot. Figure 2b shows three representative specimens from the collection. In Tributary 2 we did not capture any dace in either Ponds A or B, nor did we sample any dace when electrofishing the lower and upper reaches of Tributary 1. Only native species Brook Trout, American Eel (Anguilla rostrata), Atlantic Salmon, and Threespine Stickleback were sampled. Genetic results Of the 20 selected samples, 16 yielded sufficient DNA quantity and quality for species identification. The remaining four were excluded due to DNA qual- ity concerns. Our examination of the amplifications of the GH locus showed band size uniformity among all individuals, with a band size of approximately 250 bp. This particular fragment/band size is indica- tive of C. neogaeus (Binet and Angers 2005). A sim- ilar result was observed for the PEGI/MEST ampli- fications: band size uniformity of approximately 177 bp. Again, this particular band size is indicative of C. neogaeus (Binet and Angers 2005). There was no ie FiIGuRE 2. a. Stomach contents of a Brook Trout (Sa/velinus fontinalis) captured in pond Mina 1, central Newfoundland, on 7 June 2012, showing approximately 20 partially decomposed immature cyprinid species. b. Finescale Dace (Chrosomus neogaeus) sampled from pond Mina 2, central Newfoundland, on 14 June 2014 Photo: a. Robert Perry. Photo: b. Donald Keefe. 110 evidence of any other band size or combination in any of the 16 individuals tested at either of these two nu- clear DNA loci, indicating the absence of both hybrid C. eos-neogaeus and C. eos individuals. Discussion The record of Finescale Dace in four headwater ponds in the Exploits River watershed represents the first report of an introduced Leuciscinae spe- cies for insular Newfoundland. The species comple- ment for the island portion of the province contains only euryhaline (saltwater tolerant) species (Scott and Crossman 1964; van Zyll de Jong et al. 2004) and Finescale Dace is not considered euryhaline. Therefore, it is unlikely that the species colonized the area naturally and indicates that an exotic intro- duction has occurred. The known distribution of Finescale Dace extends across the southern and north- west parts of Canada (Scott and Crossman 1998). In the east, its distribution reaches New Brunswick and Nova Scotia while in the United States it can be found in the Mississippi and Missouri river drainages (Stasiak 1980; Page and Burr 1991). The origin for the Jumpers Brook population is uncertain. Because young-of-the-year minnows were found during stomach examination of Brook Trout it is evidence that the nonindigenous species is estab- lished. The habitat preferences for Finescale Dace in- clude acidic beaver ponds, boggy waters, and small lakes with bottom detritus and silt (Stasiak and Cunnigham 2006). The locations in which we discov- ered Finescale Dace fit the habitat requirements for the species (Mee and Rowe 2010). In accordance with our 2014 field sampling, we did not find evidence for the presence of Finescale Dace in either Tributary 1 or in the headwater ponds of Tributary 2. It would appear that the presence of the dace is currently limited to the four headwater ponds of Tributary 1. We believe the containment is owing to the presence of beaver dams in ponds Mina 1 and Mina 2 which limit the outflow of water and impedes fish movement. The outflow from Mina 2 is downstream from Mina 3 and 4 and therefore con- trols the flow of water from these two ponds. If the intent of the introduction was to provide a food source for both Brook Trout and juvenile Atlantic Salmon, it may have been short-sighted. Although we did find juvenile dace in the stomach of an adult Brook Trout, it is unclear how additional competi- tion for food sources will affect native fish popula- tions into the future. Finescale Dace are sight feeding, carnivorous predators and have been documented as eating a wide variety of food items including aqua- tic insects, vegetation, and molluscs (Stasiak and Cunnigham 2006; Mee et al. 2013). For example, adult THE CANADIAN FIELD-NATURALIST Vol. 133 fish have been documented eating midges (Diptera), beetles (Coleoptera), caddisflies (Trichoptera), may- flies (Ephemeroptera), and bugs (Hemiptera; Stasiak 1972). Native species such as Brook Trout, Atlantic Salmon, and Threespine Stickleback depend upon micro and macro invertebrates, particularly at juven- ile stages (Scott and Crossman 1998). Further, these types of minnows have a large reproductive capacity. It has been reported that Northern Redbelly Dace can produce 316000 fish per ha of surface area in a sin- gle season (Cooper 1935). Finescale Dace also has a substantial reproductive capability. For example; a 70 mm female may produce 2600 eggs (Stasiak 2011). Given the potential for these types of cyprinids to have a large reproductive capacity, it is probable that the presence of Finescale Dace will lead to inter- specific resource competition with native species. Competition among piscivores and planktivores can have a large influence on rates of primary produc- tion in an aquatic ecosystem (Carpenter ef al. 1985; Carpenter and Kitchell 1988). Following the introduc- tion of Eurasian Minnow (Phoxinus phoxinus) into a Norwegian subalpine lake, O@vre Heimdalsvatn, re- searchers witnessed major changes to both the struc- ture and composition of macroinvertebrate benthos (Naestad and Brittain 2010). These changes led to negative effects on the native Brown Trout popu- lation, including reduced annual recruitment and growth rates. Researchers attributed the changes to competition for food (Borgstrom ef al. 2010). There may also be a direct effect on native species abundance, as these types of predatory minnows have been documented eating the eggs and young of other species. A dissected large male Finescale Dace was reported having its intestinal tract full of fish eggs (presumed to be the eggs of the Northern Redbelly Dace; Stasiak 1972). Becker (1983) reported that dace will eat guppies in an aquarium setting. Furthermore, it has been reported that Northern Redbelly Dace, a species whose dietary preferences overlap with the Finescale Dace (Cochran et al. 1988), would eat Smallmouth Bass (Micropterus dolomieu) fry (Scott and Crossman 1999). Thus, it is possible that fry of Brook Trout and stickleback could also provide a food source for Finescale Dace. Further work is required to determine the short and long-term impacts of this introduction on native fauna. The discovery of Finescale Dace in this particu- lar watershed is disconcerting. The Exploits River represents one of the largest Atlantic Salmon pro- ducing rivers in North America (Pinfold 2011). The spread of this stenohaline species throughout the Exploits watershed may be limited by the presence of the Grand Falls Hydro-electric facility. The facility serves as a major barrier to fish passage and is located 2019 just upstream from where the tributary containing dace enters the main branch of the Exploits River. The origin of Finescale Dace found in the four headwater ponds is uncertain. However, there are two possible explanations. The species current docu- mented range limit in eastern Canada are the prov- inces of New Brunswick and Nova Scotia (Stasiak 1980; Page and Burr 1991). Therefore, it is possible that anglers imported the species as an illegal form of bait from the Maritime region. However, this ex- planation seems improbable given that these ponds are small in size and not known as a favoured area for recreational fishing. Another possible explanation may be that minnows kept in aquaria by area residents were released. Local residents interviewed regarding this possibility suggested that an unknown species of minnow had been housed in close proximity to the headwater ponds. Speculation by those interviewed was that minnows were imported from Labrador. Although interesting, this idea is unsubstantiated. The presence of Finescale Dace in Labrador (even if remote) is an intriguing possibility. The present range distribution for Finescale Dace does not ex- tend into Labrador and the only known species of dace existing in the province is Northern Pearl Dace (Margariscus nachtriebi). Thus, one may speculate that an undiscovered species is present or, alterna- tively, the current identification for the species in Labrador maybe an error. Further genetic work 1s re- quired to investigate these possibilities. Our findings suggest these minnow species are readily trapped and therefore could easily be moved as bait to other regions of the province. However, it is illegal in Newfoundland and Labrador to transport live fish or to use live fish as bait in inland waters. It is hoped that these regulatory restrictions will serve as a deterrent to the spread of the species. Additionally, to limit the spread of Finescale Dace, we recommend the launch of a public education and awareness pro- gram pertaining to the existence of these species in the province. Such an initiative would be beneficial and low cost for regulatory authorities. Through this effort, it is hoped that further illegal introductions or transfers could be curtailed. Additionally, the four ponds where these dace were found were small and therefore we recommend some consideration should be given to an eradication program to halt their spread. Acknowledgements We thank Dr. Daniel Ruzzante and his laboratory staff of Dalhousie University for assistance in iden- tifying the species of dace. We thank Amy Lathrop of the Royal Ontario Museum and Nathalie Dyan- Chékar of The Rooms Corporation of Newfoundland and Labrador for cataloguing specimens used in this KEEFE ET AL.: FINESCALE DACE IN NEWFOUNDLAND Tal report. Thanks to Roger Ward for his field assistance and we are grateful to Adam Hicks, Phillip Hillier, and Mark Young for laboratory assistance. Finally, thanks to Blair Adams and Shelley Garland for edi- ting the manuscript. We would like to dedicate this manuscript to the co-author’s mother, Mina Keefe (1945-2012), whose kindness and field assistance during our excursion to central Newfoundland will not be forgotten. Literature Cited Becker, G.C. 1983. Fishes of Wisconsin. University of Wis- consin Press, Madison, Wisconsin, USA. Binet, M.C., and B. Angers. 2005. Genetic identification of members of the Phoxinus eos-neogaeus hybrid com- plex. Journal of Fish Biology 67: 1169-1177. https://doi. org/10.1111/j.0022-1112.2005.00802.x Borgstrom, R., J. Museth, and J.E. Brittain. 2010. The brown trout (Sa/mo trutta) in the lake, O@vre Heimdals- vatn: long term changes in population dynamics due to exploitation and the invasive species, European minnow (Phoxinus phoxinus). Hydrobiologia 642: 82—91. https:// doi.org/10.1007/s10750-010-0161-7 Carpenter, S.R., and J.F. Kitchell. 1988. Consumer con- trol of lake productivity. BioScience 38: 764—769. https:// doi.org/10.2307/1310785 Carpenter, S.R., J.F. Kitchell, and J.R. Hodgson. 1985. Cascading trophic interactions and lake productivity. BioScience 35: 634—639. https://doi.org/10.2307/1309989 Cochran, P.A., D.M. Lodge, J.R. Hodgson, and P.G. Knapik. 1988. Diets of syntopic finescale dace, Pho- xinus neogaeus, and northern redbelly dace, Phoxinus eos: a reflection of trophic morphology. Environmental Biology of Fishes 22: 235-240. https://doi.org/10.1007/ bf00005384 Cooper, G.P. 1935. Some results of forage fish investiga- tions in Michigan. Transactions of the American Fish- eries Society 65: 132-142. https://doi.org/10.1577/1548 -8659(1935)65[132:SROFFI]2.0.CO;2 Doeringsfeld, M.R., I.J. Schlosser, J.F. Elder, and D.P. Evenson. 2004. Phenotypic consequences of genetic variation in a gynogenetic complex of Phoxinus eos- neogaeus Clonal fish (Pisces: Cyprinidae) inhabiting a heterogeneous environment. Evolution 58: 1261-1273. https://doi.org/10.1111/j.0014-3820.2004 tb01705.x Elphinstone, M.S., G.N. Hinten, M.J. Anderson, and C.J. Nock. 2003. An inexpensive and high-throughput procedure to extract and purify total genomic DNA for population studies. Molecular Ecology Notes 3: 317— 320. https://doi.org/10.1046/).1471-8286.2003.00397.x He, X., and D.M. Lodge. 1990. Using minnow traps to esti- mate fish population size: the importance of spatial dis- tribution and relative species abundance. Hydrobiologia 190: 9-14. https://doi.org/10.1007/BF00020683 Hubbs, C.L., and K.F. Lagler. 1958. Fishes of the Great Lakes Region. University of Michigan Press, Ann Ar- bor, Michigan, USA. Hustins, D. 2007. Brown Trout and Rainbow Trout: a Jour- ney into Newfoundland Waters. Tight Line Publishers, St. John’s, Newfoundland and Labrador, Canada. 112 Mee, J.A., F. Noddin, J.R. Hanisch, W.M. Tonn, and C.A. Paszkowski. 2013. Diets of sexual and sperm-de- pendent asexual dace (Chrosomus spp.): relevance to niche differentiation and mate choice hypotheses for co- existence. Oikos 122: 998-1008. https://doi.org/10.1111/ j.1600-0706.2012.00178.x Mee, J.A., and L. Rowe. 2010. Distribution of Phoxinus eos, Phoxinus neogaeus, and their asexually-reproduc- ing hybrids (Pisces:Cyprinidae) in Algonquin Provin- cial Park, Ontario. Plos One 5: e13185. https://doi.org/ 10.1371/journal.pone.0013185 Naestad, F., and J.E. Brittain. 2010. Long-term changes in the littoral benthos of a Norwegian subalpine lake fol- lowing the introduction of the European minnow (Pho- xinus phoxinus). Hydrobiologia 642: 71-79. https://doi. org/10.1007/s10750-010-0160-8 Page, L.M., and B.M. Burr. 1991. A Field Guide to the Freshwater Fishes: North America North of Mexico. Houghton Mifflin, Boston, Massachusetts, USA. Pinfold, G. 2011. Economic value of wild Atlantic Salmon. Gardner Pinfold Consultants Inc., Halifax, Nova Scotia, Canada. Scott, W.B., and E.J. Crossman. 1964. Fishes Occurring in the Fresh Waters of Insular Newfoundland. Queen’s Print- er and Controller of Stationary, Ottawa, Ontario, Canada. Scott, W.B., and E.J. Crossman. 1998. Freshwater Fishes of Atlantic Canada. Galt House Publications Ltd., Oak- THE CANADIAN FIELD-NATURALIST Vol. 133 ville, Ontario, Canada. Stasiak, R.H. 1972. The morphology and life history of the finescale dace, Pfrille neogaea, in Itasca State Park, Minnesota. Ph.D. thesis, University of Minnesota, Min- neapolis, Minnesota, USA. Stasiak, R.H. 1980. Phoxinus neogaeus Cope. Page 336 in Atlas of North American Freshwater Fishes. Edited by D.S. Lee, C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R. Stauffer. North Carolina State Museum Natural History, Raleigh, North Carolina, USA. Stasiak R.H. 2011. Reproduction, age, and growth of the finescale dace, Chrosomus neogaeus, in Minnesota, Trans- actions of the American Fisheries Society 107: 720-723. http://do1.org/fftzz9 Stasiak, R., and G.R. Cunningham. 2006. Finescale Dace (Phoxinus neogaeus): a technical conservation assess- ment. USDA Forest Service, Rocky Mountain Region. Accessed 10 October 2019. https://www.fs.usda.gov/ Internet/FSE_-DOCUMENTS‘stelprdb5206787.pdf. van Zyll de Jong, M.C., R.J. Gibson, and I.G. Cowx. 2004. Impacts of stocking and introductions on fresh- water fisheries of Newfoundland and Labrador, Canada. Fisheries Management and Ecology 11: 183-193. https:// dot.org/10.1111/j.1365-2400.2004.00390.x Received 4 October 2017 Accepted 31 July 2019 The Canadian Field-Naturalist Albinism in Orange-footed Sea Cucumber (Cucumaria frondosa) in Newfoundland EMALINE M. MONTGOMERY!”, TIFFANY SMALL!, JEAN-FRANCOIS HAMEL’, and ANNIE MERCIER! 'Department of Ocean Sciences, Memorial University, St. John’s, Newfoundland and Labrador A1C 5S7 Canada Society for Exploration and Valuing of the Environment, Portugal Cove-St. Phillips, Newfoundland and Labrador AlM 2B7 Canada “Corresponding author: e.montgomery@mun.ca Montgomery, E.M., T. Small, J-F. Hamel, and A. Mercier. 2019. Albinism in Orange-footed Sea Cucumber (Cucumaria fron- dosa) in Newfoundland. Canadian Field-Naturalist 133(2): 113-117. https://do1.org/10.22621/cfn.v13312.2047 Abstract Orange-footed Sea Cucumber (Cucumaria frondosa; Echinodermata: Holothuroidea) is a dark-brown species that is broadly distributed in North Atlantic and Arctic waters. Here, we document the rare occurrence of colour morphs showing vari- ous degrees of albinism, from totally white to faint orange pigmentation. These unusually coloured individuals were found across a broad distribution range in eastern Canada and northeastern United States, with their occurrence in Newfoundland samples ranging from 0.2% to 0.5%. Two fully albino individuals were noticeably smaller than other colour morphs. The occurrence of rare, unusually coloured sea cucumbers is important from an ecological standpoint and may also have com- mercial implications. Key words: Orange-footed Sea Cucumber; Cucumaria frondosa; albinism; colour diversity; Holothuroidea; sea cucumber; North Atlantic Introduction Unusually coloured and albino (white) sea cucum- bers are rare in nature and typically have higher market value, particularly in Asia. Albino individ- uals have sold at auction for US$23 000/kg (normal value US$2000-—2500/kg for the species; Tse 2015). To date, published records of albinism in holothu- roid echinoderms (nomenclature according to World Register of Marine Species; WoRMS 2019) include Japanese Sea Cucumber (Apostichopus japonicus) in the temperate Western Pacific (Lin et a/. 2013), White Teatfish (Holothuria fuscogilva) in the tropical Indo- Pacific (Friedman and Tekanene 2005), Brown Sea Cucumber (/sostichopus fuscus) in the tropical east- ern Pacific (Fernandez-Rivera Melo et al. 2015), Four- sided Sea Cucumber (/sostichopus badionotus) in the Caribbean (Wakida-Kusunoki et a/. 2016), Climbing Sea Cucumber (Ocnus planci) in the temperate and tropical Atlantic (Casellato et al. 2006), along with some anecdotal reports and pictures of white sea cu- cumbers scattered in the literature (Trefry 2001; Feindel et al. 2011; Benoit 2016). Colour variants, including complete lack of co- lour, in A. japonicus have been described (Yang et al. 2015). The body wall of albino A. japonicus adults contains ~0.24% melanin compared with ~3.12% in normal adults (Zhao et al. 2015). As albinism is known to occur in only a limited number of individ- uals in any given species (0.00001—0.1%; Mouahid et al. 2010), a directed search for these could fuel over- exploitation in some already overfished or vulnera- ble species (Purcell et a/. 2014). Albinism may also affect the behaviour, growth rates, and physiology of sea cucumbers, although these effects remain incom- pletely understood (Lin et al. 2013; Bai et al. 2016; Xia et al. 2017). Orange-footed Sea Cucumber (Cucumaria fron- dosa, Gunnerus 1767) is the most common sea cucu- mber along the eastern Canadian coast, with a distri- bution spanning the North Atlantic, Arctic, and Barents Sea (Paulay and Hansson 2013). A commercial fishery for C. frondosa is rapidly developing in several coun- tries (e.g., Canada, United States, Russia, Greenland, and Iceland), which makes it one of the most important wild sea cucumber fisheries in the world (Therkildsen and Petersen 2006; Nelson eft a/. 2012). As of 2016, market values for C. frondosa ranged from $120—140/ kg (Hansen 2016) up to about $300/kg (Guangzhou market, China; J.-F.H. pers. obs.); aseparate market for unusual colour morphs has not yet developed. 113 ©The Ottawa Field-Naturalists’ Club 114 Here, we describe colour morphs reported for wild C. frondosa populations from northeastern North America. We also report the results of a survey of C. frondosa populations from the Grand Banks of Newfoundland to determine the relative abundance of colour morphs to improve ecological knowledge of this important commercial species. Methods Records of unusually coloured C. frondosa were collated from numerous sources, including specimens collected by hand, SCUBA, and fishing vessels across various locations in eastern Canada (e.g., the estu- ary and Gulf of St. Lawrence, Quebec; Saint-Pierre Bank and southeastern coast of Newfoundland; Fi- gure 1, Table 1). Two batches of sea cucumbers (” = 600) were ex- amined in June and July 2017 to establish the per- centage of orange and white individuals relative to normally coloured ones (dark-brown and grey- brown); no colour standards were used. Samples were obtained from the commercial fishery (trawled using modified scallop gear) in the Grand Banks area of Newfoundland, Canada. No restrictions are placed on animal size, shape, or colour in the fish- ery; thus, harvests are usually a good representation of local populations. Total wet weights of sea cucum- bers, as well as the contracted lengths and widths were measured from fully contracted individuals ac- cording to Singh ef al. (2001). These measurements were taken for all white and orange individuals and from 150 haphazardly selected dark-brown and grey- brown individuals. For white and orange individuals collected in the Grand Banks area of Newfoundland in 2013, two additional measurements were included. 70.000°W 65.000°W Quebec 70.000°W 65.000°W THE CANADIAN FIELD-NATURALIST Vol. 133 Internal organ colours were also compared in pale versus normal individuals. Results Summary of colour morphs in northeastern North America Four main colour variants have been reported for C. frondosa in northeastern North America: dark- brown, grey-brown, orange, and albino (1.e., white and off-white individuals; Figure 2). Dark-brown and grey-brown are the most common colours for this species and, thus, represent normally coloured individuals (Figure 2a,b). The tentacles and tube feet of dark-brown and grey-brown C. frondosa are dark-brown and dark-orange to brown, respectively. Orange-tinged tube feet are common, hence the com- mon name (Paulay and Hansson 2013). Orange and white individuals are the rarest colour morphs and can be described as a gradient of little to no melanin pigment deposition (Figure 2c—g). The tentacles and tube feet of orange and white individuals are typically pale orange and white, respectively. Frequency of colour morphs in Newfoundland trawls All four colour morphs were found off the Grand Banks, Newfoundland (trawled at 40-70 m depth) and in coastal areas near the Avalon Peninsula (collected by SCUBA at <10 m). The survey from the Grand Banks area revealed that orange-coloured individu- als made up 0.5% of the population (7 = 3 in 600), but that white individuals were rarer in that sample (n = 1 in 600, <0.2%). The internal organs (muscle bands, gonad, and intestine) of these white and orange indi- viduals were the same colour as those of dark-brown and grey-brown individuals. Orange individuals were also similar in size to brown individuals (mean wet 60.000°W 60.000°W 55.000°W FicureE 1. Locations in northeastern North America where unusually coloured Orange-footed Sea Cucumbers (Cucumaria Jrondosa) were observed and/or collected. 2019 MONTGOMERY ET AL.: ALBINISM IN ORANGE-FOOTED SEA CUCUMBER WS weight 246 + 87 [SD] g versus 235 + 74 g: Table 2). 2013 (B. Gianasi pers. comm. October 2017), was 78.6 The mean wet weight of the two white individuals, in- +3.4 g (versus mean wet weight of 235 g in normally cluding the one in our 2017 survey and another from _ coloured individuals; Table 2). TABLE 1. Locations in North America where unusually coloured Orange-footed Sea Cucumber (Cucumaria frondosa) occur, based on reports from the literature and the present study. Province or Colour No. (total state, country Region Coordinates observed sampled) Date Source Maine, USA NA NA Orange NA NA Feindel et a/. 2011 NA NA Albino NA NA_ Feindel et a/. 2011 Newfoundland, Fortune Bay, 47.18597933°N, Orange 3 (600) 2017 E. Montgomery (current study) Canada Grand Banks 55.86135864°W 47.18597933°N, Albino 1 (NA) 2017 E. Montgomery (current study) 55.86135864°W St. Pierre 46.67205647°N, Orange 3 (NA) 2017 E. Montgomery (current study) Bank, Grand 55.92315674°W Banks 46.67205647°N, Albino NA 2017 E. Montgomery (current study) 55.92315674°W Unnamed site, 46.89023157°N, Orange 3 (NA) 2013 B. Gianasi (current study) Grand Banks 58.05450439°W 46.89023157°N, Albino 1 (NA) 2013 B. Gianasi (current study) 58.05450439°W Logy Bay 47.6295369°N, Orange NA NA Memorial Field Services 52.66073227°W 47.6295369°N, Albino NA NA Memorial Field Services 52.66073227°W Bay Bulls 47.3048439°N, Orange NA NA Memorial Field Services 52.77677536°W 47.3048439°N, Albino NA NA Memorial Field Services 52.77677536°W Quebec, Canada Mingan 50.02538762°N, Albino NA 1990s _ J.-F. Hamel Archipelago 63.50372314°W Gaspé 49.73513141°N, Albino NA 1990s J.-F. Hamel Peninsula 65.78887939°W Les Escoumins 48.30512072°N, Albino NA 1990s _ J.-F. Hamel 69.24407958°W New Brunswick, Bay of Fundy 44 96479793°N, Orange NA NA. S. Robinson (DFO) Canada 65.80261230°W 44.96479793°N, Albino NA NA_ S. Robinson (DFO) 65.80261230°W Note: DFO = Department of Fisheries and Oceans, NA = data not available. FicurE 2. Colour diversity in adults of Orange-footed Sea Cucumber (Cucumaria frondosa) collected off Newfoundland, eastern Canada. a and b. Typical colours: dark-brown and grey-brown. c to g. Increasing discolouration/albinism: pale brown, orange, pale orange, beige, and white. Sea cucumbers that match c, d, or e are considered “orange”, and those like f and g are considered “white”. Photos were taken with uniform illumination and have not been edited for brightness, con- trast, or colour. Scale bar represents 5 cm for a—f and 3 cm for g. Photos: E. Montgomery. 116 TABLE 2. Size of individuals of Orange-footed Sea Cucum- ber (Cucumaria frondosa) sampled in southeastern New- foundland during the present study. Mean values for orange and white colour morphs are pooled across individuals col- lected in this area, 2013-2017. Mean values for fully pig- mented morphs were obtained from individuals collected in 2017. Mean weight Pisa our Colour morph g+ SD ’ tracted length, cm + SD White (fully albino) 2 787+ 3.4 8.2515 Orange (partly albino) 9 24634871 12.7419 Dark-brown/ 150% 235. 0740: 1ss05 grey-brown (fully pigmented) Discussion Four main colour morphs of the body wall have been reported for C. frondosa from northeastern North America. However, it is important to note that a con- tinuum of degree of pigmentation exists in this spe- cies, ranging from dark-brown to white. White and orange individuals from Newfoundland had normal- ly coloured internal structures and organs, suggest- ing that this species may display leucism (partial or complete loss of pigment), rather than true albinism (no pigment deposition). In fact, some of the dark in- dividuals housed in the laboratory for several months displayed a tendency to bleach over time (B. Gianasi pers. comm. 2017). In the present study, white and orange individu- als were seen and collected in all known populations of C. frondosa sampled in eastern Canada, suggest- ing that loss or decrease of pigmentation is not geo- graphically constrained. An anecdotal reference to unusually coloured individuals is also made in a fish- eries report from Maine (Feindel et a/. 2011), further supporting the suggestion that the phenomenon likely occurs across populations of C. frondosa in North America, northern Europe, and the Arctic, as a gene flow study of So et al. (2011) indicates a supply source along southeastern Newfoundland. Further study will be needed to confirm the factors involved in pigment absence and/or loss in this species, as the phenome- non seems to be geographically widespread. We noted that different colour morphs of C. fron- dosa were collected in Newfoundland from differ- ent depths. Dark-brown sea cucumbers occurred in shallow water <10 m deep (collected by SCUBA) and grey-brown individuals in deeper water (40-70 m; collected by trawl) where less ultraviolet (UV) light penetrates, and weaker pigment protection may be re- quired, as also proposed for A. japonicus (Jiang et al. 2015). Also of note is the fact that white individuals were present in all sampled locations, mixed with the normally coloured morphs (Figure 1). Despite this, it THE CANADIAN FIELD-NATURALIST Vol. 133 remains possible that these individuals may be more susceptible to UV light than normally coloured in- dividuals and may seek cover more readily than darker individuals. In the lower subtidal area of the St. Lawrence Estuary and the Mingan Archipelago of Quebec, pale individuals were reported mainly from discrete areas at the base of rocks or under dense algal cover (J.-F.H. pers. obs.), suggesting that they are ac- tively seeking shelter against exposure to strong light, similar to increased covering behaviour reported in albino urchins in the Caribbean (Kehas et al. 2005). Paler sea cucumbers reported from the Bay of Fun- dy, New Brunswick, did not appear to differ in size from dark-brown and brown individuals (S. Robinson pers. comm. October 2017), which is consistent with the current samples. Although any size difference re- mains to be confirmed with further sampling, white sea cucumbers stand out against the background sub- strate colour, which may generate more stress and, together with decreased UV protection, might explain why they would be generally smaller than normally coloured individuals (Table 2). Their relatively small- er size may also be explained by metabolic factors, as white sea cucumbers have previously been reported as less efficient at protein metabolism than other co- lour morphs (e.g., 4. japonicus, Bai et al. 2016). Our data document the presence of uncommon colour morphs of C. frondosa across most of its east- ern Canadian distribution (Figure 1) and other areas from its general geographic distribution. This de- serves further investigation from both ecological and economical perspectives. Author Contributions Writing — Original Draft: E.M. and T.S.; Writing — Review & Editing: E.M., T.S., J.-F. H., and A.M. Investigation: E.M., T.S., J.-F. H., and A.M.; Formal Analysis: E.M. and T.S.; Funding Acquisition: A.M. Acknowledgements We thank Shawn Robinson (Department of Fish- eries and Oceans Canada) and Bruno Gianasi (Me- morial University) for their data contributions and Jiamin Sun (Memorial University) and Justin So (Amec) for their feedback. We also thank Bill Molloy and the staff of Quin-Sea Fisheries Ltd. for their as- sistance with logistics and animal collections. This research was supported by the Natural Sciences and Engineering Research Council of Canada (grants 311406 and 508323 to A.M). Literature Cited Bai, Y., L. Zhang, S. Xia, S. Liu, X. Ru, Q. Xu, T. Zhang, and H. Yang. 2016. Effects of dietary protein levels on the growth, energy budget, and physiological and im- munological performance of green, white and purple 2019 color morphs of sea cucumber, Apostichopus japonicus. Aquaculture 450: 375-382. https://doi.org/10.1016/j.aqua culture.2015.08.021 Benoit, G. 2016. An albino sea cucumber! Is that possible? Musée du Fjord, La Baie, Quebec, Canada. Accessed January 2019. http://museedufjord.com/en/an-albino- sea-cucumber- is-that-possible. Casellato, S., L. Massiero, and S. Soresi. 2006. Un caso di albinismo in Ocnus planci (Brandt, 1835) (Echinoder- mata: Holothuroidea) nelle “tegnue” dell’Alto Adriatico. Biologia Marina Mediterranea 13: 1059-1062. https:// doi.org/10.7325/galemys.2012.n02 Feindel, S., T. Bennett, and Kk. Kanwit. 2011. The Maine sea cucumber (Cucumaria frondosa) fishery. Depart- ment of Marine Resources, Augusta, Maine, USA. Ac- cessed January 2019. https://www.maine.gov/dmr/science- research/species/cukes/documents/feindeletal2011.pdf. Fernandez-Rivera Melo, F.J., H. Reyes-Bonilla, A. Can- tu, and J. Urias. 2015. First record of albinism in the brown sea cucumber J/sostichopus fuscus in the Gulf of California, Mexico. Marine Biodiversity Records 8: e14. https://doi.org/10.1017/s1755267214001353 Friedman, K., and M. Tekanene. 2005. White teatfish at Kiribati sea cucumber hatchery: local technicians get- ting them out again. SPC Beche-de-Mer Information Bulletin 21: 32-33. Accessed 24 May 2019. https://tiny url.com/y26jnjok. Hansen, J. 2016. Sea cucumber facility in Hackett’s Cove hoping to entice Chinese market. CBC Radio-Canada, Ottawa, Ontario, Canada. Accessed January 2019. https://www.cbc.ca/news/canada/nova-scotia/sea- cucumber-plant-china-market-marine-animal-1.3699286. Jiang, S., S. Dong, Q. Gao, Y. Ren, and F. Wang. 2015. Ef- fects of water depth and substrate color on the growth and body color of the red sea cucumber, Apostichopus ja- ponicus. Chinese Journal of Oceanology and Limnology 33: 616-623. https://doi.org/10.1007/s00343-015-41 78-7 Kehas, A.J., K.A. Theoharides, and J.J. Gilbert. 2005. Effect of sunlight intensity and albinism on the covering response of the Caribbean Sea urchin 7ripneustes ven- tricosus. Marine Biology 146: 1111-1117. https://do1.org/ 10.1007/s00227-004-1514-4 Lin, C., L. Zhang, S. Liu, S. Gao, Q. Xu, and H. Yang. 2013. A comparison of the effects of light intensity on movement and growth of albino and normal sea cucum- bers (Apostichopus japonicus Selenka). Marine and Freshwater Behaviour and Physiology 46: 351-366. https://doi.org/10.1080/10236244.2013.841350 Mouahid, G., R.M. Nguema, M.A. Idris, M.A. Shaban, S. Al Yafee, J. Langand, M. Verdoit-Jarraya, R. Gali- nier, and H. Moné. 2010. High phenotypic frequencies of complete albinism in wild populations of Biomphalaria Dfeifferi (Gastropoda: Pulmonata). Malacologia 53: 161— 166. https://doi.org/10.4002/040.053.0109 Nelson, E.J., B.A. MacDonald, and S.M. Robinson. 2012. A review of the Northern sea cucumber Cucumaria fron- dosa (Gunnerus, 1767) as a potential aquaculture spe- cies. Reviews in Fisheries Science 20: 212-219. Paulay, G., and H. Hansson. 2013. Cucumaria frondosa (Gunnerus, 1767). Jn World Register of Marine Species. Flanders Marine Institute, Ostend, Belgium. Accessed MONTGOMERY ET AL.: ALBINISM IN ORANGE-FOOTED SEA CUCUMBER 117 January 2019. http://www.marinespecies.org/aphia.php ?p=taxdetails&id=124612. Purcell, S.W., B.A. Polidoro, J.-F. Hamel, R.U. Gamboa, and A. Mercier. 2014. The cost of being valuable: pre- dictors of extinction risk in marine invertebrates ex- ploited as luxury seafood. Proceedings of the Royal Society B 281: 20133296. https://doi.org/10.1098/rspb. 2013.3296 Singh, R., B.A. MacDonald, P. Lawton, and M.L.H. Thomas. 2001. The reproductive biology of the dendro- chirote sea cucumber Cucumaria frondosa (Echinoder- mata: Holothuroidea) using new quantitative methods. Invertebrate Reproduction and Development 40: 125— 141. https://doi.org/10.1080/07924259.2001.9652713 So, J.J., S. Uthicke, J.-F. Hamel, and A. Mercier. 2011. Genetic population structure in a commercial marine in- vertebrate with long-lived lecithotrophic larvae: Cucuma- ria frondosa (Echinodermata: Holothuroidea). Marine Biology 158: 859-870. https://doi.org/10.1007/s00227-01 0-1613-3 Therkildsen, N.O., and C.W. Petersen. 2006. A review of the emerging fishery for the sea cucumber Cucumaria frondosa. biology, policy, and future prospects. SPC Beche-de-Mer Information Bulletin 23: 16—25. Accessed 24 May 2019. https://tinyurl.com/yS5mp97wd. Trefry, S. 2001. Parastichopus californicus: Race Rocks Taxonomy, Pearson College, Victoria, British Columbia, Canada. Accessed January 2019. http://www.racerocks. com/racerock/eco/taxalab/saraht.htm. Tse, C. 2015. Albino sea cucumbers, a delicacy, could be- come a lot less rare. New York Times, Aug. 6. Accessed January 2019. https://sinosphere.blogs.nytimes.com/ 2015/08/06/albino-sea-cucumbers-a-delicacy-could- become-a-lot-less-rare/. Wakida-Kusunoki, A., A. Poot-Salazar, and R. J.D. Mena- Loria. 2016. First record of albinism in three-rowed sea cucumber, /sostichopus badionotus. Bulletin of Ma- rine Science 92: 285-290. https://doi.org/10.5343/bms. 2015.1051 WoRMS (World Register of Marine Species). 2019. An Authoritative Classification and Catalogue of Marine Names. Flanders Marine Institute, Ostend, Belgium. http://www.marinespecies.org. Xia, S.D., M. Li, L.B. Zhang, M. Rahman, Q.Z. Xu, L.N. Sun, S.L. Liu, and H.S. Yang. 2017. Growth, im- munity and ammonia excretion of albino and normal Apostichopus japonicus (Selenka) feeding with various experimental diets. Aquaculture Nutrition 2017: 1-9. https://doi.org/10.1111/anu.12646 Yang, H., J.-F. Hamel, and A. Mercier. 2015. The Sea Cu- cumber Apostichopus japonicus: History, Biology and Aquaculture. Academic Press, New York, New York, USA. Zhao, H., C. Muyan, and H. Yang. 2015. Albinism. Pages 211-226 in The Sea Cucumber Apostichopus japonicus: History, Biology and Aquaculture. Edited by H. Yang, J.-F. Hamel, and A. Mercier. Academic Press, New York, New York, USA. Received 9 February 2018 Accepted 22 January 2019 The Canadian Field-Naturalist Wintercresses (Barbarea W.T. Aiton, Brassicaceae) of the Canadian Maritimes CoLIn J. CHAPMAN!’, C. SEAN BLANEY!, and Davip M. MAZEROLLE!? ‘Atlantic Canada Conservation Data Centre, P.O. Box 6416, Sackville, New Brunswick E4L 1G6 Canada ?Current address: Kouchibouguac National Park, 186 Route 117, Kouchibougouac, New Brunswick E4X 2P1 Canada “Corresponding author: colin.chapman@accdc.ca Chapman, C.J.,C.S. Blaney, and D.M. Mazerolle. 2019. Wintercresses (Barbarea W.T. Aiton, Brassicaceae) of the Canadian Maritimes. Canadian Field-Naturalist 133(2): 118-124. http://doi.org/10.22621/cfn.v13312.2235 Abstract We conducted a review of herbarium collections of the Wintercress genus (Barbarea W.T. Aiton) from the Maritime prov- inces. Most specimens previously determined to be the regionally rare native species Erect-fruit Wintercress (Barbarea orthoceras Ledebour) are in fact the uncommon exotic Small-flowered Wintercress (Barbarea stricta Andrzejowski). The latter species is here reported as new to Atlantic Canada, where it is scattered but widespread in the three Maritime prov- inces. Only three collections (two from New Brunswick and one from Nova Scotia) were confirmed as B. orthoceras. Its known range extent and area of occupancy in the Maritimes has been significantly revised, and B. orthoceras 1s now con- sidered potentially extirpated in New Brunswick and extremely rare in Nova Scotia. One collection from Nova Scotia was referred to another rare exotic species, Early Wintercress (Barbarea verna (Miller) Ascherson), which represents the first record for the Maritimes. Key words: Cruciferae; Brassicaceae; new record; floristics; Barbarea stricta, Barbarea orthoceras, Barbarea verna, con- servation; Maritimes; Canada; wintercress Introduction The wintercress genus (Barbarea W.T. Aiton) has long been a source of confusion in North America, in part due to taxonomy and to somewhat variable species with overlapping morphology (Fernald 1909; Mulligan 2002; Al-Shehbaz 2010). The sole native North American member of the genus, Erect-fruit Wintercress (Barbarea orthoceras Ledebour), was at One point considered a native form of the Eurasian spe- cies Bitter Wintercress (Barbarea vulgaris W.T. Aiton) or Small-flowered Wintercress (Barbarea stricta An- drzejowski; discussed in Fernald 1909). Fernald (1909), who concluded reports of B. stricta were mis- identified individuals of B. vulgaris with appressed fruit, excluded the former species from the North American flora. Mulligan (1978) reported the first con- firmed North American records of B. stricta based on specimens collected from Quebec in 1944, although the species was not included in The Flora of Canada (Scoggan 1978). Barbarea stricta has subsequently been documented in Ontario (based on a 1922 speci- men; Dorofeev 1998), Colorado, Connecticut, Maine, Michigan, Massachusetts, New Hampshire, New York, Rhode Island, Vermont, and Wisconsin (AI- Shehbaz 2010). Prior to our study, only two species of Barbarea were listed in the flora of the Canadian Maritimes (New Brunswick [NB], Nova Scotia [NS], and Prince Edward Island [PEI]): B. vulgaris and B. orthoceras (Roland and Smith 1969; Scoggan 1978; Zinck 1998; Hinds 2000; Munro ef al. 2014). Barbarea vulgaris is a common and widespread weedy species of Eurasian origin, well documented from throughout NB, NS, and PEI (Erskine 1960; Roland and Smith 1969; Zinck 1998; Hinds 2000; AC CDC 2019; Figure 1). Barbarea orthoceras is na- tive to boreal North America and eastern and cen- tral Asia (Al-Shehbaz 2010). Haines (2011) con- siders B. orthoceras a calciphile associated in New England with high-pH bedrock or till, although a var- iety of habitats, including grasslands, forests, boggy ground, and railroad embankments are reportedly suitable (Al-Shehbaz 2010). Though relatively se- cure in the western and northern portion of its range, B. orthoceras 1s rare in eastern North America and is of conservation concern in all jurisdictions of oc- currence east of Ontario (NatureServe 2017). As of 2000, it was reported in the Maritimes from five NB locations on “stream banks, sandy beaches, gravel river strands, and rocky shores” (Hinds 2000: 225). An additional nine NB collections and two NS col- 118 ©The Ottawa Field-Naturalists’ Club 2019 lections initially identified as B. orthoceras were de- posited at regional herbaria between 2001 and 2015 (AC CDC 2019; Figure 2), but in 2015 D.M.M. sus- pected some Maritimes records involved B. stricta, so we undertook a thorough specimen review to deter- mine the regional status of B. orthoceras, B. stricta, and B. vulgaris. Methods Approximately 170 specimens from four regional herbaria (E.C. Smith Herbarium—Acadia University [ACAD], New Brunswick Museum [NBM], Nova Scotia Museum of Natural History [NSPM], Connell Memorial Herbarium—wUniversity of New Bruns- wick [UNB]) and one national herbarium (Canadian Museum of Nature [CAN]) were examined by C.J.C. in 2018. Specimens from Agriculture and Agri-Food Canada’s National Collection of Vascular Plants [ DAO] were unavailable for examination due to facil- ity renovations. Plants were determined based on the treatments in Al-Shehbaz (2010) and Haines (2011). A simplified key is presented here: CHAPMAN ET AL.: MARITIME BARBAREA 119 la. Stylar beaks narrow, longer than 1.5 mm (Fi- gure S1); auricles of distal leaves glabrous..... Lae Mth ARAL AR AAR AT NG AAAS ie B. vulgaris 1b. Stylar beaks stout, less than 1.5 mm long (Fi- gure S2); auricles of distal leaves at least SMarSely-callaee 7s. the asadusa stots noedas trees taest 2 2a. Uppermost leaves dentate (Figure S3); petals less than 4.5 mm long (Figure S4); fruit mostly shorter than 28 mm long................00.0... B. stricta 2b. Uppermost leaves pinnatifid (Figure S5); pe- tals greater than 5 mm long; fruit mostly oOrealer than SIMI Ong ekki. 3 3a. Basal leaves with 1—4 pairs lateral lobes; fruit usually under 40 mm long; fruiting pedicels narrower than fruit 0.0.0.0... B. orthoceras 3b. Basal leaves with 4—10 pairs lateral lobes; fruit usually greater than 53 mm long; fruiting pedicels as broad as fruit ....0..00c. B. verna Clear determinations could be made for most spec- imens, but some collections presented conflicting morphology, as is mentioned of NB (Hinds 2000), New England (Al-Shehbaz 2010), and Michigan (Voss 0 100 200 km Ficur 1. Distribution of Bitter Wintercress (Barbarea vulgaris) in the Canadian Maritimes based on specimens (solid circles) and Atlantic Canada Conservation Data Centre sight records (hollow circles) determined and verified during the present study. 120 and Reznicek 2012) material. We sent a small sub- set of six specimens to Ihsan Al-Shehbaz, Missouri Botanical Garden, and he confirmed all six as the species initially determined by C.J.C. All records in this paper are either supported by voucher specimens or are photographic or sight re- cords made by Atlantic Canada Conservation Data Centre (AC CDC) botanists. Results and Discussion Barbarea vulgaris W.T. Aiton Barbarea vulgaris remains the most frequently encountered and widespread species of Barbarea in the Maritimes (Figure 1). As a weedy species, B. vul- garis is found in a variety of anthropogenic habitats such as fields and roadsides (Erskine 1960; Roland and Smith 1969; Hinds 2000). It is also frequent on river shores, where it can co-occur with B. stricta and potentially with B. orthoceras. The taxon is morph- ologically variable, and though many varieties have been described (treated in Fernald 1909), none are presently recognized in North America (Al-Shehbaz 2010). Some specimens present mixed or intermedi- ate morphology in style length, uppermost leaf shape, and auricle pubescence. Hinds (2000) reported ap- THE CANADIAN FIELD-NATURALIST A, 4 =55/ A Specimens Vol. 133 parently intermediate NB specimens and suspected hybridization might be involved. Hybrids of B. vul- garis and B. stricta (= B. x schulzeana Haussknecht) are recorded for Europe, although occurrences are very infrequently recorded and highly sterile (Rich 1987). If hybrids were also sterile in the Maritimes, we might expect a lower frequency of plants inter- mediate between B. vulgaris and B. orthoceras than has been observed. These intermediates may thus be morphological extremes of the highly variable B. vulgaris. Barbarea orthoceras Ledebour We found that only one specimen initially identi- fied as B. orthoceras was determined correctly. The remainder were reassigned to B. stricta or in very few cases B. vulgaris. An additional B. orthoceras rec- ord was discovered upon redetermination of a speci- men originally identified as B. vulgaris. The two Specimens now confirmed for NB were collected in 1944 along water-runs in an old pasture on Grand Manan Island (C.A. Weatherby & Una F. Weatherby 7343), and in 1964 on a roadside at the edge of a Black Spruce (Picea mariana (Miller) Britton, Sterns & Poggenburgh) forest in Kings County (P.R. Roberts = @ Verified B. orthoceras redetermined O Maine B. orthoceras FiGure 2. Distribution of the rare native Erect-fruit Wintercress (Barbarea orthoceras) in New Brunswick (NB), Nova Scotia (NS), and Prince Edward Island (PE) based on specimens determined and verified during the present study. Historical records from Maine provided by the Maine Natural Heritage Program were not verified, but one Maine collection at the New Brunswick Museum was redetermined as B. orthoceras. 2019 & N. Bateman 64-361). C.S.B. discovered the first and only known NS population on a steep, seepy rav- ine slope under shrubs in northern Cape Breton in 2016 (collection number 8978). Barbarea orthoceras was first reported for NB based on two specimens revised by H.J. Scoggan in 1955 (R. Chalmers 305a [now determined as B. stricta] and C.A. Weatherby and Una F: Weatherby 7343). Scoggan appears unlikely to have considered B. strictaas a possible identity because it was not con- firmed in North America until much later (Mulligan 1978). Indeed, his key in Scoggan (1978) describes the petals in B. orthoceras as “at most 5 mm long”, a key character of B. stricta (Al-Shehbaz 2010). Hinds (2000) similarly described the petals of B. orthoceras as “less than 5 mm long”. In fact, the petals are 5—7 mm long in B. orthoceras, which is the most reliable character separating it from B. stricta (I. Al-Shehbaz pers. comm. 8 May 2018). One historical collection of B. orthoceras (initially misidentified as B. vulgaris and later as B. stricta) was uncovered from Fort Kent, Maine (G.U. Hay, s.n.), along the Saint John River across from Madawaska County, NB. The species is known from two addi- tional historical records in northern Maine, however all recent collections of potential B. orthoceras in the state have turned out to be B. stricta (L. St. Hilaire and D. Cameron pers. comm. 20 February 2019). All re- cent records from extensive AC CDC fieldwork along northern NB rivers have also been B. stricta, sug- gesting that, if B. orthoceras is present on rivers in the region, it is quite rare. However, the now-confirmed records of B. orthoceras from pasture and roadsides in NB suggest it could be overlooked in disturbed sites because of assumptions that Barbarea in ruderal habitats must be B. vulgaris or B. stricta, and because botanists tend to spend less time in ruderal habitats. Speculation aside, our study greatly decreases the known range extent and area of occupancy for B. orthoceras in the Maritime provinces (Figure 2). No recent records exist in NB, where its provincial status has been changed from imperilled/vulnerable (S2S3) to possibly extirpated (SH). It has been confirmed as extremely rare in NS and remains unknown on PEI. Barbarea stricta Andrzejowski This study confirmed the presence of B. stricta in Atlantic Canada, where it is scattered but widely dis- tributed in NB, NS, and PEI (Figure 3). Haines (2011) and Cayouette (1984) describe B. stricta as having in- vaded river shores, lake shores, and wet, disturbed areas in New England, and Quebec respectively, and it has mostly been collected from similar habitats in the Maritimes. Many B. stricta specimens examined in this study had morphology that was at the larger ex- tremes for the species (as also reported in Al-Shehbaz CHAPMAN ET AL.: MARITIME BARBAREA 121 2010), which is potentially suggestive of genetic in- fluence from the larger B. orthoceras. The morphol- ogy of these plants might also be explained by the founder principle (Mayr 1942) if North American populations happened to have been founded by un- usually large individuals. The morphological sim- ilarity of B. orthoceras and B. stricta would make hybridization difficult to demonstrate without mo- lecular investigation. We failed to locate historical specimens of B. stricta in NS or PEI, suggesting it may have dis- persed more recently to these provinces. However, it was introduced in NB as early as 1877 (R. Chalmers 305a;, now the earliest Canadian record), where col- lections were misidentified as B. orthoceras, or more rarely as B. vulgaris. Early introduction of B. stricta and overlapping habitat requirements of Barbarea species suggests ample opportunity for hybridization in NB. However, because Hinds (2000) appears not to have considered B. stricta as a possibility in iden- tifying NB material, his specimens of “intermediate morphology” between B. vulgaris and B. orthoceras may simply have represented B. stricta. Barbarea verna (Miller) Ascherson We referred one collection from Kentville, NS (WS. Erskine s.n.) to Early Wintercress (Barbarea verna (Miller) Ascherson). Originally identified as B. vul- garis, it was distinguished by its leaves with five to seven pairs of lateral lobes, pinnatifid uppermost leaves, and conspicuously ciliate leaf auricles (Al- Shehbaz 2010; Haines 2011). In fruit, B. verna can also be distinguished by its large fruit (5.3—7 cm), and pedicels as broad as the fruit they subtend (AI- Shehbaz 2010). This European species is cultivated as a salad plant in North America, where it escapes to disturbed habitats such as fields and meadows (Mulligan 2002; Haines 2011). This represents the first Maritimes record of this rarely reported introduction, otherwise known in Canada only from Newfoundland and British Columbia (Brouillet et a/. 2010+). Voucher specimens Barbarea orthoceras Ledebour—NEW BRUNS- WICK: Charlotte Co., Grand Manan, Between Long Pond & Red Pt., weed along water-runs in old pasture, 6 August 1944, C.A. Weatherby and U.F. Weatherby 7343 (CAN); Kings Co., Lower Millstream, roadside at Black Spruce forest edge, 28 May 1964, P.R. Rob- erts & N. Bateman 64-361 (UNB); NOVA SCOTIA: Inverness Co., Pleasant Bay, Lower Delaneys Brook, 46.966519°N, 60.654339°W, steep, seepy ravine slope under shrubs, not seen elsewhere between High Capes & Delaneys Point, with Red-osier Dogwood (Cornus sericea L.), Tall Meadow-rue (Thalictrum pubescens Pursh), 14 July 2016, C.S. Blaney 8978 (NSPM); 122 THE CANADIAN FIELD-NATURALIST Vol. 133 100 FicurE 3. Distribution of the uncommon exotic species Small-flowered Wintercress (Barbarea stricta) in the Canadian Maritimes based on specimens (solid circles) and Atlantic Canada Conservation Data Centre photographic or sight rec- ords (hollow circles; the western sample is shown in Figure S3) determined, verified, or revised during the present study. MAINE: Aroostook Co., Fort Kent, 8 July 1904, G.U. Hay s.n. (NBM). Barbarea stricta Andrzejowski—NEW BRUNS- WICK: Madawaska Co., St. Francis River, 47.182855°N, 68.896639°W, wet mud, in grassy meadow along shore of floodplain island, 11 July 2013, D.M. Mazerolle DM4032 (NBM, UNB, DAO); Madawaska Co., St. Francis River, 47.285603°N, 69.050753°W, sandy riverbank with sparse al- ders, 9 July 2013, C.S. Blaney & D.M. Mazerolle 8267 (NBM), Madawaska Co., St. Francis River, 47.276528°N, 69.049497°W, sandy-bouldery shore under Eastern White Cedar (Thuja occidentalis L.), 9 July 2013, C.S. Blaney 8268 (NBM); Madawaska Co., St. Francis River, 47.26046°N, 69.050219°W, silty sand-gravel rivershore, 9 July 2013, C.S. Blaney 8270 (NBM), Restigouche Co., Campbellton, 20 July 1877, R. Chalmers 305a (CAN), Victoria Co., Grand Falls, 47.05527°N, 67.759561°W, 31 July 2018, D.M. Mazerolle DM7808 (NBM); Restigouche Co., Heron Island, 47.9823°N, 66.11649°W, sandy hummocks, 29 September 1982, H.R. Hinds 5798 (UNB), Res- tigouche Co., Eel River, 48.02520°N, 66.39349°W, gravel shore, 5 August 1982, H.R. Hinds 5400 (UNB), Restigouche Co., Morrissey Rock, 47.983327°N, 66.832622°W, alongside railway tracks, 10 Au- gust 1965, P.R. Roberts & B. Pugh 65-5384 (UNB); Restigouche Co., Restigouche River, 47.990321°N, 66.769302°W, sandy floodplain terrace above river bank, on island in upper estuary near head of tide, 18 September 2013, D.M. Mazerolle DM4143 (NBM); Restigouche Co., Tidehead, 47.985532°N, 66.767721°W, silty bar in fresh tidal zone, 18 Sep- tember 2013, C.S. Blaney 8485 (NBM), Restigouche Co., Eel River, 48.023715°N, 66.412898°W, gramin- oid floodplain meadow on elevated terrace along brackish tidal river, 26 August 2015, D.M. Mazerolle DM6967 (NBM, UNB); Restigouche Co., Upsalquitch River, 47.883333°N, 66.951000°W, 12 July 1957, E.C. Smith 16312 (ACAD), Victoria Co., Arthurette, 46.81718°N, 67.43563°W, gravelly north shore, 27 July 1982, H.R. Hinds 5361 (UNB), Victoria Co., Grand Falls, 47.059342°N, 67.778800°W, 31 July 2018, C.J. Chapman 1152 (NBM), Victoria Co., 2019 Grand Falls, 47.064978°N, 67.788144°W, 31 July 2018, C.J. Chapman 1158 (UNB); Northumber- land Co., Upper Blackville Bridge, 46.629606°N, 65.865519°W, eroding sandy-gravel shore slope, 14 August 2007, C.S. Blaney & D. Whittam 6646 (UNB); Northumberland Co., Beaubears Island, Muirami- chi, 46.978856°N, 65.561207°W, On open backshore of east point of island, 7 September 2005, D. Mc- Leod & C. Merrithew 5271 (UNB); Carleton Co., Big Presque Isle Stream, 46.424062°N, 67.703127°W, moist depression in densely vegetated weedy Bal- sam Poplar (Populus balsamifera L.)/Black Ash (Fraxinus nigra Marshall) floodplain, 7 July 2015, D.M. Mazerolle DM6711 (NBM, UNB); Kent Co., Kouchibouguac NP, 46.86312°N, 64.95513°W, gar- bage dump, 11 June 1977, B. Lyons & D. LaFon- taine 564 (UNB, ACAD), Kings Co., Hatfield Point, 45.63523°N, 65.88335°W, rocky and gravelly shore, 28 June 1980, H.R. Hinds 3234 (UNB); Kings Co., Ham- mond River, 45.471532°N, 65.9056750°W, rich soil in understorey of Silver Maple (Acer saccharinum L.) floodplain forest, on floodplain island, 23 August 2017, D.M. Mazerolle DM7598 (NBM); Westmor- land Co., North River, 46.038486°N, 65.138669°W, rich, silty upper river bank, 19 July 2017, CS. Blaney 9148 (NBM); Westmorland Co., North River, 46.041878°N, 65.135242°W, silty river bank, 19 July 2017, C.S. Blaney 9153 (NBM); Westmorland Co., Petitcodiac River, 45.949465°N, 65.166692°W, muddy gravel river bar, 8 September 2017, C.S. Blaney 9198 (NBM); NOVA SCOTIA: Antigonish Co., Antig- onish Harbour, 45.659103°N, 61.895856°W, open, cattle-grazed White Spruce (Picea glauca (Moench) Voss) forest on gypsum at edge of saltmarsh, 30 June 2014, C.S. Blaney 8575 (ACAD), Antigonish Co., Antigonish Harbour, 45.658474°N, 61.899508°W, muddy edge of brackish marsh, 30 June 2014, C.S. Blaney 8578 (ACAD, NSPM, DAO, UNB); Inver- ness Co., Glenora, 45.742390°N, 61.293412°W, edge of rich shrubby floodplain terrace, 31 July 2015, D.M. Mazerolle DM6879 (ACAD, NSPM, DAO); PRINCE EDWARD ISLAND): Prince Co., Miminegash River, 46.860973°N, 64.169513°W, forb and graminoid floodplain meadow, 28 July 2017, D.M. Mazerolle DM/7483 (ACAD, DAO). Barbarea verna (Miller) Ascherson—NOVA SCO- TIA: Kings Co., Kentville, sandy grassy slope, 22 May 1950, JS. Erskine s.n. (NSPM). Author Contributions Writing—Original Draft: C.J.C.; Writing—Re- view & Editing: C.S.B., D.M.M., and C.J.C.; Concep- tualization: C.S.B. and D.M.M.,; Investigation: C.J.C.; Methodology: C.J.C.; Funding Acquisition: C.S.B., D.M.M., and C.J.C. CHAPMAN ET AL.: MARITIME BARBAREA 123 Acknowledgements We thank Ihsan Al-Shehbaz, Missouri Botanical Garden, for discussing morphological variation and for verifying a subset of specimens. Thank you to herbarium and research staff for making available Barbarea specimens: Alain Belliveau (ACAD, Acadia University), Stephen Clayden and Amanda Bremner (NBM, New Brunswick Museum), Katherine Ogden (NSPM, Nova Scotia Museum), Robyn Shortt (UNB, University of New Brunswick), and Jennifer Doubt, Lyndsey Sharp, and Paul Sokoloff (CAN, National Herbarium of Canada, Canadian Museum of Nature). Thank you to Lisa St. Hilaire and Don Cameron of the Maine Natural Areas Program for their comments on B. orthoceras in Maine, and to Mike Oldham, Jim Pringle, and Paul Catling for helpful review of the manuscript. Fieldwork that generated the Atlantic Canada Conservation Data Centre specimens exam- ined in this study was supported primarily by the New Brunswick Wildlife Trust Fund, the Nova Scotia Crown Share Land Legacy Trust, Environment and Climate Change Canada’s Atlantic Ecosystem Initi- atives program, and the Nova Scotia Department of Natural Resources (now Lands and Forestry). Literature Cited AC CDC (Atlantic Canada Conservation Data Centre). 2019. Digital database of species occurrence records for the Maritime provinces. Atlantic Canada Conservation Data Centre, Sackville, New Brunswick, Canada. Al-Shehbaz, I.A. 2010. Barbarea. Pages 460—463 in Flora of North America North of Mexico. Volume 7. Edited by Flora of North America Editorial Committee. Flora of North America Association, New York and Oxford. Brouillet, L., F. Coursol, M. Favreau, M. Anions, P. Bélisle, and P. Desmet. 2010+. VASCAN, the database of vascular plants of Canada. Accessed 10 May 2018. http://data.canadensys.net/vascan. Cayouette, J. 1984. Nouvelles stations du Barbarea stricta Andrz. au Québec. Naturaliste canadien 111: 207—209. Dorofeey, V.I. 1998. The four new species of Brassicaceae for North America. Botanicheskii Zhurnal 83: 133-135. Erskine, D.S. 1960. The Plants of Prince Edward Island. Plant Research Institute, Research Branch, Canada De- partment of Agriculture. Publication 1088. https://doi. org/10.5962/bhI.title.53729 Fernald, M.L. 1909. The North American species of Bar- barea. Rhodora 11: 134-141. Haines, A. 2011. Flora Novae Angliae, A Manual for the Identification of Native and Naturalized Higher Vas- cular Plants of New England. Yale University Press, New Haven, Connecticut, USA. Hinds, H.R. 2000. Flora of New Brunswick, Second Edi- tion: A Manual for Identification of the Vascular Plants of New Brunswick. Department of Biology, University of New Brunswick, Fredericton, New Brunswick, Canada. Mayr, E. 1942. Systematics and the Origin of Species. Co- lumbia University Press, New York, USA. 124 Mulligan, G.A. 1978. Barbarea stricta Andrz., a new intro- duction to Quebec. Naturaliste canadien 105: 298-299. Mulligan, G.A. 2002. Weedy introduced mustards (Brassic- aceae) of Canada. Canadian Field-Naturalist 116: 623— 631. Accessed 12 October 2019. https://biodiversitylibrary. org/page/35151869. Munro, M.C., R.E. Newell, and N.M. Hill. 2014. Nova Scotia Plants: 3-14 Brassicaceae, mustard family. Nova Scotia Museum, Halifax, Nova Scotia. Accessed 10 May 2018. https://ojs.library.dal.ca/NSM/article/view/5478. NatureServe. 2017. NatureServe Explorer: an online en- cyclopedia of life. Version 7.1. NatureServe, Arlington, Virginia, USA. Accessed 10 May 2018. http://explorer. natureserve.org. Rich, T.C.G. 1987. The genus Barbarea R.Br. (Cruciferae) SUPPLEMENTARY MATERIAL: THE CANADIAN FIELD-NATURALIST Vol. 133 in Britain and Ireland. Watsonia 16: 389-396. Roland, A.E., and E.C. Smith. 1969. The Flora of Nova Sco- tia. Nova Scotia Museum, Halifax, Nova Scotia, Canada. Scoggan, H.J. 1978. The Flora of Canada, Part 3: Dicotyle- donae (Saururaceae to Violaceae). National Museums of Canada, Ottawa, Canada. https://doi.org/10.5962/bh1 title. 122890 Voss, E.G., and A.A. Reznicek. 2012. Field Manual of Mi- chigan Flora. University of Michigan Press, Ann Arbor, Michigan, USA. https://doi.org/10.3998/mpub.345399 Zinck, M. 1998. Roland’s Flora of Nova Scotia. Nimbus Publishing, Halifax, Nova Scotia, Canada. Received 28 March 2019 Accepted 8 August 2019 Ficure S1. Bitter Wintercress (Barbarea vulgaris), with characteristic slender, relatively long stylar beaks (>1.5 mm long). FiGuRE 82. Small-flowered Wintercress (Barbarea stricta), with short and stout stylar beaks (<1.5 mm long) and appressed siliques. FiGurE S83. Small-flowered Wintercress (Barbarea stricta), with dentate upper stem leaves and ciliate auricles. Figure S4. Small-flowered Wintercress (Barbarea stricta), with relatively short petals (<<4.5 mm long). Figur S5. Large-fruit Wintercress (Barbarea orthoceras), with pinnatifid uppermost stem leaves. Specimen: National Her- barium of Canada, Canadian Museum of Nature (CAN 60422). The Canadian Field-Naturalist Note Summer movements of a radio-tagged Hoary Bat (Lasiurus cinereus) captured in southwestern Ontario DEREK MORNINGSTAR" and AL SANDILANDS” 'Myotistar, 51 Silverthorne Drive, Cambridge, Ontario N3C 0B4 Canada 71356 Lockie Road, Branchton, Ontario NOB 1L0 Canada *Corresponding author: myotistar@gmail.com Morningstar, D., and A. Sandilands. 2019. Summer movements of a radio-tagged Hoary Bat (Lasiurus cinereus) captured in southwestern Ontario. Canadian Field-Naturalist 133(2): 125-129. https://doi.org/10.22621/cfn.v13312.2148 Abstract Hoary Bat (Lasiurus cinereus) is a migratory species known to travel long distances during migration. Little is known about its movement patterns during other periods. An adult male Hoary Bat that we radio-tagged in southwestern Ontario in sum- mer was tracked using the Motus Wildlife Tracking System. It travelled a minimum of 827 km in a circular route over a 2-week period and was last recorded 46 km from the original capture site. Hoary Bat is highly vulnerable to being killed at wind turbines and its propensity to travel great distances during summer and migration may exacerbate the impacts of wind farms. Key words: Hoary Bat; Lasiurus cinereus, summer movement; Motus; radio-tracking; southwestern Ontario Hoary Bat (Lasiurus cinereus) is widely distributed throughout the Western Hemisphere, and may be com- mon in the Great Lakes Region. However, it is among the least frequently encountered species in studies of regional bat communities (e.g., Jung et al. 1999). Hoary Bat travels long distances through its life cycle (Cryan et al. 2014). Migratory movements of this species have mostly been inferred from the sea- sonal distribution of museum specimens (Cryan 2003; Cryan et al. 2014), but there are several biases in these data. Kurta (2010) demonstrated that stud- ies based on museum specimens did not match actual distribution and sex ratios of Hoary Bats captured by mist-netting in Michigan. This species is frequently studied through acoustic inventories (e.g., Barclay er al. 1999) and mortality studies at wind power facili- ties (e.g., Kunz et al. 2007). Such studies provide no information on individual movement patterns, which can only be determined through physical handling and tracking of individuals. Only one study has docu- mented the long-distance movements of Hoary Bats in North America (Weller et a/. 2016), and it was con- ducted in autumn rather than summer. The non-migratory movements of males have not been well documented. Banfield (1974) stated that males seem to wander erratically during spring and summer and do not associate with females while the latter are caring for their young. Here we provide data on the short-term movements of a radio-tracked male Hoary Bat during mid-summer. On 9 July 2016, we captured an adult male Hoary Bat in a 12-m triple high mist net near a Little Brown Myotis (Myotis /ucifugus) maternal roost near Branch- ton, Ontario (43.2986°N, 80.2900°W). The Hoary Bat was 29.2 g at the time of capture and had a fore- arm length of 51 mm. A Nanotag radio transmitter (Lotek, Newmarket, Ontario, Canada) was affixed to the back of the bat by shaving a small area and apply- ing a small amount of Osto-Bond (Montreal Ostomy Inc., Vaudreuil-Dorion, Quebec, Canada) glue to the bat and the tag. The mass of the tag reported by the manufacturer was 0.33 g, representing 1.1% of the body mass of the bat, less than the maximum of 5% recommended by Aldridge and Brigham (1988). This brand of skin cement is effective for adhesion of the tag to the bat for at least several days (Carter et al. 2009), and the transmitter battery life is expected to be as long as 21 days. Once the glue had dried, the bat was released at the capture site. We attempted to relocate the bat by driving roads within 5 km of the capture site for seven days after capture, using two 4-element Yagi antennae fixed in opposite directions to the roof of a truck and con- nected to an SRX800 receiver (Lotek). The bat was A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 125 ©The Ottawa Field-Naturalists’ Club 126 never detected from the ground; thus, tracking its movements relied on detections on the Motus Wild- life Tracking System. The Motus system is an international collabor- ative research network that uses coordinated auto- mated radio telemetry arrays to study movements of small animals (Taylor et a/. 2017); it has been used for tracking migratory bats (Lagerveld et al. 2017). Bird Studies Canada maintains an array of more than 100 automated radio telemetry stations in Ontario, and The Ohio State University maintains other sta- tions in Ohio used by this study. Although these do not provide complete coverage of all areas and are not suitable for precise triangulation, detections show large landscape movements. Detection of tags at a re- ceiver station could be within 15 km of the station de- pending on station strength and whether the animal is flying in the open or near obstructions (Taylor er MICHIGAN Detroit Windsor \— , “x THE CANADIAN FIELD-NATURALIST “London Vol. 133 al. 2017). The nearest receiver station to the capture site (Onondaga) was ~2 km south, but it was not acti- vated until 12 July 2016 (seven days after we tagged the bat); the bat was never detected at this station. Figure 1 depicts the movements of the bat, al- though it did not necessarily travel in a straight line between stations. The specific locations of the sta- tions where it was detected are provided in Table 1 along with the dates and times when the bat was de- tected. Over two weeks, the Hoary Bat travelled a minimum distance of 827 km and was last detected only 46 km from the banding location. The landscape that it moved through was predominantly agricultural with scattered remnants of forest and wetland. The male Hoary Bat made significant movements within short periods (Table 1): the longest were a minimum of 253 km over one night and 316 km over three nights (although it is unknown how long the A Toronto ) 9, Formosa ee % ~ ¢ 9 10, Conestogo Waterloo. ly »— Tag Location, / @North Dumfries , ' i Hamilton | 2 11, Curries r | sye@ 1, Waterford Quarry 2, Falconer 3, Falconer 4, Falconer - oo * on *, oo SIs, Ottawa Weir * 6, Stangee Seo aXe, Cleveland SEE INSET MAP INSET MAP 7, Winous Point FiGurE 1. The map indicates the location where the male Hoary Bat (Lasiurus cinereus) was tagged on 9 July 2016 (North Dumfries) and its subsequent detections at the various Motus stations: 1. Waterford Quarry; 2, 3, 4. Falconer; 5. Ottawa Weir; 6. Stangee; 7. Winous Point; 8. Cedar Point; 9. Formosa; 10. Conestogo; 11. Curries. The inset map shows its move- ments in Ohio on 19 July 2016. 2019 MORNINGSTAR AND SANDILANDS: SUMMER MOVEMENTS OF A HOARY BAT IZ, TABLE 1. Movements of a radio-tagged Hoary Bat (Lasiurus cinereus) over a 2-week period in July 2016 in southwestern Ontario. Motus station* North Dumfries (tagging location) 1. Waterford Quarry 9 July, 2350 14 July, 1943-2200 2. Falconer 15 July, 2229-2303 3. Falconer 16 July, 2005 4. Falconer 18 July, 1912-1929 5. Ottawa Weir 19 July, 1728-1730 6. Stangee 19 July, 1732-1743 7. Winous Point 8. Cedar Point 18 July, 1731-1736 19 July, 1743-1744 9. Formosa 22 July, 1954-1955 10. Conestogo 22 July, 2129-2133 11. Curries 23 July, 2229-2349 24 July, 0156-0201 *Numbers correspond to those on the map in Figure 1. bat took to fly these distances). The movements were not in a clear latitudinal direction. The bat initially moved to the southwest along or over Lake Erie, then flew quickly back to the northeast along or over Lake Huron and started another trip toward the southwest. It spent some time north of the Long Point area at Falconer and in the Cedar Point area of Ohio and was last detected at the Curries station (also near Long Point). The bat was detected consistently early in the night and often during daylight hours. If the bat was not moving at this time, we would have expected it to be detected repeatedly at individual stations until it moved out of the detection range. The bat did not ap- pear to return to the location where it was captured during our sample period, because it was never de- tected at the Onondaga or other nearby stations. Our study reports the first documented move- ments of an adult male Hoary Bat during the summer months. It has been widely believed that Hoary Bat makes long-distance movements during migration, but concrete evidence of this is sparse; even less 1n- formation is available on local summer movements, such as those documented here. The few recoveries of banded Hoary Bats show maximum distances be- tween banding and recovery sites of 150—450 km, representing more local movements (Davis 1969, 1970), although isotope analysis has shown that Hoary Bats make long-distance seasonal movements (Arias 2014; Baerwald et al. 2014). In autumn in California, Weller et a/. (2016) re- covered three adult male Hoary Bats with global pos- itioning system tags. One remained sedentary during the study, one made local movements of less than 100 km, and the third travelled over 1000 km in a month. Similar to the bat that we tracked, it moved in a cir- cular manner and ended up less than 150 km from the original capture location. Results from this observa- Date and time detected Cumulative distance travelled, km Distance from last station, km 0 0 43 43 25, 68 0 68 0 68 253 321 3 324 Js) 349 ao 388 316 704 56 760 67 827 tion and Weller et a/. (2016) indicate that male Hoary Bats may occasionally make circular or other long- distance movements lasting several days and cover- ing distances as great as 1000 km. Although sample size 1s still very small (two of four tracked bats), this indicates that these bats have a large home range and, therefore, habitat protection or conservation for the species must similarly be on a large scale. The pur- pose of these flights is uncertain: bats may be search- ing for females to mate with before migration (Weller et al. 2016), although this bat was not scrotal (show- ing signs of sexual reproduction by having distended testes). More studies that use the Motus system or other methods capable of tracking Hoary Bats are recommended to better understand the space use by this fast, high-flying bat. Hoary Bats are frequently killed at wind power facilities, particularly during late summer and au- tumn migration (Arnett et al. 2007; Bird Studies Canada et al. 2016). Hayes et al. (2015) concluded that Hoary Bats make up approximately 40-50% of all bat mortalities at wind farms; in Canada, this fig- ure is 30.9% (Bird Studies Canada et al. 2016). Cryan (2011) estimated that as many as 225 000 Hoary Bats might be killed annually at North American wind farms, and it is predicted that the Hoary Bat popu- lation could decline by 90% in the next 50 years be- cause of mortality related to wind turbines (Frick ef al. 2017). Over the landscape that this bat travelled, a considerable number of wind turbines are in oper- ation. Ontario has the greatest wind power genera- tion in Canada with more than 4781 MWh of annual power production as of 2016 (Canadian Wind Energy Association 2017); many of the turbines are in south- western Ontario. This bat either avoided those tur- bines or flew through them without being killed dur- ing the period that it was tracked. Our data provide 128 evidence that Hoary Bats are at risk of encountering a large number of wind turbines during their summer movements, not just during migration. Acknowledgements This study was funded in part by the Canadian Wildlife Service. We thank Stu Mackenzie for his thoughtful review and insight on an earlier draft. Field assistance was provided by Lucas Greville, Luke Owens, Roberto Valdizon, Alejandra Ceballos- Vasquez, Chris Risley, and Patricia Ronald. We are indebted to Bird Studies Canada and collabor- ators in the Motus Wildlife Tracking System for the Ontario stations and to Chris Tonra and The Ohio State University for operation of the Ohio stations; without the Motus system, this work would not have been possible. We thank Andy McLennan of Cartographics Mapping & Design Inc. for prepar- ing the map. Capture and handling of bats was con- ducted under authorization of Wildlife Animal Care Protocol #16-335 and Wildlife Scientific Collector’s Authorization #1083372, both issued by the Ontario Ministry of Natural Resources and Forestry. Comments from D.A.W. Lepitzki, T.S. Jung, and two anonymous reviewers improved previous drafts of this manuscript. Literature Cited Aldridge, H.D.J.N., and R.M. Brigham. 1988. Load carrying and maneuverability in an insectivorous bat: a test of the 5% “rule” of radio-telemetry. Journal of Mam- malogy 69: 379-382. https://doi.org/10.2307/1381393 Arias, L. 2014. Using stable isotope analysis to study alti- tudinal and latitudinal bat migration. Ph.D. thesis, Indiana State University, Terre Haute, Indiana, USA. Arnett, E.B., D.B. Inkley, D.H. Johnson, R.P. Larkin, S. Manes, A.M. Manville, R. Mason, M. Morrison, M.D. Strickland, and R. Thresher. 2007. Impacts of wind energy facilities on wildlife and wildlife habi- tat. Technical review 07-2. Wildlife Society, Bethesda, Maryland, USA. Accessed 7 May 2017. https://wildlife. org/wp-content/uploads/2014/05/Wind07-2.pdf. Baerwald, E.F., W.P. Patterson, and R.M.R. Barclay. 2014. Origins and migratory patterns of bats killed by wind turbines in southern Alberta: evidence from stable isotopes. Ecosphere 5: 1-17. https://doi.org/10.1890/ES 13-00380.1 Banfield, A.W.F. 1974. The Mammals of Canada. National Museums of Canada, Ottawa, Ontario, Canada. Barclay, R.M.R., J.H. Fullard, and D.S. Jacobs. 1999. Variation in the echolocation calls of the hoary bat (La- siurus cinereus). influence of body size, habitat struc- ture, and geographic location. Canadian Journal of Zoology 77: 530-534. https://do1.org/10.1139/z99-008 Bird Studies Canada, Canadian Wind Energy Associ- ation, Environment Canada, and Ontario Ministry of Natural Resources. 2016. Wind energy bird and bat monitoring database: summary of the findings from THE CANADIAN FIELD-NATURALIST Vol. 133 post-construction monitoring reports. Port Rowan, Ontario, Canada. Accessed 26 September 2019. https:// docs.wind-watch.org/Bird-Studies-CAN-Jul2016_ Wind.pdf. Carter, T.C., T.J. Sichmeller, and M.G. Hohmann. 2009. A field- and laboratory-based comparison of adhesives for attaching radiotransmitters to small insectivorous bats. Bat Research News 50: 81-85. Canadian Wind Energy Association. 2017. Wind is Ca- nada’s largest source of new electricity generation for more than a decade. Canadian Wind Energy Asso- ciation, Ottawa, Ontario, Canada. Accessed 3 October 2018. https://canwea.ca/news-release/2017/01/31/wind- energy-continues-strong-growth-canada-2016/. Cryan, P.M. 2003. Seasonal distribution of migratory tree bats (Lasiurus and Lasionycteris) in North America. Journal of Mammalogy 84: 579-593. https://doi.org/10. 1644/1545-1542(2003)084<0579:sdomtb>2.0.co;2 Cryan, P.M. 2011. Wind turbines as landscape impediments to the migratory connectivity of bats. Environmental Law Review 41: 355-370. Accessed 6 July 2017. http:// elawreview.org/articles/volume-41/issue-41-2/wind- turbines-as-landscape-impediments-to-the-migratory- connectivity-of-bats/?hilite="cryan". Cryan, P.M., C.A. Stricker, and M.B. Wunder. 2014. Continental-scale, seasonal movements of a hetero- thermic migratory tree bat. Ecological Applications 24: 602-616. https://doi.org/10.1890/13-0752.1 Davis, W.H. 1969. A recovery of a banded Hoary Bat. Bat Research News 10: 41. Davis, W.H. 1970. Recovery of a banded Lasiurus cinereus. Bat Research News 11: 30. Frick, W.F., E.F. Baerwald, J.F. Pollock, R.M.R. Bar- clay, J.A. Szymanski, T.J. Weller, A.L. Russell, S.C. Loeb, R.A. Medellin, and L.P. McGuire. 2017. Fatalities at wind turbines may threaten population vi- ability of a migratory bat. Biological Conservation 209: 172-177. https://doi.org/10.1016/j.biocon.2017.02.023 Hayes, M.A., P.M. Cryer, and M.B. Wunder. 2015. Seasonally-dynamic presence-only species distribution models for a cryptic migratory bat impacted by wind energy development. PLoS One 10: e0132599. https:// doi.org/10.1371/journal.pone.0132599 Jung, T.S., 1.D. Thompson, R.D. Titman, and A.P. Apple- john. 1999. Habitat selection by forest bats in relation to mixed-wood stand types and structure in central Ontario. Journal of Wildlife Management 63: 1306— 1319. https://doi.org/10.2307/3802849 Kunz, T.H., E.B. Arnett, B.M. Cooper, W.P. Erickson, R.P. Larkin, T. Mabee, M.L. Morrison, M.D. Strick- land, and J.M. Szewezak. 2007. Assessing impacts of wind-energy development on nocturnally active birds and bats: a guidance document. Journal of Wildlife Management 71: 2449-2486. https://doi.org/10.2193/ 2007-270 Kurta, A. 2010. Reproductive timing, distribution, and sex ratios of tree bats in Lower Michigan. Journal of Mammalogy 91: 586-592. https://doi.org/10.1644/09- mamm-a-308.1 Lagerveld, S., R. Janssen, J. Manshanden, A.-J. Haars- ma, S. De Vries, R. Brabant, and M. Scholl. 2017. 2019 MORNINGSTAR AND SANDILANDS: SUMMER MOVEMENTS OF A HOARY BAT 129 Telemetry for migratory bats—a feasibility study. Tech- nical report. Wageningen University, Den Helder, Ne- therlands. https://doi.org/10.18174/417092 Taylor, P.D., T.L. Crewe, S.A. Mackenzie, D. Lepage, Y. Aubry, Z. Crysler, G. Finney, C.M. Francis, C.G. Guglielmo, D.J. Hamilton, R.L. Holberton, P.H. Loring, G.W. Mitchell, D.R. Norris, J. Paquet, R.A. Roncini, J.R. Smetzer, P.A. Smith, L.J. Welsh, and B.K. Woodworth. 2017. The Motus Wildlife Tracking System: a collaborative research network to enhance the understanding of wildlife movement. Avian Con- servation and Ecology 12: 8. https://doi.org/10.5751/ ace-00953-120108 Weller, T.J., K.T. Castle, F. Liechti, C.D. Hein, M.R. Schirmacher, and P.M. Cryer. 2016. First direct evi- dence of long-distance seasonal movements and hiber- nation in a migratory bat. Scientific Reports 6: 34585. https://doi.org/10.1038/srep34585 Received 10 October 2018 Accepted 31 July 2019 The Canadian Field-Naturalist A reconnaissance survey for Collared Pika (Ochotona collaris) in northern Yukon SYDNEY G. CANNINGS!, THOMAS S. JUNG””, JEFFREY H. SKEVINGTON’, ISABELLE DUCLOs*, and SALEEM Dar! 'Canadian Wildlife Service, Environment and Climate Change Canada, Whitehorse, Yukon Y1A 5X7 Canada “Yukon Department of Environment, Whitehorse, Yukon Y1A 2C6 Canada -Agriculture and Agri-food Canada, Ottawa, Ontario K1A 0C6 Canada ‘Canadian Wildlife Service, Environment and Climate Change Canada, Yellowknife, Northwest Territories X1A 2P7 Canada Corresponding author: thomas jung@gov.yk.ca Cannings, S.G., T.S. Jung, J.-H. Skevington, I. Duclos, and S. Dar. 2019. A reconnaissance survey for Collared Pika (Ochotona collaris) in northern Yukon. Canadian Field-Naturalist 133(2): 130-135. https://doi.org/10.22621/cfn.v13312.2109 Abstract Collared Pika (Ochotona collaris) is a cold-adapted Beringian species that occurs on talus slopes and is sensitive to climate warming. Collared Pikas are patchily distributed throughout the sub-Arctic mountains of northwestern Canada and Alaska; however, information on their occurrence in the northern part of their distributional range is limited. In particular, no sur- vey information is known from the southern Richardson Mountains and the Nahoni Mountains. We conducted aerial- and ground-based surveys to document Collared Pika occurrence and general habitat suitability in northern Yukon. We flew 505 km of aerial survey (not including ferrying to targetted survey areas) and performed ground surveys at 22 sites within the Richardson Mountains (including a portion of Daadzaii Van Territorial Park) and the Nahoni Mountains in and adjacent to NViinli Njik (Fishing Branch) Territorial Park. Overall, suitable habitat for Collared Pikas was patchy in the mountains of northern Yukon—talus was sparse and many patches of talus appeared to be unsuitable. Collared Pikas were detected at eight of 22 (36%) sites visited, representing important new records for the species in the northern portion of their range. Our reconnaissance provides a first approximation of habitat suitability for Collared Pikas of the mountains of northern Yukon, as well as new records for the species in the region. These data are useful in better determining the contemporary distribu- tion of Collared Pika through species distribution modelling, and may serve to identify areas for more detailed survey and monitoring initiatives for this climate-sensitive mammal. Key words: Collared Pika; Daadzaii Van Territorial Park; distributional range; NV inlii Njik (Fishing Branch) Territorial Park; Ochotona collaris Introduction Collared Pika (Ochotona collaris) is a small, cold- adapted mammal that is Beringian in origin (Lanier and Olson 2013; Lanier et a/. 2015) and patchily dis- tributed throughout the sub-Arctic mountains of Alaska and northwestern Canada (MacDonald and Jones 1987). Collared Pika are closely associated with talus (1.e., boulder fields) that is interspersed by alpine meadows (MacDonald and Jones 1987; Franken and Hik 2004; Morrison and Hik 2007). Talus provides Collared Pikas with critical protection from predators and inclement weather; as such, they are rarely found far from this habitat. However, not all talus is suitable for Collared Pika. In Tombstone Territorial Park (cen- tral Yukon, Canada), for instance, Collared Pika oc- cupancy was positively associated with large patches of talus that had an average rock size of 30-100 cm, and where Dryas spp. and Carex spp. were available within and adjacent to the patch (L.M. Andresen et al. unpubl. data). Given that talus is naturally patchy on the landscape, Collared Pikas have a fragmented distribution. They have limited dispersal ability and are subject to metapopulation dynamics—whereby local populations may periodically become extinct— leaving apparently suitable habitat variably occupied (Franken and Hik 2004; Morrison and Hik 2007). In Canada, Collard Pika has been assessed as a spe- cies of Special Concern by the Committee on the Sta- tus of Endangered Wildlife in Canada (COSEWIC), largely because of the threat of climate change (CO- SEWIC 2011). The region where the Collared Pika occurs 1s “experiencing climate-driven shifts in habi- tat, temperature, and precipitation at faster rates than elsewhere in Canada” (COSEWIC 2011). Climate-in- duced shrubification of alpine tundra (Danby and Hik 2007; Myers-Smith er al. 2011) is of chief concern re- 130 ©The Ottawa Field-Naturalists’ Club 2019 garding the persistence of Collared Pika populations, as is the depth and duration of snowpack (Morrison and Hik 2007). Local populations of Collared Pikas in southwestern Yukon have declined due to variabil- ity in snowpack (Morrison and Hik 2007). This dem- onstrated sensitivity to climate-induced changes to their habitat, coupled with poor dispersal ability and the fragmented nature of their habitats, make Col- lared Pikas particularly vulnerable to climate change (Morrison and Hik 2007, 2008; COSEWIC 2011). As such, Collared Pika may be a useful bioindicator of climate change impacts to alpine ecosystems (Morri- son and Hik 2008). To assess the range-wide impact of climate change on Collared Pika, wildlife managers require better in- formation on the current species distribution. Precise location data also may be used to develop accurate spatial distribution models that can then be used to predict changes in distribution under different climate change scenarios (e.g., Li et al. 2015; Struebig et al. 2015). Detailed monitoring and systematic surveys of Collared Pika have occurred in southwestern Yukon (Morrison and Hik 2008) and Tombstone Territorial Park (Kukka et al. 2014); however, information on their occurrence in the northern part of their distri- butional range is limited. In particular, no survey in- formation was known from the southern Richardson Mountains and the Nahoni Mountains (1.e., NVinlii Njik [Fishing Branch] Territorial Park). The purpose of this study was two-fold: 1) to sur- vey for the presence of Collared Pika in the northern portion of their range, and 2) to conduct a rapid as- sessment of the habitat suitability for this species in the Richardson Mountains (including Daadzaii Van Territorial Park) and in the Nahoni Mountains (in- cluding in Nriinlii Njik [Fishing Branch] Territorial Park) in northern Yukon. To do so, we undertook a re- connaissance survey for Collared Pika and their habi- tat, using aerial- and ground-based surveys. Our aim was to provide new information on Collared Pikas in the northern portion of their distributional range so that these data can inform habitat modelling, mon- itoring, and management planning, initiatives for this species at risk. Methods We surveyed for Collared Pikas and their habi- tat in northern Yukon, Canada, during 3-6 July 2018. Specifically, we searched for suitable habitat in the Richardson Mountains east and north of Eagle Plains, Yukon (including a portion of Daadzaii Van Territorial Park), as well as in the Nahoni Mountains (including Nriinlii Njik [Fishing Branch] Territorial Park), south of Old Crow, Yukon (Figure 1). We used an AStar helicopter (AS350B3; Eurocopter, Mari- CANNINGS ET AL.: COLLARED PIKA IN NORTHERN YUKON 131 gnane, France) to provide an aerial overview of the habitat conditions in the survey area, and to locate the apparently most suitable habitat for ground-based surveys. We flew 100—400 m above ground level at slow speeds (i.e., 100-120 km/h) in suitable terrain (1.e., mountains) and searched for areas of exten- sive talus slopes and investigated these more closely. Based on occupancy models developed for Collared Pika in Tombstone Territorial Park (L.M. Andresen et al. unpubl. data) that identified predictive habitat covariates, we created four habitat suitability ranks to apply to observed talus slopes (Table 1). We ap- plied these to broadly characterize the suitability of the talus as Collared Pika habitat. At select sites (n = 22) we landed and searched talus areas for Collared Pika presence. We attempted to select the most suitable talus sites for ground sur- veys (i.e., habitat suitability rank 3 or 4; Table 1); how- ever, where no such habitat was apparent we elected to search lower ranked areas of talus to ensure that we covered the possibility that Collared Pikas were se- lecting these sites based on their availability. At each site 4-5 observers searched separate talus patches for approximately 30—60 minutes to detect the pres- ence of Collared Pikas. We walked along the perim- eter of the talus patch, and also traversed portions of the talus to intersect potential Collared Pika territor- ies. Pikas (Ochotona spp.) are highly territorial and vocalize when conspecifics or other species (includ- ing humans) enter their territory (Conner 1984; Trefry and Hik 2009). As such, we largely relied on acous- tically detecting Collared Pika (Moyer-Horner eg al. 2012). We also used binoculars to periodically scan for Collared Pika within the talus; however, Collared Pika are cryptically-coloured to match talus, and may be difficult to visually observe if they are not moving or vocalizing. Finally, pikas build easily recognizable hay piles and latrines within the talus (MacDonald and Jones 1987), and we also used these signs to de- tect their presence (Morrison and Hik 2008; Moyer- Horner et al. 2012; L.M. Andresen et a/. unpubl. data). For each site surveyed we assigned a habitat suita- bility rank of 1—4 (poor to excellent; Table 1). Results and Discussion We flew 505 km of survey effort for Collared Pika in northern Yukon (not including ferrying to target- ted survey areas). This effort included low-level aerial survey of 158 and 178 km of potential Collared Pika habitat (i.e., mountain slopes) in the southern and northern Richardson Mountains, respectively, and 169 km of potential habitat in NVinlii Njik (Fishing Branch) Territorial Park (Figure 1). While some mountains observed had large patches of talus (e.g., approximately >5 ha; Figure 2), in gen- 132 eral we did not observe extensive boulder fields in the same relative abundance as that found in Tomb- stone Territorial Park, likely because much of the Richardson and Nahoni mountains were unglaciated during the Last Glacial Maximum (Catto 1996). Quan- tity of talus, in general, was greatest during our sur- vey in the northern Richardson Mountains, in and ad- jacent to Daadzaii Van Territorial Park, with much of that observed being ranked as 3—4 (good—excel- lent) as Collared Pika habitat. In contrast, quantity of talus was low in the southern Richardson Mountains, and its suitability as Collared Pika habitat was ranked only as 1—2 (poor-marginal). Mountain peaks there 147°W 139°W Old Crow 57.0°N f, ie rishi Branch ’ Territorial Park AE i 55.0°N a We: cA Jtembstone Territorial Park ASE Tie? THE CANADIAN FIELD-NATURALIST Vol. 133 were low and rounded and most talus observed was small (<30 cm) and unsuitable as Collared Pika habi- tat. Potential Collared Pika habitat was variable in the Nahoni Mountains, with some local areas hav- ing abundant talus of suitable characteristics (rank 2—4; marginal-excellent) and other areas have little talus available. Overall, we would rate the northern Richardson Mountains being most suitable as Col- lared Pika habitat in the areas we surveyed, although local areas in the Nahoni Mountains were also suit- able. We observed little suitable Collared Pika habitat in the southern Richardson Mountains. We detected Collared Pika presence at nine of 22 137°W >) | ac -) \ Eagle Plains = Figure 1. Flight path (black transect) of an aerial survey for Collared Pika (Ochotona collaris) and their habitat in northern Yukon. Closed circles are sites surveyed on the ground where Collared Pika was detected, and open circles where they were not detected. The stippled polygon represents the putative distributional range of Collared Pika (Lanier and Hik 2016). Stars indicate human settlements. Site numbers are in Table 2. The insert shows the study area situated within Yukon, Canada. 2019 CANNINGS ET AL.: COLLARED PIKA IN NORTHERN YUKON 183 TABLE 1. Description of habitat suitability ranks given to areas surveyed for Collared Pika (Ochotona collaris) in northern Yukon, July 2018. Bapieayuilaeility General description rank 1 Poor habitat quality: Average rock size <30 cm or >100 cm with many smaller rocks that fill in the interstitial spaces between the larger rocks. Marginal habitat quality: Average rock size 30—100 cm, but with areas of extensive shrub cover in 2 and around the talus, or with no to small amounts of Dryas spp. cover adjacent to talus, or many smaller rocks that fill in the interstitial spaces between the larger rocks. 3 Good habitat quality: Average rock size 30—50 cm; large area covered by talus slopes that are interspersed with non-shrubby patches of vegetation including Dryas spp. a Excellent habitat quality: Average rock size 50-100 cm; large area covered by talus slopes that are interspersed with non-shrubby patches of vegetation including extensive Dryas spp. cover. FiGure 2. Photograph of site 8 (see Table 2)—an example of the site characteristics where Collared Pika (Ochotona collaris) were observed in northern Yukon, Canada. Photo: J.H. Skevington. sites surveyed on the ground (Table 2; Figure 1). One of these sites was just outside the northern boundary of Tombstone Territorial Park, where Collared Pika are already known and monitored (Kukka et al. 2014; Figure 1). We detected Collared Pika at three of five and one of seven sites surveyed in the northern and southern Richardson Mountains, respectively, and four of eight sites in the Nahoni Mountains (Figure 1). At two sites (sites 4 [Figure 3a] and 11; Table 2) we detected only old sign of Collared Pika, indicating that the population may have been extirpated. This included the single site where we detected Collared Pika in the southern Richardson Mountains (site 4, Table 2). Although our survey was not designed to es- timate the density of Collared Pika at our sites, it ap- peared that they persisted at low densities at all the sites where they were detected. In only one instance did we detect more than a single individual at a site. In suitable habitat in southwestern Yukon, Collared Pika density is estimated at <1 to 4 individuals per ha (Morrison and Hik 2007). Habitat suitability was variable among the 22 sites we surveyed on the ground, with 36% of them having poor-marginal (1-2 ranks) and 64% having good-ex- cellent (3—4 ranks) habitat suitability ranks (Table 2). Collared Pikas were detected at only one of eight sites that were of poor-marginal habitat suitability, but they were detected at seven of 14 sites we classified as being of good-excellent habitat suitability (Figures 2 and 3). Anecdotally, the most limiting habitat fea- tures at the sites we surveyed were likely the aver- age rock size being <50 cm, coupled with many sites having extensive shrubby vegetation in and around the talus (as opposed to Dryas spp. and other forage plants; L.M. Andresen ef a/. unpubl. data). The ele- vation where we detected Collared Pika was variable with a mean of 961.1 + 175.8 m (SD; range = 685-— 1329 m; Table 2). Other mammals or their sign (e.g., burrows, dig- gings, dens, scat, antlers) detected at our survey sites for Collared Pika included Grizzly Bear (Ursus arctos), Wolverine (Gulo gulo), Dall’s Sheep (Ovis dalli), Caribou (Rangifer tarandus), Muskox (Ovi- bos moschatus), Moose (Alces americanus), Arctic Ground Squirrel (Urocitellus parryii), and small ro- dents (likely Singing Vole [Microtus miurus], Tundra Vole [Microtus oeconomus|, Northern Red-backed Vole [Myodes rutilus], or Brown Lemming [Lemmus trimucronatus]; Table 2). Notably, we did not observe any sign of Hoary Marmot (Marmota caligata) dur- ing our survey. Moreover, Arctic Ground Squirrels were surprisingly not abundant at any of the sites we surveyed north of the Ogilvie Mountains, where there they are often on mountains associated with Collared Pika (T.S.J. pers. obs.). We detected Arctic Ground Squirrels at only six of 21 (29%) sites that were sur- veyed north of the Ogilvie Mountains. Our reconnaissance of the mountains of north- ern Yukon provides a first approximation of habi- tat suitability for Collared Pikas, as well as impor- tant new records for the species, in the northern portion of their distributional range. These data are useful in determining the contemporary distribu- tion of Collared Pika through species distribution modelling and may serve to identify areas for more detailed survey and monitoring initiatives for this 134 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Description of sites surveyed for Collared Pika (Ochotona collaris) in northern Yukon, 3—6 July 2018. Si Habitat Collared Pika Detection : Elevation Other mammals ite Location rank* typet (m) observed 1 3 Yes RED 64.77657°N, 139.08717°W 1329 [e2386e7 2 1 No 66.24000°N, 136.09439°W 1092 IF Dee 3 3 No — 66.22826°N, 135.84087°W 771 \Remsers 4 2 Yes ) 66.36445°N, 136.21266°W 944 StS, 7 5 1 No 66.44556°N, 136.11227°W 851 7 6 1 No — 66.62994°N, 136.20248°W 1205 aoe 7 2 No — 66.89725°N, 136.13144°W 565 ise, 8 3 Yes 2 66.96921°N, 136.14555°W 927 |e ae: 9 4 No — 67.13135°N, 136.26424°W 1011 1 Soe 10 3 Yes 2 67.13584°N, 136.27515°W 862 5,6 ‘ia 4 Yes 3 67.65533°N, 137.01370°W 1047 | S28 Ome]. 12 5 No — 67.67641°N, 137.08986°W 941 2 pay 13 4 No — 67.60668°N, 137.31078°W 957 he 28 Bea 14 2 No — 66.61674°N, 136.79306°W 873 3 ifs 3 No — 66.13930°N, 139.29184°W 994 15657 16 3 Yes pe 66.14023°N, 139.39793°W 817 1,5, 6, 7, 8 17 3 Yes es 66.21690°N, 139.53180°W 685 Sa0n 7 18 4 Yes 2.8 66.31663°N, 139.78535°W 1120 N22 Omer, 19 1 No — 66.37148°N, 139.77873°W 871 1 Wes: 20 2 No — 66.89491°N, 139.85330°W 984 [eo OE Om Dp 21 1 No — 66.65713°N, 140.79265°W 680 out 22 3 Yes 2 66.34312°N, 140.20544°W 919 ee *See Table | for habitat suitability rank descriptions. tDetection types as follows: 1 = visual, 2 = acoustic, 3 = sign (haypiles or latrines). £Codes for other mammals as follows: 1 = Grizzly Bear (Ursus arctos), 2 = Dall’s Sheep (Ovis dalli), 3 = Caribou (Rangifer tarandus), 4 = Muskox (Ovibos moschatus), 5 = Moose (Alces americanus), 6 = Arctic Ground Squirrel (Spermophilius parryii), 7 = voles or lemmings, 8 = Wolverine (Gulo gulo). 5 bi. ae FIGURE 3. RR of the ay habitat conditions at select survey sites for Collared Pika (Ochotona collaris) in the southern Richardson Mountains (a is site 4) and Nahoni Mountains (b is site 15) in northern Yukon, Canada. Photos: J.H. Skevington. climate-sensitive mammal. We suggest that Collared Pika presence and habitat suitability was good in the northern Richardson Mountains, moderate-to-good in the Nahoni Mountains, and poor in the southern Richardson Mountains. We emphasize, however, that our work is preliminary in nature and our abil- ity to thoroughly survey our target areas was lim- ited; thus, this region would benefit from further survey effort. Preliminary habitat suitability map- ping in northern Yukon, using imagery from remote sensing to map large patches of talus, would likely be helpful in determining other sites with a high probability of Collared Pika occurrence. We suggest that the northern Richardson Mountains (in and ad- jacent to Daadzaii Van Territorial Park) would be an important area to focus future survey efforts, per- haps in conjunction with similar surveys for other small mammals of conservation interest in the re- gion (e.g., collared lemmings [Discrostonyx spp.]; Jung et al. 2014). 2019 Acknowledgements We thank Vince Edmonds (TransNorth Helicop- ters) for safe piloting and effectively contributing to the survey effort. Darius Elias (Vuntut Gwich’in First Nation) kindly assisted in the field during a portion of our surveys and provided logistical support. Piia Kukka (Government of Yukon) made the maps. Shannon Stotyn (Environment and Climate Change Canada), Bruce McLean, Kirby Meister, and Carrie Mierau (Government of Yukon) are also thanked for providing logistical support. We thank Environment and Climate Change Canada, Polar Continental Shelf Project, and Environment Yukon, for supporting this work. Bruce Bennett, Dwayne Lepitzki, Graham Forbes, and two anonymous reviewers kindly pro- vided comments on an earlier draft of this manuscript that improved its quality. Literature Cited Catto, N.R. 1996. Richardson Mountains, Yukon-North- west Territories: the northern portal of the postulated ‘ice-free corridor’. Quaternary International 32: 3-19. https://doi.org/10.1016/1040-6182(95)00062-3 Conner, D.A. 1984. The role of an acoustic display in ter- ritorial maintenance in the pika. Canadian Journal of Zoology 62: 1906-1909. https://doi.org/10.1139/z84-280 COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2011. COSEWIC assessment and status report on the Collared Pika Ochotona collaris in Canada. Committee on the Status of Endangered Wild- life in Canada, Ottawa, Ontario, Canada. Danby, R.K., and D.S. Hik. 2007. Variability, contingency and rapid change in subarctic alpine tree line dynamics. Journal of Ecology 95: 352-363. https://doi.org/10.1111/ j.1365-2745.2006.01200.x Franken, R.J., and D.S. Hik. 2004. Influence of habitat quality, patch size and extinction dynamics of collared pika Ochotona collaris. Journal of Animal Ecology 73: 889-896. https://doi.org/10.1111/j.0021-8790.2004.00865.x Jung, T.S., B.G. Slough, D.W. Nagorsen, and P.M. Kukka. 2014. New records of the Ogilvie Mountains Collared Lemming (Dicrostonyx nuntakensis) in central Yukon. Canadian Field-Naturalist 128: 265-268. https://doi.org/ 10.22621/cfn.v128i3.1605 Kukka, P.M., A. McCulley, M. Suitor, C.D. Eckert, and T.S. Jung. 2014. Collared pika (Ochotona collaris) occu- pancy in Tombstone Territorial Park, Yukon: 2013 sur- vey results. Yukon Department of Environment Survey Report SR-14-01, Whitehorse, Yukon, Canada. Lanier, H.C., and D.S. Hik. 2016. Ochotona collaris. The IUCN Red List of Threatened Species 2016: e.T41257A 45182533. https://doi.org/10.2305/IUCN.uk.2016-3.rlts. t41257a45182533.en Lanier, H.C., R. Massati, Q. He, L.E. Olson, and L.L. Knowles. 2015. Colonization from divergent ances- tors: glaciation signatures on contemporary patterns CANNINGS ET AL.: COLLARED PIKA IN NORTHERN YUKON 135 of genomic variation in collared pikas (Ochotona col- laris). Molecular Ecology 24: 3688-3705. https://do1.org/ 10.1111/mec.13270 Lanier, H.C., and L.E. Olson. 2013. Deep barriers, shal- low divergences: reduced phylogeographic structure in the collared pika (Mammalia: Lagomorpha: Ochotona collaris). Journal of Biogeography 40: 466—476. https:// doi.org/10.1111/jbi.12035 Li, R., M. Xu, M. Hang, G. Wong, S. Qiu, X. Li, D. Ehrenfeld, and D. Li. 2015. Climate change threatens giant panda protection in the 21st Century. Biological Conservation 182: 93-101. https://doi.org/10.1016/).bio con.2014.11.037 MacDonald, S.O., and C. Jones. 1987. Ochotona collaris. Mammalian Species 281: 1—4. https://doi.org/10.2307/ 3503971 Morrison, S.F., and D.S. Hik. 2007. Demographic analysis of a declining pika Ochotona collaris population: link- ing survival to broad-scale climate patterns via spring snowmelt patterns. Journal of Animal Ecology 76: 899— 907. https://doi.org/10.1111/).1365-2656.2007.01276.x Morrison, S.F., and D.S. Hik. 2008. When? Where? And for how long? Census design considerations for an alp- ine lagomorph, the collared pika (Ochotona collaris). In Lagomorph Biology. Edited by P.C. Alves, N. Ferrand, and K. Hacklander. Springer, Berlin, Heidelberg, Ger- many. Moyer-Horner, L., M.M. Smith, and J. Belt. 2012. Citizen science and observer variability during American pika surveys. Journal of Wildlife Management 76: 1472— 1479. https://doi.org/10.1002/jwmg.373 Myers-Smith, I.H., B.C Forbes, M. Wilmking, M. Hallinger, T. Lantz, D. Blok, K.D Tape, M. Macias- Fauria, U. Sass-Klaassen, E. Lévesque, S. Boudreau, P. Ropars, L. Hermanutz, A. Trant, L. Siegwart Collier, S. Weijers, J. Rozema, S.A. Rayback, N.M. Schmidt, G. Schaepman-Strub, S. Wipf, C. Rixen, C.B. Menard, S. Venn, S. Goetz, L. Andreu-Hayles, S. Elmendorf, V. Ravolainen, J. Welker, P. Grogan, H.E. Epstein, and D.S. Hik. 2011. Shrub expansion in tundra ecosystems: dynamics, impacts and research priorities. Environmental Research Letters 6: 045509. https://doi.org/10.1088/1748-9326/6/4/045509 Struebig, M.J., M. Fischer, D.L.A. Gaveau, E. Meijaard, S.A. Wich, C. Gonner, R. Sykes, A. Wilting, and S. Kramer-Schadt. 2015. Anticipated climate and land- cover changes reveal refuge areas for Borneo’s orang- utans. Global Change Biology 21: 2891—2904. https://doi .org/10.1111/gcb.12814 Trefry, S.A., and D.S. Hik. 2009. Eavesdropping on the neighbourhood: collared pika (Ochotona collaris) re- sponses to playback calls of conspecifics and heterospe- cifics. Ethology 115: 928-938. https://doi.org/10.1111/. 1439-0310.2009.01675.x Received 23 July 2018 Accepted 8 August 2019 The Canadian Field-Naturalist Note Occurrence of the rare marine littoral millipede, Thalassisobates littoralis (Diplopoda: Nematosomatidae), in Canada DONALD F. MCALPINE New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada; email: donald.mcalpine@ nbm-mnb.ca McAlpine, D.F. 2019. Occurrence of the rare marine littoral millipede, Thalassisobates littoralis (Diplopoda: Nemato- somatidae), in Canada. Canadian Field-Naturalist 133(2): 136-138. https://doi.org/10.22621/cfn.v13312.2215. Abstract The first Canadian occurrence of the rare, marine littoral millipede, Thalassisobates littoralis, is reported from Campobello Island in the outer Bay of Fundy, New Brunswick. One of only a few North American occurrences, this is the most north- erly to date from the continent. Key words: New Brunswick; marine habitat; marine millipede; dispersal; anthropochorus species; Thalassisobates littoralis World-wide, Thalassisobates littoralis (Silvestri, 1903) (no common name) is one of the few marine lit- toral millipedes and is considered rare (Blower 1985; Barber 2009). The species occurs under stones and seaweed, in rock crevices or shingle in or above the tidal zone, and sometimes in coastal caves (Enghoff 1987, Cawley 1997). Thalassisobates littoralis has a wide but scattered distribution, with reports (of- ten single) from the coasts of Europe, the Balearic Islands, Algeria, and the eastern United States (Enghoff 2013). The centre of its distribution ap- pears to be the western Mediterranean basin (Kime 1999), with Kime (1999) suggesting that 7° /ittoralis may have been introduced to northwestern Europe. Reporting the first North American occurrences, Enghoff (1987) speculated that the species was of European origin, but was uncertain whether its am- phi-Atlantic distribution was natural or the result of human introduction. He noted that all amphi-Atlantic millipedes previously reported from North America, with one possible exception, can be regarded as intro- ductions to the continent and that the direction of the Gulf Stream is not conducive to the natural dispersal of 7: /ittoralis from Europe to North America. Thus, not unreasonably, 7: /ittoralis has been considered of anthropochorus origin in North America (Kime 1999; Golovatch and Kime 2009). Previously, 7: /ittoralis has been recorded in North America only from the southwest shore of Chincoteague Island, Virginia, in 1964, from an un- known locale in Massachusetts (date unknown; Eng- hoff 1987), and, more recently (2005-2009), from six of the 34 islands that make up the Boston Harbor Recreation Area (Boston Harbor Islands 2014). Here, I document the first occurrence of 7° /ittoralis from Canada and the most northerly to date on the North American continent. On 24 September 2017, I found 7° /ittoralis to be present, but patchily distributed, along a cob- ble shoreline at Herring Cove, in Herring Cove Provincial Park, Campobello Island, New Brunswick (44.85956°N, 66.93188°W), along the western shore of the Bay of Fundy. Millipedes were present un- der patches of decomposing Bladderwrack (Fuscus vesiculosus) above and below the high-water mark (Figure la). Where present, millipedes were abun- dant (Figure 1b). In a sample of 104 specimens, 55 females and 49 males were present, close to a 1:1 sex ratio. The whole body of a single male (Figure 1c) and a series of views of the male peltogonopods, di- agnostic for 7: littoralis, are shown in Figure 1d-f. Voucher specimens have been deposited in the New Brunswick Museum (NBM 10776). After the discovery of 7! /ittoralis on Campobello Island, two other cobble beach sites in New Bruns- wick were searched along the western coast of the Bay of Fundy (Alma, 45.596739°N, 64.948728°W, and Browns Beach, West Quaco, 45.319236°N, 65.551114°W), for 7: littoralis without success. Fur- ther field investigations will be required to determine 136 ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). MCALPINE: 7HALASSISOBATES LITTORALIS IN CANADA FicurE 1. a. Shoreline at Herring Cove Provincial Park, New Brunswick, showing habitat for Thalassisobates littoralis. b. Concentration of millipedes under Bladderwrack (Fuscus vesiculosus). c. Habitus of male 7: /ittoralis; scanning elec- tron microscope images show d. superior, e. inferior, and f. distal views of the diagnostic peltogonopods. Photos: a—b. D.F. McAlpine. Photos: c—f. Nhu Trieu. 138 the full distribution and true abundance of 7: /ittora- lis in Atlantic Canada. Acknowledgements I thank Amber McAIpine-Mills for help in the field, investigating sites at Alma and West Quaco, New Brunswick. I also thank Nhu Trieu, of the University of New Brunswick Microscopy and Microanalysis Facility, for providing Figure Ic-f. Literature Cited Barber, A.D. 2009. Littoral myriapods: a review. Soil Organisms 81: 735-760. Blower, J.G. 1985. Millipedes. E.J. Brill, London, United Kingdom. Boston Harbor Islands. 2014. All taxa biodiversity inven- tory: list of specimens: Thalassisobates littoralis. Pre- sident and Fellows of Harvard College, Boston, Mas- sachusetts, USA. Accessed 14 October 2019. http://140. 247.96.247/boston_islands/mantisweb/specimen_list_ bhi.php?id=59261. THE CANADIAN FIELD-NATURALIST Vol. 133 Cawley, M. 1997. Distribution records for uncommon milli- pedes (Diplopoda) including Thalassisobates littoralis (Silvestri) new to Ireland. Irish Naturalist Journal 25: 380-382. Enghoff, H. 1987. Thalassisobates littoralis (Silvestri— an amphiatlantic millipede (Diplopoda, Julida, Nema- tosomatidae). Entomologist’s Monthly Magazine 123: 205-206. Enghoff, H. 2013. New montane, subterranean congen- ers of a littoral millipede, genus Thalassisobates (Di- plopoda: Julida: Nemasomatidae). Journal of Natural History 47: 1613-1625. https://doi.org/10.1080/00222933. 2012.759289 Golovatch, S.I., and R.D. Kime. 2009. Millipede (Diplo- poda) distributions: a review. Soil Organisms 81: 565— 597. Kime, R.D. 1999. The continental distribution of British and Irish millipedes. Bulletin of the British Myriapod Group 15: 33-76. Received 15 January 2019 Accepted 31 July 2019 The Canadian Field-Naturalist A practical technique for preserving specimens of duckmeal, Wolffia (Araceae) DANIEL F. BRUNTON 216 Lincoln Heights Roads, Ottawa, Ontario KIA 8A8; email: bruntonconsulting@rogers.com Brunton, D.F. 2019. A practical technique for preserving specimens of duckmeal, Wolffia (Araceae). Canadian Field-Naturalist 133(2): 139-143. https://doi.org/10.22621/cfn.v133i2.2108 Abstract Making identifiable herbarium vouchers of the minute aquatic vascular plant duckmeal, Wolffia (Lemnoideae; Araceae) has typically required plants to be preserved in transparent, space-consuming vials that are fragile, difficult to work with, and labourious to prepare. An alternative technique for dry-mounting Wolffia within a layer of transparent, acid-free glue pre- sents a promising alternative. Although the largely water-filled individual plants still compress substantially, this prepara- tion technique results in specimens that retain their colour, size, and, most important, their shape. This greatly enhances the possibility of confident identification and simplifies both specimen preparation and storage. Key words: Wolffia, Lemnoideae; Araceae; herbarium specimen preparation; storage Introduction Dore (1957) described the difficulty collectors face in securing voucher specimens of duckmeal (Wolffia spp., Lemnoideae, Araceae; also known as water- meal) in a condition that permits their re-examination and identification. Three species of this minute, sim- ple plant, which is uncommon to rare throughout most of its Canadian range, typically form dense, floating mats (Figure 1) consisting of thousands and even millions of individuals (Brunton 2018; Brunton and Bickerton 2018). At least two Wolffia species (Figure 2) are native in southern Ontario and Quebec (Crow and Hellquist 2000). Individuals of this, the smallest flowering plant in the world, shrivel up into greenish- brown dust when air dried as conventional vascular plant herbarium specimens (Figure 3). This deforma- tion eliminates the possibility of further identifica- tion, which is heavily dependent on plant shape char- acteristics (Crow and Hellquist 2000). Accordingly, the potential use of such material for taxonomic or phytogeographic analysis is substantially reduced. Dore’s (1957) solution to that curatorial challenge was to place Wolffia samples in a preservative fluid (alcohol) in a sealed glass vial, which was then at- tached to a standard herbarium sheet. Although this technique permits the voucher plants to retain their shape and size, it is an involved process that results in a slurry of colourless plants floating in a cumber- some, fragile vessel that takes up considerable space in a conventional herbarium arrangement. Individual plants also move freely within the vial and are diffi- cult to follow or relocate. Unless the vial is opened (necessitating repetition of the entire voucher prep- aration process), the view of individual plants is also obscured and distorted by the glass container. There is arisk of breakage or leakage of the vials, as is evi- dent in Dore’s collection preserved in the Agriculture and Agri-Food Canada herbarium (DAO). In addi- tion, the alcohol-filled vials are sealed with a flame or heat, a process that requires some experience and poses potential hazards (P.M. Catling pers. comm. 14 March 2019). Although preferable to simply dry- ing plants into an unrecognizable condition, the vial solution is a cumbersome and unsatisfying curator- ial response. The easy availability of high-quality digital im- agery in the field now provides a practical enhance- ment of the traditional air-dried voucher technique, because hard-copy images of fresh (pre-dried) vou- cher material can be affixed to the voucher sheet. However, photographs are only two-dimensional rep- resentations and cannot be used for detailed morpho- logical studies. Discussions with numerous field botanists in east- ern Canada have confirmed that the difficulty in securing reasonable quality Wolffia vouchers signifi- cantly discourages their collection. This represents a floristic and conservation problem in Ontario and Quebec where Northern Duckmeal (Wolffia borealis (Engelmann) Landolt & Wildi ex Gandhi, Wiersema & Brouillet) @ Wolffia punctata Grisebach) and Co- A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 139 ©The Ottawa Field-Naturalists’ Club 140 THE CANADIAN FIELD-NATURALIST Vol. 133 = cd = em < oe = a = we oe pies” ee “< = z ae 6 f a eee ste sy s a a a = = Se] ot i pace See = = “ ae Shana FicurE 1. Dense duckmeal (Wolffia spp.) mat completely covering creek surface. 8 August 2011, Beachburg, Renfrew County, Ontario. Photo: D.F. Brunton. Wolffia columbiana Wolffia borealis FIGURE 2. Fresh (live) Ontario native duckmeal (Wolffia) species. 31 July 2011, Peterborough County, Ontario, D.F. Brunton & K.L. McIntosh 17,896B. Photo: D.F. Brunton. lumbia Duckmeal (Wolffia columbiana H. Kirsten) (= Wolffia arrhiza auct., non (L.) Horkel) are almost always regionally uncommon (Dore 1957; Soper 1962; Oldham 2017) and the possibly introduced Papil- late Watermeal (Wolffia brasiliensis Weddell) is rare (Thomson 2005). Both W. borealis and W. colum- biana are considered rare and of conservation con- cern in Quebec (Tardif et a/. 2005). In addition, popu- lations are increasingly being found beyond Wolffia’s traditional northern limit along the southern edge of the Canadian Shield in Ontario (Soper 1962), sug- gesting ongoing range expansion (Brunton 2018). Does this change represent climate change? In- —_ iy Be Be a Figure 3. Shrivelled air-dried Northern Duckmeal (Wolffia borealis). 31 July 2011, Peterborough County, Ontario, D.F. Brunton & K.L. McIntosh 17,896B. Photo: D.F. Brunton. creased populations of certain waterfowl species? Transportation by human activity? All of the above? That is not clear, but the range of Wolffia unquestion- ably is enlarging in Canada and new Wolffia oc- currences must be recorded with voucher speci- mens to document this and assess its implications. Accordingly, a practical procedure for producing in- formative vouchers is an important part of addressing these questions. 2019 The dry-mounted specimen preparation technique described in this paper is an adaptation of the method described by Brunton (1990) for securing mass sam- ples of even smaller and more fragile megaspores of the aquatic lycophyte /soetes (quillwort) in a readily examinable state. This new dry-mounting procedure has been used to prepare samples in approximately two dozen Wolffia collections to date. It has resulted in vouchers of superior quality and utility in com- parison to most vouchers prepared using traditional methods. Methods A small (5—10 mL) sample of fresh Wolffia plants is placed in a plastic or glass vial with enough of the water-diluted (ca. 75% of full strength), acid-free glue commonly used in herbarium specimen preparation to maintain the sample free floating. The glue used here was commercially available Weldbond Universal Adhesive (Frank T. Ross & Sons Ltd., Markham Ontario, Canada). No attempt was made to assess the performance of other commercial brands of adhesive, but it is suspected that any water-soluble white herb- arium glue would perform comparably. The sample is left to soak for ~48 h. The container is covered to pre- vent the glue from drying and solidifying. Because Wolffia plants are exceedingly buoyant, the slurry of plants is stirred periodically (two or three times in the course of the soaking period) to ensure thorough con- tact with the glue solution. After about 25-30 h, the colour of the glue solu- tion takes on a distinctively green cast. The Wolffia plants also appear to be less buoyant, floating lower in the solution and remaining submerged slightly longer when stirred. This is interpreted as indicating that the plants have become slightly waterlogged and that the glue has penetrated their tissues. After ~48 h the slurry is spread (poured) onto a sheet of plain paper (a heavier gauge, pH neutral paper is recommended) or directly onto a herbarium sheet (White 100 Percent Rag Bond — neutral pH 20 or 24 Ib; Herbarium Supply Co., Bozeman, Montana, USA) in as thin a layer as possible and allowed to air dry. Concentrations of plants on the mounting sheet can be thinned by gently passing a dissection needle or probe through them. This must be done immedi- ately after the slurry has been poured, however, as the solution becomes too tacky for smooth manipu- lation within as little as 10 min. The sample (Figure 4) is dry and ready for examination or processing as a herbarium specimen within about 2 h. Results Applying this voucher preparation technique to over 30 samples of native Ontario Wolffia involving BRUNTON: PRESERVING SPECIMENS OF DUCKMEAL 141 FiGurE 4. Sample of dried, glue-impregnated Columbia Duckmeal (Wolffia columbiana), larger intermixed plants are Spirodella polyrhiza (L.) Schleiden. 25 August 2017, Renfrew, Renfrew County, Ontario, D.F. Brunton & K. Fleming 19,783. Photo: D.F. Brunton. 23 separate collections (Appendix 1) has produced encouraging results. Although the dry, glue-impreg- nated plants are completely deflated and flattened, they have, for the most part, retained their original size, shape, and colour (Figure 5). Some aspects of their morphology remain evident: for example, the punctate (dotted) surface of W. borealis is clearly vis- ible in the dried plants. Rehydrating shrivelled, long-dried herbarium ma- terial may also be worthwhile. In the two samples tested, the glue solution did not become as green as it did with fresh material, suggesting that the rehy- drating plants do not become as completely satur- ated. Nonetheless, the rehydrated material was more distinguishable than it would have been if specimens had only been air dried. It is recommended that collectors making Wolffia vouchers include a packet of air-dried plants (“green dust”) prepared in the traditional way as well. This typically indistinguishable air-dried material is pot- entially useful for chemical and molecular analyses. Ideally, photos of fresh plants should be included in the voucher. When used in combination with glue- impregnated plants, these additional specimen com- ponents provide the most complete vouchers possible for multidisciplinary investigations. More extensive experimentation would refine the technique described here. It may well provide simi- larly satisfying results, for example, for other aquatic plants that occur in vast numbers of minute individ- uals, such as duckweed (Lemna), duck-meat (Spiro- dela), and mud-midget (Wolffiella). A variation on this technique to accommodate larger specimens might also prove useful for the preservation of exceptionally delicate and/or fragile water nymph (Najas), ditch- grass (Ruppia), horned pondweed (Zannichellia), and 142 THE CANADIAN FIELD-NATURALIST Vol. 133 Figure 5. Close-up of glue-impregnated Columbia Duckmeal (Wolffia columbiana) retaining shape and size. 25 August 2017, Renfrew, Renfrew County, Ontario, D.F. Brunton & K. Fleming 19,783. Photo: D.F. Brunton. pondweed (Potamogeton and Stuckenia) plants in a more natural form. As currently understood, however, this technique provides a logistically simple, space- efficient, permanent preservation alternative for mak- ing Wolffia vouchers, for which there is an immediate need. The technique has the added benefit of being hazard free for the preparator and posing no curator- ial risks to associated herbarium material. Acknowledgements This paper benefited significantly from input by Associate Editor, Paul M. Catling, and reviewers, Bruce Ford, University of Manitoba, Winnipeg, Mani- toba, Garret Crow, Calvin College, Grand Rapids, Mi- chigan, and Michael J. Oldham, Ontario Ministry of Natural Resources and Forestry, Peterborough, On- tario. Their valuable comments and observations were based on extensive first-hand field experience with Wolffia. The author was inspired to investigate and collect these curious and challenging aquatic plants in the first place by the enthusiastic encouragement and assistance of the late W.G. (Bill) Dore (1912-1996), then of Agriculture Canada, Ottawa. Literature Cited Brunton, D.F. 1990. A device for the protection of spore samples from /soetes (Isoetaceae) voucher specimens. Taxon 39: 226-228. https://doi.org/10.2307/1223021 Brunton, D.F. 2018. The changing distribution and extra- ordinary abundance of Wo/ffia in Ontario. Field Botanists of Ontario Newsletter 30(2). Accessed 15 March 2019. https://www.researchgate.net/publication/326719302 _ Wolffia_in Ontario FBO 2018. Brunton, D.F., and H. Bickerton. 2018. New records for the 2019 Eastern Mosquito-fern (Azolla cristata: Salviniaceae) in Canada. Canadian Field-Naturalist 132: 350-359. https:// doi.org/10.22621/cfn.v13214.2033 Crow, G.E., and C.B. Hellquist. 2000. Aquatic and wet- land plants of northeastern North America, Volume Two Angiosperms: Monocotyledons. University of Wiscon- sin Press, Madison, Wisconsin, USA. Dore, W.G. 1957. Wolffia in Canada. Canadian Field-Natu- ralist 71: 10-16. Accessed 15 March 2019. https://www. biodiversitylibrary.org/item/89178#page/22. Oldham, M.J. 2017. List of the vascular plants of Ontario’s Carolinian Zone (Ecoregion 7E). Ontario Ministry of Natural Resources and Forestry, Peterborough. https:// doi. org/10.13140/RG.2.2.34637.33764 Soper, J.H. 1962. Some genera of restricted range in the BRUNTON: PRESERVING SPECIMENS OF DUCKMEAL 143 Carolinian flora of Canada. Transactions of the Royal Canadian Institute 34 (Part 1): 3-56. Tardif, B., G. Lavoie, et Y. Lachance. 2005. Atlas de la bio- diversité du Québec. Ministere du Développement du- rable, de l’ Environnement et des Parcs, Direction du dé- veloppement durable, du patrimone écologique et des Parcs, Québec, Canada. Accessed 15 March 2019. https:// tinyurl.com/y3794fcj. Thomson, E.R. 2005. Papillate Watermeal, Wolffia brasili- ensis, in eastern Ontario: an addition to the Flora of Canada. Canadian Field-Naturalist 119: 137-138. https:// doi.org/10.22621/cfn.v11911.97 Received 20 July 2018 Accepted 25 July 2019 Appendix 1: Wolffia specimens prepared, partly or entirely, using the Weldbond technique. Herbarium acro- nyms of Thiers (continuously updated). Wolffia borealis Ontario: City of Ottawa, 45.36843°N, 75.79522°W, west end of Turtle Bay, southwest portion of Mud Lake, Britannia Conservation Area, D.F. Brunton 17,106, 7 September 2007 (DFB); Ontario: City of Ottawa, 45.33572°N, 75.92777°W, in Shirley’s Brook along south side of Trillium Woods, South March Highlands, Kanata, D.F. Brunton 17,125, 5 October 2007 (DFB),; Ontario: City of Ottawa, 45.39846°N, 75.96812°W, south end of Constance Lake by inlet of Constance Creek, West Carleton, D.F. Brunton 18,036, 13 Au- gust 2011 (CAN, DFB); Ontario: City of Ottawa, 45.34581°N, 75.86959°W, abandoned sewage treatment plant 395 m south southeast of Range Road/Carling Avenue inter- section, Nepean, D.F. Brunton 18,913A, 8 October 2014 (WIN, DFB); Ontario: Frontenac County, 44.27868°N, 76.59344°W, west side of Collins Creek north side of Creeksford Road, Kingston, D.F. Brunton 18,050A, 24 August 2011 (WIN, DFB); Ontario: Peterborough County, 44.76643°N, 78.49064°W, west side of White’s Lake Road, north side of Cry- stal Lake, Galway & Cavendish Township, D.F. Brunton and K.L. McIntosh 17,986B, 12 July 2011 (DFB); Ontario: Peterborough County, 44.38742°N, 77.97751°W, east side of mill pond by outlet, north side of High- way 7, east end of Norwood, Asphodel Township, D.F. Brunton 19,916, 6 September 2017 (DFB). Wolffia columbiana Ontario: City of Ottawa, 45.36843°N, 75.79522°W, west end of Turtle Bay, southwest portion of Mud Lake, Britannia Conservation Area, D.F. Brunton 17,107, 7 September 2007 (DFB); Ontario: City of Ottawa, 45.33572°N, 75.92777°W, in Shirley’s Brook along south side of Trilltum Woods, South March Highlands, Kanata, D.F. Brunton 17,126, 5 October 2007 (DFB); Ontario: City of Ottawa, 45.39846°N, 75.96812°W, south end of Constance Lake by inlet of Constance Creek, West Carleton, D.F. Brunton 18,035, 13 August 2011 (CAN, DFB),; Ontario: City of Ottawa, 45.34581°N, 75.86959°W, abandoned sewage treatment plant 395 m south- southeast of Range Road/Carling Avenue inter- section, Nepean, D.F. Brunton 18,913B, 8 October 2014 (WIN, DFB); Ontario: Frontenac County, 44.27868°N, 76.59344°W, west side of Collins Creek north side of Creeksford Road, Kingston, D.F. Brunton 18,050B, 24 August 2011 (WIN, DFB),; Ontario: Peterborough County, 44.38742°N, 77.97751°W, east side of mill pond by outlet, north side of High- way 7, east end of Norwood, Asphodel Township, D.F. Brunton 19,917, 6 September 2017 (DFB); Ontario: Peterborough County, 44.76643°N, 78.49064°W, west side of White’s Lake Road, north side of Cry- stal Lake, Galway & Cavendish Township, D.F. Brunton and K.L. McIntosh 17,986B, 312 July 2011 (WIN, UBC, DFB),; Ontario: Renfrew County, 45.5128°N, 76.7591°W, 650 m north of Highway 60 at southeast corner of Clubhouse Lake, west side of Renfrew Golf Club, Horton Geographic Township, D.F. Brunton and K. Fleming 19,873, 27 August 2017 (DFB). Literature Cited Thiers, B. [continuously updated]. Index herbariorum: a glo- bal directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Accessed 15 March 2019. http://sweetgum.nybg.org/science/ih/ The Canadian Field-Naturalist Note Sighting rates and prey of Minke Whales (Balaenoptera acutorostrata) and other cetaceans off Cormorant Island, British Columbia JARED R. TOWERS!”*, CHRISTIE J. MCMILLAN!, and REBECCA S. PIERCEY? y) 'Marine Education and Research Society, P.O. Box 1347, Port McNeill, British Columbia VON 2RO Canada *Bay Cetology, P.O. Box 554, Alert Bay, British Columbia VON 1A0 Canada >Hakai Institute, P.O. Box 309, Heriot Bay, British Columbia VOP 1HO Canada “Corresponding author: jrtowers@gmail.com Towers, J.R., C.J. McMillan, and R.S. Piercey. 2019. Sighting rates and prey of Minke Whales (Balaenoptera acutorostrata) and other cetaceans off Cormorant Island, British Columbia. Canadian Field-Naturalist 133(2): 144-150. https://do1. org/10.22621/cfn.v13312.2103 Abstract From June to August 2012, we conducted over 500 h of visual surveys from Cormorant Island, British Columbia, to determine behaviour and habitat use patterns of nearby cetaceans. Seven species were documented, but Minke Whales (Balaenoptera acutorostrata) were by far the most common and were observed lunge feeding at the surface on 15 occa- sions. In addition, this species was documented surface lunge feeding on Pacific Herring (C/upea pallasi) and Pacific Sand Lance (Ammodytes personatus) on 32 occasions during vessel-based cetacean surveys around Cormorant Island between 2010 and 2014. Although Minke Whales are relatively uncommon in British Columbia, these results indicate that they can regularly be found in specific feeding areas during the summer. Key words: Minke Whale; Balaenoptera acutorostrata, habitat use; feeding ground; Pacific Herring; Clupea pallasi; Pacific Sand Lance; Ammodytes personatus, Cormorant Island; British Columbia The coastal waters of the eastern North Pacific are home to several species of cetaceans. Many have been exploited by humans and conservation concerns have made them the focus of extensive field studies in re- cent decades (Ford 2014). Important foraging habitats considered critical for recovery have been identified for depleted populations of Killer Whale (Orcinus orca), Humpback Whale (Megaptera novaeangliae), and Fin Whale (Balaenoptera physalus), and some populations of these species have become increas- ingly abundant and widespread in coastal waters in recent years (Ford et al. 2009, 2013, 2017; Nichol et al. 2018; Towers et al. 2015, 2018, 2019). Although the distribution and behaviour of less common cetacean species have been recorded (Ford 2014), the foraging ecology and habitat use patterns of some, such as Minke Whale (Balaenoptera acutorostrata), remain poorly understood. Minke Whale is a small, migratory baleen whale that normally occurs in coastal waters of the eastern North Pacific between spring and fall (Towers et al. 2013). Despite a lack of human exploitation history in these waters, Minke Whales are relatively rare along the west coast of North America. Ship-based cetacean surveys between California and Washington have documented Minke Whales 18 times between 1991 and 2008 (Barlow and Forney 2007; Barlow 2010). Further north, in the coastal waters of British Columbia, sur- veys conducted by Williams and Thomas (2007) and Ford et al. (2010) detected only a combined total of 35 Minke Whales during 34 290 km of survey effort. Directed photo-identification research on Minke Whales in British Columbia and Washington has also documented relatively few unique individuals with totals of only 38 identified between 1977 and 1987 (Dorsey et al. 1990) and 44 between 2005 and 2012 (Towers et al. 2013). Photo-identification data have not been compared between these periods, but at least one individual identified in the 1980s was also docu- mented in the 2010s (J.R.T. unpubl. data). Several other individuals documented on more than one oc- casion show high degrees of inter- and intra-annual fi- delity to certain coastal areas over long periods (Dor- sey et al. 1990; Towers et al. 2013). 144 ©The Ottawa Field-Naturalists’ Club 2019 The feeding behaviour of Minke Whales can be difficult to determine because most of it occurs below the water surface. However, in Washington, Minke Whales have been reported surface feeding on Pacific Herring (Clupea pallasi) and Pacific Sand Lance (Ammodytes personatus) on 10 and two occasions, respectively (Hoelzel et al. 1989). Few data on Minke Whale prey have been reported in western Canada, but Minke Whales have also been observed lunge feeding at the surface around Cormorant Island off northeastern Vancouver Island (Towers ef al. 2013). The waters around Cormorant Island are relatively shallow compared to depths of other nearby water- ways and the substrate is made up of glaciofluvial grav- els and sand. Large tidal fluctuations (5 m) and varied bathymetry in this area result in strong currents (<<10 km/h = <5 kts with anything over 2 kts generally con- sidered to have an impact on the movements and behav- iour of most commercial and recreational vessel traffic as well as most fish and marine mammals) and upwell- ings. These environmental variables combine to create favourable habitat for an array of marine species, in- cluding several cetaceans in addition to Minke Whales. 50°38'N 5O°S4 Observation site = Visibility range Predation events: @ Herring O Sand lance a ferea TOWERS ET AL.: MINKE WHALE SIGHTINGS AND PREY 145 To study the behaviour and occurrence of these species non-invasively, we conducted systematic shore-based cetacean observations from Cormorant Island in the summer of 2012 while making concur- rent underwater acoustic recordings. This note pre- sents the visual results of this study combined with data from surface predation events by Minke Whales that were opportunistically documented from small research vessels around Cormorant Island from 2010 to 2014. The acoustic results of this study are reported in Nikolich and Towers (2018). Visual observations of cetaceans were made from the north shore of Cormorant Island at 50°36.140'N, 126°56.820'W (Figure 1). Observers used naked eyes and 15 x 80 binoculars (Steiner, Greely, Colorado, USA) with built in magnetic (M) compass and ret- icle bar mounted on a leveled tripod (Velbon, To- kyo, Japan) to detect cetaceans over an arc of 160° (272°M-—072°M). Visual surveys were conducted by two or more alternating observers for up to 13 h/day when Beaufort sea state levels were <2 (see http:// www.wdcs.org/submissions_bin/WDCS_Shore watch_Seastate.pdf for explanation of Beaufort sea 126°54 126°48'W Ficure 1. The study area off northeastern Vancouver Island, with a. showing the point on Cormorant Island from which observations were made, the arc of visibility, and the locations of Minke Whale (Balaenoptera acutorostrata) predation events and prey species documented from research vessels from 2010 to 2014 in relation to b. the south and central coasts of British Columbia. 146 states). Data, including Beaufort sea state, tide height, and notes on visibility, were recorded every 30 min- utes of survey effort. The time, compass bearing, and reticle distance were recorded each time a cet- acean surfaced (Nikolich and Towers 2018), but when Minke Whales and other species were present in the study area, priority was given to recording the surfa- cings of Minke Whales. The behaviour of cetaceans was noted when apparent, and an effort was made to identify individual whales visually when they were within range. Individual Minke Whale identifica- tions were based on unique natural markings as de- scribed and shown in Towers (2011). To reduce any biases arising from animals being missed when sight- ing effort was focussed in another section of the study area, cetacean occurrence is portrayed in this paper as presence per unit effort (PPUE). A unit of effort was Classified as 55-60 minutes of uninterrupted vis- ual surveys and PPUE was defined as the number of effort units that the species was present divided by the total number of effort units. Between 11 June and 15 August 2012, weather con- ditions allowed for observers to make shore-based vis- ual surveys on 54 days: 20 in June, 19 in July, and 15 in August (Figure 2). In total, 513 units of visual survey effort were conducted. Seven species of cet- aceans were documented: Minke Whale, Humpback Whale, Fin Whale, Bigg’s Killer Whale (Orcinus orca, also known as West Coast Transient popula- tion), Dall’s Porpoise (Phocoenoides dalli), Harbour Porpoise (Phocoena phocoena), and Pacific White- sided Dolphin (Lagenorhynchus obliquidens,; Figure 2). Minke Whales had the highest PPUE (0.44), as they were observed during 224 units of effort (Figure 2; Table 1). Six previously known and individually recognizable Minke Whales were visually identi- fied (M001, M002, M003, M004, M006, and M022; Towers 2011). Harbour Porpoises were the second most commonly present species in the study area fol- lowed by Humpback Whales and Dall’s Porpoises (Figure 2; Table 1). The species with the lowest PPUE was Fin Whale (Figure 2; Table 1). All species were documented transiting; although most species likely foraged at depth in the study area, only Minke Whales, THE CANADIAN FIELD-NATURALIST Vol. 133 Humpback Whales, and Bigg’s Killer Whales were observed feeding at the surface. Bigg’s Killer Whales were documented preying on a Dall’s Porpoise on one occasion, and Minke and Humpback Whales were observed feeding on small schooling fish on 15 and two occasions, respectively. No relation between tidal activity and the foraging behaviour or occurrence of any species was detected. Between 10 September 2010 and 7 August 2014, predation at the surface by Minke Whales around Cor- morant Island was also opportunistically documented from small research vessels on 32 occasions (Figure 3; Table 2). Prey species were visually identified from photographs, video, or samples of remains collected near the surface with a dip-net after the feeding event. Of the predation events, 20 were on Pacific Herring and 12 were on Pacific Sand Lance (Figures | and 3; Table 2). Of the six photo-identified Minke Whales involved in these events (Table 2), five were visually documented during the shore-based component of this study. — Effort +++ HP -- HW -— BKW MW — DP - PWSD FW Units of survey effort T 30-Jul T 15-Jul Date FIGURE 2. Units of survey effort and the presence per unit effort (PPUE) of each species of cetacean sighted between 11 June and 15 August 2012 off Cormorant Island, British Columbia. MW = Minke Whale (Balaenoptera acuto- rostrata), HP = Harbour Porpoise (Phocoena phocoena), HW = Humpback Whale (Megaptera novaeangliae), DP = Dall’s Porpoise (Phocoenoides dalli), PWSD = Pacific White-sided Dolphin (Lagenorhynchus obliquidens), BK W = Bigg’s Killer Whale (Orcinus orca), and FW = Fin Whale (Balaenoptera physalus). TABLE 1. Occurence of cetaceans and presence per unit effort (PPUE) based on sightings from the north shore of Cormorant Island, British Columbia, 11 June to 15 August 2012. MW HP HW Days observed 45 39 14 Sightings ste 109 M3 Units present 224 80 DT, PPUE 0.44 0.16 0.05 BP. PWSD BKW FW 20 5 4 1 30 23 40 3 23 8 8 2 0.04 0.02 0.02 0 Note: BKW =Bigg’s Killer Whale (Orcinus orca), DP = Dall’s Porpoise (Phocoenoides dalli), FW = Fin Whale (Balaenoptera physalus), HP = Harbour Porpoise (Phocoena phocoena), HW = Humpback Whale (Megaptera novaeangliae), MW = Minke Whale (Balaenoptera acutorostrata), and PWSD = Pacific White-sided Dolphin (Lagenorhynchus obliquidens). TOWERS ET AL.: MINKE WHALE SIGHTINGS AND PREY Figure 3. Minke Whale (Balaenoptera acutorostrata) MO01 lunging on a. juvenile Pacific Herring (C/upea pallasi) on 18 July 2011 and b. juvenile Pacific Sand Lance (Ammodytes personatus) on 11 June 2014. Photos: Jared Towers. Of the other species documented in this study, all but Bigg’s Killer Whales are also known to feed on Pacific Herring (Walker et al. 1998; Morton 2000; Ni- chol et al. 2013; McMillan et al. 2018; Towers et al. 2018). Among them, at least the second, third, and fourth most commonly observed species in this study, Harbour Porpoise, Humpback Whale, and Dall’s Por- poise also feed on Pacific Sand Lance (Walker et al. 1998; Nichol et a/. 2013; Marine Education and Re- search Society unpubl. data). All species documented are relatively common in many coastal regions of the eastern North Pacific with the exception of Fin Whale. This species is rarely observed in the inshore waters around Vancouver Island and the individual 148 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Dates of foraging events by identified Minke Whales (Balaenoptera acutorostrata) around Cormorant Island, British Columbia, from 2010 to 2014, and prey species consumed: Pacific Herring (Clupea pallasi) and Pacific Sand Lance (Ammodytes personatus). Date Individual whale Prey species 10 Sep. 2010 M004 Pacific Herring 10 Sep. 2010 M006 Pacific Herring 18 Jul. 2011 MO01 Pacific Herring 27 Jul. 2011 M006 Pacific Herring 1 Aug. 2011 M022 Pacific Herring 18 Sep. 2011 M007 Pacific Herring 18 Sep. 2011 M007 Pacific Herring 15 Jul. 2012 MOOI Pacific Sand Lance 15 Jul. 2012 M004 Pacific Sand Lance 15 Jul. 2012 MOOl1 Pacific Sand Lance 15 Jul. 2012 M002 Pacific Sand Lance 16 Jul. 2012 MOOI Pacific Sand Lance 16 Jul. 2012 M004 Pacific Sand Lance 18 Jul. 2012 M004 Pacific Sand Lance 18 Jul. 2012 M004 Pacific Sand Lance 18 Jul. 2012 M004 Pacific Sand Lance observed is likely the same one photographed nearby the previous day as reported in Towers ef al. (2018). Although the species observed in this study are widely distributed throughout the eastern North Pa- cific (Ford 2014), the high rates of Minke Whale sight- ings compared with other cetacean species and numer- ous predation events recorded confirm this area as an important summer feeding ground for some individual Minke Whales that show intra- and inter-annual fidel- ity to these waters. To our knowledge, no other shore- based cetacean study in the eastern North Pacific has documented such high rates of Minke Whale occur- rence, and the feeding observations reported here are among the first of this species in western Canada. The environmental variables in the waters around Cormorant Island that create favourable habitat for Minke Whales and their prey are not unlike those documented in Minke Whale habitat in other regions. Minke Whale occurrence is positively correlated with shallow water that is near deeper water in Washington (Hoelzel et al. 1989) and Scotland (Robinson ef al. 2009). The dynamic bottom topography of such areas often result in strong currents that drive upwelling and increase ocean productivity (Croll et al. 2005; Tynan et al. 2005). In addition, the sand and gravel substrate around Cormorant Island and in other areas where Minke Whales are often found in the west- ern and eastern North Atlantic (Naud ef al. 2003; MacLeod et al. 2004; Robinson et al. 2009; de Boer 2010) can provide suitable burrowing or rearing habi- tat for Minke Whale prey, such as Pacific Sand Lance (Bizzarro et al. 2016) and Pacific Herring (Hay and McCarter 2006), respectively. Further studies on Minke Whales off Cormorant Island could focus on relations between their occurrence and temporal fluc- Date Individual whale Prey species 18 Jul. 2012 M004 Pacific Sand Lance 20 Jul. 2012 M022 Pacific Herring 25 Jul. 2012 MOOI Pacific Herring 26 Aug. 2012 M006 or M004 si Pacific Herring 17 Jun. 2013 MOOI Pacific Herring 17 Jun. 2013 MOOI Pacific Herring 17 Jun. 2013 MOOI Pacific Herring 17 Jun. 2013 MOOI Pacific Herring 11 Sep. 2013 MOOI Pacific Herring 11 Jun. 2014 MOO] Pacific Sand Lance 11 Jun. 2014 MOO] Pacific Sand Lance 14 Jun. 2014 M006 Pacific Herring 1 Jul. 2014 M022 Pacific Herring 1 Jul. 2014 M022 Pacific Herring 31 Jul. 2014 M006 Pacific Herring 7 Aug. 2014 MOOI Pacific Herring tuations in the abundance and distribution of their prey and other environmental variables. Author Contributions Writing — Original Draft: J.T.; Writing — Review & Editing: JT., C.M., and R.P.; Conceptualization: J.T.; Investigation: J.'T., C.M., and R.P.; Methodology: J.T. and C.M.; Formal Analysis: J.T., C.M., and R.P.; Funding Acquisition: J.T. Acknowledgements This research was funded in part by Mountain Equipment Co-op. We thank Debra Hughes, Jo Mro- zewski, Leticiaa Legat, Bart Willis, Angelica Rose, Ivan Ng, Samuel Ng, Joe Ng, and Nicole Borowczak for helping with the collection of field data, Bill Her- bert and Wendy Thompson for supplying some ma- terials, Dave Towers for logistical support, John Ford for input toward study design, and George Speck for permission to conduct this study from ’Namgis First Nation land. Literature Cited Barlow, J. 2010. Cetacean abundance in the California current estimated from a 2008 ship-based line-transect survey. Technical memorandum NOAA-TM-NMFS-SWESC- 456. United States Department of Commerce, National Oceanic and Atmospheric Administration, La Jolla, Cali- fornia, USA. Barlow, J., and K. Forney. 2007. Abundance and popula- tion density of cetaceans in the California current eco- system. Fishery Bulletin 105: 509-26. Bizzarro, J.J., A.N. Peterson, J.M. Blaine, J.P. Balaban, H.G. Greene, and A.P. Summers. 2016. Burrowing behaviour, habitat, and functional morphology of the Pacific sand lance (Ammodytes personatus). Fishery Bul- letin 114: 445—460. https://do1.org/10.7755/fb.114.4.7 2019 Croll, D.A., B. Marinovic, S. Benson, F.P. Chavez, N. Black, R. Ternullo, and B.R. Tershy. 2005. From wind to whales: trophic links in a coastal upwelling system. Marine Ecology Progress Series 289: 117-130. https:// doi.org/10.3354/meps289117 de Boer, M.N. 2010. Spring distribution and density of min- ke whale Balaenoptera acutorostrata along an offshore bank in the central North Sea. Marine Ecology Pro- gress Series 408: 265-274. https://doi.org/10.3354/meps 08598 Dorsey, E.M., S.J. Stern, A.R. Hoelzel, and J. Jacobsen. 1990. Minke whale (Balaenoptera acutorostrata) from the west coast of North America: individual recognition and small-scale site fidelity. Report of the International Whaling Commission (special issue) 12: 357-368. Ford, J.K.B. 2014. Marine Mammals of British Columbia. Royal BC Museum, Victoria, British Columbia, Canada. Ford, J.K.B., R.M. Abernethy, A.V. Phillips, J. Calam- bokidis, G.M. Ellis, and L.M. Nichol. 2010. Distri- bution and relative abundance of cetaceans in western Canadian waters from ship surveys, 2002—2008. Cana- dian technical report of fisheries and aquatic sciences 2913. Fisheries and Oceans Canada, Nanaimo, British Columbia, Canada. Ford, J.K.B., J.F. Pilkington, A. Reira, M. Otsuki, B. Gisborne, R.M. Abernethy, E.H. Stredulinsky, J.R. Towers, and G.M. Ellis. 2017. Habitats of special im- portance to resident Killer Whales (Orcinus orca) off the west coast of Canada. Canadian Science Advisory Secretariat research document 2017/035. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Ford, J.K.B., A.L. Rambeau, R.M. Abernethy, M.D. Boogaards, L.M. Nichol, and L.D. Spaven. 2009. An assessment of the potential for recovery of Humpback Whales off the Pacific coast of Canada. Canadian Science Advisory Secretariat research document 2009/015. Fish- eries and Oceans Canada, Ottawa, Ontario, Canada. Ford, J.K.B., E.H. Stredulinsky, J.R. Towers, and G.M. Ellis. 2013. Information in support of the identification of critical habitat for transient Killer Whales (Orcinus orca) off the west coast of Canada. Canadian Science Advisory Secretariat research document 2012/155. Fish- eries and Oceans Canada, Ottawa, Ontario, Canada. Hay, D.E., and P.B. McCarter. 2006. Herring spawning areas of British Columbia: a review, geographic analysis and classification. Fisheries and Oceans Canada, Pacific Biological Station, Nanaimo, British Columbia, Canada. Hoelzel, A.R., E.M. Dorsey, and S.J. Stern. 1989. The for- aging specializations of individual minke whales. Ani- mal Behaviour 38: 786—794. https://doi.org/10.1016/s00 03-3472(89)801 11-3 Macleod, K., R. Fairbairns, A. Gill, B. Fairbairns, J. Gordon, C. Blair-Myers, and E.C.M. Parsons. 2004. Seasonal distribution of minke whales Balaenoptera acutorostrata in relation to physiography and prey off the Isle of Mull, Scotland. Marine Ecology Progress Series 277. 263-274. https://doi.org/10.3354/meps277263 McMillan, C.J., J.R. Towers, and J. Hildering. 2018. The innovation and diffusion of “trap feeding,” a novel humpback whale foraging strategy. Marine Mammal Science 35: 779-796. https://doi.org/10.1111/mms.12557 TOWERS ET AL.: MINKE WHALE SIGHTINGS AND PREY 149 Morton, A. 2000. Occurrence, photo-identification and prey of Pacific white-sided dolphins (Lagenorhyncus obli- quidens) in the Broughton Archipelago, Canada 1984— 1998. Marine Mammal Science 16: 80—93. https://do1. org/10.1111/j.1748-7692.2000.tb00905.x Naud, M., B. Long, J. Bréthes, and R. Sears. 2003. In- fluences of underwater bottom topography and geomor- phology on minke whale (Balaenoptera acutorostrata) distribution in the Mingan Island (Canada). Journal of the Marine Biological Association of the United King- dom 83: 889-896. http://doi.org/dskq9c Nichol, L.M., R.M. Abernethy, B.M. Wright, S. Heaslip, L.D. Spaven, J.R. Towers, J.F. Pilkington, E.H. Stredulinsky, and J.K.B. Ford. 2018. Distribution, movements and habitat fidelity patterns of Fin Whales (Balaenoptera physalus) in Canadian Pacific Waters. Canadian Science Advisory Secretariat research docu- ment 2017/004. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Nichol, L.M., A.M. Hall, G.M. Ellis, E. Stredulinsky, M. Boogaards, and J.K.B. Ford. 2013. Dietary over- lap and niche partitioning of sympatric harbor por- poises and Dall’s porpoises in the Salish Sea. Progress in Oceanography 115: 202-210. https://doi.org/10.1016/j. pocean.2013.05.016 Nikolich, K., and J.R. Towers. 2018. Vocalizations of common minke whales (Balaenoptera acutorostrata) in an eastern North Pacific feeding ground. Bioacoustics. https://doi.org/10.1080/09524622.2018.1555716 Robinson, K.P., M.J. Tetley, and E.G. Mitchelson-Jacob. 2009. The distribution and habitat preference of coast- ally occurring minke whales (Balaenoptera acutoro- strata) in the outer southern Moray Firth, northeast Scotland. Journal of Coastal Conservation 13: 39-48. https://doi.org/10.1007/s11852-009-0050-2 Towers, J.R. 2011. Minke Whales of the Straits off North- eastern Vancouver Island (Second Edition). Marine Edu- cation and Research Society, Alert Bay, British Colum- bia, Canada. Towers, J.R., G.M. Ellis, and J.K.B. Ford. 2015. Photo- identification catalogue and status of the northern resi- dent Killer Whale population in 2014. Canadian tech- nical report of fisheries and aquatic sciences 3139. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Towers, J.R., M. Malleson, C.J. McMillan, J. Cogan, S. Berta, and C. Birdsall. 2018. Occurrence of fin whales (Balaenoptera physalus) between Vancouver Island and continental North America. Northwestern Naturalist 99: 49-57. https://doi.org/10.1898/nwn17-16.1 Towers, J.R., C.J. McMillan, M. Malleson, J. Hildering, J.K.B. Ford, and G.M. Ellis. 2013. Seasonal move- ments and ecological markers as evidence for migra- tion of common minke whales photo-identified in the eastern North Pacific. Journal of Cetacean Research and Management 13: 221-229. Towers, J.R., G.J. Sutton, T.J.H. Shaw, M. Malleson, D. Matkin, B. Gisborne, J. Forde, D. Ellifrit, G.M. Ellis, J.K.B. Ford, and T. Doniol-Valcroze. 2019. Photo- identification catalogue, population status, and distri- bution of Bigg’s killer whales known from coastal wa- ters of British Columbia, Canada. Canadian technical 150 report of fisheries and aquatic sciences 3311. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Tynan, C.T., D.G. Ainley, J.A. Barth, T.J. Cowles, S.D. Pierce, and L.B. Spear. 2005. Cetacean distributions relative to ocean processes in the northern California Current system. Deep-Sea Research II 52: 145-167. https://doi.org/10.1016/j.dsr2.2004.09.024 Walker, W.A., M.B. Hanson, R.W. Baird, and T.J. Guenther. 1998. Food habits of the harbor porpoise, Phocoena phocoena, and the Dall’s porpoise, Phocoe- noides dalli, in the inland waters of British Columbia and THE CANADIAN FIELD-NATURALIST Vol. 133 Washington. Pages 63—75 in Marine Mammal Protection Act and Endangered Species Act Implementation Pro- gram 1997. AFSC processed report 98-10. National Ma- rine Fisheries Service, Seattle, Washington, USA. Williams, R., and L. Thomas. 2007. Distribution and abundance of marine mammals in the coastal wa- ters of British Columbia, Canada. Journal of Cetacean Research and Management 9: 15-28. Received 11 June 2018 Accepted 17 January 2019 The Canadian Field-Naturalist Use of salmon (Oncorhynchus spp.) by Brown Bears (Ursus arctos) in an Arctic, interior, montane environment MatTHew S. Sorum!", KYLE JOLy', and MATTHEW D. CAMERON! 'National Park Service, Gates of the Arctic National Park and Preserve, 4175 Geist Road, Fairbanks, Alaska 99709 USA “Corresponding author: mathew_sorum@nps.gov Sorum, M.S., K. Joly, and M.D. Cameron. 2019. Use of salmon (Oncorhynchus spp.) by Brown Bears (Ursus arctos) in an Arctic, interior, montane environment. Canadian Field-Naturalist 133(2): 151-155. https://doi.org/10.22621/cfn. v13312.2114 Abstract Salmon (Oncorhynchus spp.) is a key dietary item for temperate coastal Brown Bears (Ursus arctos) across much of their circumpolar range. Brown Bears living in Arctic, interior, and montane environments without large annual runs of salmon tend to be smaller bodied and occur at much lower densities than coastal populations. We conducted ground and aerial sur- veys to assess whether Brown Bears fished for salmon above the Arctic Circle, in and around Gates of the Arctic National Park and Preserve. Here, we document the use of salmon by interior Brown Bears in the Arctic mountains of the central Brooks Range of Alaska. We believe our findings could be important for understanding the breadth of the species’ diet across major biomes, as well as visitor safety in the park and Brown Bear conservation in the region. Key words: Alaska; Brown Bear; diet; fishing; mountains; Oncorhynchus spp.; salmon; Ursus arctos, Brooks Range Introduction In temperate environments, Brown Bears (Ursus arctos) feed extensively on salmon (Oncorhynchus spp.; Hilderbrand et al. 1999a; Mowat and Heard 2006). The abundance of salmon in some river sys- tems and their high nutritional value allow Brown Bears in these areas to grow much larger and live at much higher densities than Brown Bear populations without predictable access to seasonal marine resour- ces (Hilderbrand et al. 1999a,b; Mowat and Heard 2006). For example, Brown Bears living in interior re- gions are often much smaller than coastal conspecif- ics and occur at much lower densities (Hildebrand et al. 2018a,b). Substantively lower resource availability is thought to be a major determinant of both body size and population density (Hilderbrand et al. 1999a,b; Mowat and Heard 2006). Bears use terrestrial protein sources (Caribou [Rangifer tarandus], Arctic Ground Squirrels [Spermophilus parryii], and Moose [Alces americanus]), aS well as plants, nuts, and other foods when access to marine resources, such as salmon, is limited (Welch et al. 1997; Rode et al. 2001; Gau et al. 2002; Mowat and Heard 2006), in areas such as the Arctic, interior, montane environments of the central Brooks Range of Alaska. Reports from local aircraft pilots of Brown Bears congregating along Arctic rivers during August and September have been received by the National Park 151 Service (NPS) as early as 2008, although use of sal- mon by Brown Bears has never been reported in the central Brooks Range or in many other Arctic regions (e.g., MacHutchon and Wellwood 2003; Mowat and Heard 2006). In 2014, the NPS initiated a Brown Bear monitoring project. Global positioning system (GPS) data collected by that project indicated that Brown Bears in the interior mountains of the Brooks Range did indeed spend extended periods along larger river corridors from July through September. Our goal was to document the timing and location of salmon use by Brown Bears in the central Brooks Range by direct observation. A better understand- ing of Brown Bear resource use in the region could be useful in determining the breadth of the species’ diet across major biomes, as well as locally, for visitor management and Brown Bear conservation. Methods We conducted a survey of Brown Bears fishing in and around Gates of the Arctic National Park and Preserve (GANPP; Figure 1). GANPP encompasses an interior Arctic ecosystem characterized by the mountainous terrain of the Brooks Range, exten- sive spruce (Picea spp.) forests and lowland ripar- ian areas on the range’s southern flanks, and tundra on its northern side (Wilson et al. 2014). The head- waters of three major river systems begin in GANPP: ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 152 Chukchi Sea ud ae : ns Pa a 156°W THE CANADIAN FIELD-NATURALIST Vol. 133 + 67°N t66°N 152°W Ficure 1. Study area (black oval) in the central Brooks Range mountains in Arctic Alaska. the Kobuk, Koyukuk, and Noatak rivers. The Kobuk River drains west about 550 km (river length) to the Chukchi Sea (part of the Arctic Ocean) with most of the major tributaries arising in the southern Brooks Range. The Koyukuk River, a major tributary of the Yukon River, drains southwest for 1870 river-km from GANPP to the Bering Sea. The Noatak River drains northwest about 660 river-km to the Chukchi Sea, with all of its major tributaries arising in the northern Brooks Range. Site-specific salmon escapement numbers are not available for our study areas, as these counts usu- ally focus on commercial fisheries in salt water and specific freshwater streams important for sport or subsistence fishing. However, each of GANPP’s three main rivers has runs of Chum Salmon (Oncorhynchus keta) and Chinook Salmon (Oncorhynchus tshawy- tscha) from mid-July to early September (O’Brien and Berkgiler 2005). We visited one tributary in each of GANPP’s three main river systems to document use of sal- mon by Brown Bears in the region. The specific lo- cations visited were determined from GPS collared bears (n = 33) or reports by pilots and were access- ible by wheeled or float plane. (For more information about our collaring efforts, which were in accord- ance with the guidelines of the American Society of Mammalogists [Sikes and Gannon 2011] and approved by United States Geological Survey and NPS Institutional Animal Care and Use Committees [I[ACUCs; 2014-1 and 2014A2, respectively], see Hil- derbrand ef al. 2018a.) In 2016, we surveyed 8 km of the Kobuk River system (Table 1). In 2017, we surveyed 2 km of the Noatak River system and 8 km of the Koyukuk River system (Table 1). Our surveys involved walking along river banks and/or floating down streams in a raft to identify signs of bears fishing (e.g., fish carcass, scat, or observations of bears fishing) and document the presence of salmon species. Along the Noatak River system, we used a vantage point above an area that we suspected bears used for fishing to look for bears using a 20 x 60 spotting scope. Results During each stream visit, we observed bears fish- ing or identified recent signs of fishing activity (i.e., salmon gill plate, mandible, carcass, or bear scat with fish remains in it; Table 1). On the Kobuk River sys- tem, 17 August 2016, we found a large pile of fresh gill plates (Figure 2) along a sharp bend in the river where a deep pool had formed. Although we did not observe a bear fishing, we surmised that bears were diving for the salmon carcasses that we observed lying at the bottom of the pool. On the Koyukuk River system, 2 August 2017, we observed an adult male fishing by “snorkeling” up- stream in ~1.5 m of water and another adult male fish- 2019 SORUM ET AL.. BROWN BEARS’ USE OF SALMON 153 TABLE 1. Observations of Brown Bears (Ursus arctos) using salmon (Oncorhynchus spp.) in the central Brooks Range, Alaska, 2016-2017. River system Observation Dates Kobuk Salmon gill plates 17 August 2016 Fresh bear tracks 1-4, 17, 29 August 2016 9-10 September 2016 1 bear* 29 August 2016 Koyukuk 1 bear* 4 August 2016 1 male bear 2 August 2017 1 male bear 2 August 2017 1 male bear 21 August 2017 Salmon gill plates 22 August 2017 Noatak 3 bears 28 July 2017 11 bears 29 August 2017 1 male bear 30 August 2017 Notes Fresh pile of plates found adjacent to a deep pool formed at bend in river Single bears and family groups Adult fishing near Chum Salmon Adult fishing near Chum Salmon Large male snorkeling upstream in 1.5 m of water Large male fishing along a shallow side stream Large male fishing along a shallow portion of river Fresh pile of plates found along bank of river A Brown Bear family group fishing at the confluence 8 independent bears and | sow with 2 2-year-old cubs fishing along 2 km of river Male bear fishing along a shallow side stream *Aerial survey; all other observations were made during ground-based surveys. ing along a shallow (<0.5 m) portion of the stream. On 29 August 2017, we spent 4 h observing bears fishing from a vantage point above the Noatak River. We observed 11 bears (eight independent adults and one female with two 2-year-old cubs) using ~2 km of stream. Each adult was observed catching at least one salmon. Most bears either walked upstream along the bank or in the stream to locate and catch spawning Chum Salmon (Figure 3). All bears fished along a FiGurE 2. Salmon remains (jaws and gill plates) found along a tributary of the Koyukuk River, Alaska, 22 August 2017. Piles such as these were common along well-worn game trails at the river edge. Photo: Matthew Cameron, National Park Service. small (<250 m) section of stream and were often close to other bears. We also documented the presence and spawning activity of Chum Salmon at each stream. Discussion Our surveys substantiate earlier reports that inter- ior, montane Brown Bears in the Arctic fish for sal- mon. Although the Brown Bears we studied lived about 400 km inland (or 550-1800 river-km from the coast), Chum Salmon were present and used by Brown Bears. Our findings build on past research that evaluated broad diet patterns of Brown Bears across North America (Mowat and Heard 2006). Even though salmon occur in streams throughout the Brooks Range, Mowat and Heard (2006) reported no use of salmon by Brown Bears in the central Brooks Range. Our observations of Brown Bears fishing for salmon across multiple river systems throughout the central Brooks Range provide evidence that salmon are not only used by bears here, but are likely an im- portant seasonal food resource in the region. Many interior populations of Brown Bears rely solely on terrestrial food resources, such as green vegetation, berries, and ungulates to satisfy their nu- tritional requirements (Gau ef al. 2002; MacHutchon and Wellwood 2003; Mowat and Heard 2006). How- ever, if available, some interior bear populations con- sume salmon (Belant et a/. 2006). In the Arctic inter- ior, food is even more limited for bears, as the growing season is short and ungulate densities are quite low and sparsely distributed over vast areas (Gasaway et al. 1992). Thus, where salmon resources are available, they likely play an important dietary role for bears liv- ing in the Arctic interior and may alter their distribu- tion, body size, and population density (Hilderbrand et al. 1999a:; Deacy et al. 2016, 2019). 154 : f 4 ¥ ‘a at BN eae) BA Sai Nari be teehee THE CANADIAN FIELD-NATURALIST Vol. 133 Figure 3. A Brown Bear (Ursus arctos) successfully acquiring a salmon (Oncorhynchus spp.) from a tributary of the Noatak River, central Brooks Range Alaska, 30 August 2017. Photo: Mathew Sorum, National Park Service. Consumption of salmon by Brown Bears provides a direct avenue for nutrient and energy transfers from marine to lotic to terrestrial systems (Hilderbrand et al. 1999c). Inputs of marine-derived nutrients into a terrestrial system creates cascading effects across trophic levels via increased productivity (Mathewson et al. 2003; Winder et al. 2005), and the effects are likely compounded in a nutrient-limited system, such as GANPP. Ultimately, this relationship cre- ates biological hotspots with higher productivity, species diversity, and richness (Naiman et al. 2002). Identification of these areas is important for consery- ation and preservation of these important ecological systems. For example, areas of congregating Brown Bears in this low-density system may warrant addi- tional hunting and/or visitation restrictions in the fu- ture to avoid overhunting and/or disturbance during the critical period of hyperphagia. Identification and further elucidation of the rela- tion between salmon and bears in interior ecosystems will improve the understanding and management of population dynamics of both predators and their prey. Future research should consider estimating the com- position of salmon in the diet of Brown Bears and the influence of salmon on seasonal distribution and habitat selection patterns of bears. Acknowledgements Funding for this work was provided by the United States National Park Service. We thank pilots E. Sieh, S. Sample, and C. Cebulski for reporting their obser- vations of Brown Bears that helped lead to this study. Dave Gustine and Grant Hilderbrand were instru- mental in Brown Bear collaring efforts. We thank Lavern Beier, Mark Melham, Jared Hughey, and Ken Hill for field assistance. We also thank Thomas Jung, Dwayne Lepitzki, William Leacock, and an anonym- ous reviewer for providing recommendations, which improved this manuscript. Literature Cited Belant, J.L., K. Kielland, E.H. Follmann, and L.G. Adams. 2006. Interspecific resource partitioning in sympatric urs- ids. Ecological Applications 16: 2333-2343. https://doi. org/10.1890/1051-0761(2006)016[2333:irpisu]2.0.co;2 Deacy, W., W. Leacock, J.B. Armstrong, and J.A. Stanford. 2016. Kodiak brown bears surf the salmon red wave: direct evidence from GPS collared individuals. Ecology 97: 1091-1098. https://doi.org/10.1890/15- 1060.1 Deacy, W.W., W. Leacock, J.A. Stanford, and J.B. Arm- strong. 2019. Variation in spawning phenology within salmon populations influences landscape-level patterns of brown bear activity. Ecosphere 10: e02575. https:// doi.org/10.1002/ecs2.2575 2019 Gasaway, W.C., R.D. Boertje, D.V. Grangaard, D.G. Kelleyhouse, R.O. Stephenson, and D.G. Larsen. 1992. The role of predation in limiting moose at low densities in Alaska and Yukon and implications for con- servation. Wildlife Monographs 120: 3-59. Gau, R.J., R. Case, D.F. Penner, and P.D. McLoughlin. 2002. Feeding patterns of barren-ground grizzly bears in the central Canadian Arctic. Arctic 55: 339-344. https://doi.org/10.14430/arctic717 Hilderbrand, G.V., D.D. Gustine, B.A. Mangipane, K. Joly, W. Leacock, L.S. Mangipane, J. Erlenbach, M.S. Sorum, M.D. Cameron, J.L. Belant, and T. Cambier. 2018a. Body size and lean mass of brown bears across and within four diverse ecosystems. Journal of Zoology 305: 53-62. https://do1.org/10.1111/jzo. 12536 Hilderbrand, G.V., D.D. Gustine, B. Mangipane, K. Joly, W. Leacock, L. Mangipane, J. Erlenbach, M.S. Sorum, M.D. Cameron, J.L. Belant, and T. Cambier. 2018b. Plasticity in physiological condition of female brown bears across diverse ecosystems. Polar Biology 41: 773-780. https://doi.org/10.1007/s00300-017-2238-5 Hilderbrand, G.V., T.A. Hanley, C.T. Robbins, and C.C. Schwartz. 1999c. Role of brown bears (Ursus arctos) in the flow of marine nitrogen into a terrestrial eco- system. Oecologia 121: 546-550. https://doi.org/10.1007 /s004420050961 Hilderbrand, G.V, S.G. Jenkins, C.C. Schwartz, T.A. Hanley, and C.T. Robbins. 1999b. Effect of seasonal differences in dietary meat intake on changes in body mass and composition in wild and captive brown bears. Canadian Journal of Zoology 77: 1623-1630. https://doi. org/10.1139/z99-133 Hilderbrand, G.V, C.C. Schwartz, C.T. Robbins, M.E. Jacoby, T.A. Hanley, S.M. Arthur, and C. Servheen. 1999a. The importance of meat, particularly salmon, to body size, population productivity, and conservation of North American brown bears. Canadian Journal of Zoology 77: 132-138. https://doi.org/10.1139/cjz-77-1-132 MacHutchon, G.A., and D.W. Wellwood. 2003. Grizzly bear food habits in northern Yukon, Canada. Ursus 14: 225-235. SORUM ET AL.. BROWN BEARS’ USE OF SALMON 155 Mathewson, D.D., M.D. Hocking, and T.E. Reimchen. 2003. Nitrogen uptake in riparian plant communities across a sharp ecological boundary of salmon densi- ty. BMC Ecology 3: 4-15. https://doi.org/10.1186/14 72-6785-3-4 Mowat, G., and D.C. Heard. 2006. Major components of grizzly bear diet across North America. Canadian Journal of Zoology 84: 473—489. https://doi.org/10.1139/ z06-016 Naiman, R.J., R.E. Bilby, D.E. Schindler, and J.M. Helfield. 2002. Pacific salmon, nutrients, and the dyna- mics of freshwater and riparian ecosystems. Ecosystems 5: 399-417. https://doi.org/10.1007/s10021-001-0083-3 O’Brien, J.P., and B.L. Berkgiler. 2005. Abundance and run timing of adult salmon in Henshaw Creek, Kanuti National Wildlife Refuge, Alaska, 2004: 23. Alaska fisheries data series 2005-15. United States Fish and Wildlife Service, Fairbanks, Alaska, USA. Rode, K.D., C.T. Robbins, and L.A. Shipley. 2001. Con- straints on herbivory by grizzly bears. Oecologia 128: 62-71. https://doi.org/10.1007/s004420100637 Sikes, R.S., and W.L. Gannon. 2011. Guidelines of the American Society of Mammalogists for the use of wild mammals in research. Journal of Mammalogy 92: 235— 253. https://doi.org/10.1644/10-mamm-f-355.1 Welch, C.A., J. Keay, K.C. Kendall, and C.T. Robbins. 1997. Constraints on frugivory by bears. Ecology 78: 1105-1119. https://doi.org/10.1890/0012-9658(1997)078 [1105:cofbb]2.0.co;2 Wilson, R.R., D.D. Gustine, and K. Joly. 2014. Evaluating potential effects of an industrial road on winter habitat of caribou in north-central Alaska. Arctic 67: 472-482. https://doi.org/10.14430/arctic4421 Winder, M., D.E. Schindler, J.W. Moore, S.P. Johnson, and W.J. Palen. 2005. Do bears facilitate transfer of salmon resources to aquatic macroinvertebrates? Cana- dian Journal of Fisheries Aquatic Science 62: 2285— 2293. https://doi.org/10.1139/f05-136 Received 24 August 2018 Accepted 30 May 2019 The Canadian Field-Naturalist Note Roman Snail, Helix pomatia (Mollusca: Helicidae), in Canada ROBERT G. ForsytTu'’, and JAMES KAMSTRA” 'New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada 7AECOM, 105 Commerce Valley Drive West, Markham, Ontario L3T 7W3 Canada “Corresponding author: rforsyth.bc.ca@gmail.com Forsyth, R.G., and J. Kamstra. 2019. Roman Snail, Helix pomatia (Mollusca: Helicidae), in Canada. Canadian Field-Naturalist 133(2): 156-159. https://doi.org/10.22621/v13312.2150 Abstract Populations of Roman Snail, Helix pomatia, a large European land snail, are reported for the first time in Canada from disturbed habitats in two distant locations: Sarnia, Ontario and Montrose, British Columbia. As Roman Snail is an edible species subject to international commercial trade, its deliberate, but illegal, introduction into Canada and intentional or unintentional releases are possible sources of these populations. Key words: Mollusca; terrestrial snail; gastropod; new provincial records; Ontario; British Columbia; biogeography The genus Helix L., 1758, in its modern, restricted sense, is a group of large-bodied land snails of Eur- ope, some parts of western Asia, and North Africa (Schileyko 2006; Mumladze et al. 2008; Fiorentino et al. 2016). Larger land snails, including species of the genus Helix and its type species, Helix pomatia L., 1758 (Figure 1), are among those that have been consumed by humans for millennia (Bar 1977; Lubell 2004). The range of H. pomatia, Roman or Burgundy Snail, includes central and southern Europe, but also extends into western and northwestern Europe to coastal areas of countries on the Baltic Sea and west to France and England (Kerney 1999; Neubert 2013, 2014), with the ancestral range in the Balkan region (Fiorentino et al. 2016). The presence of this species in Britain is generally believed to represent a Roman introduction (Taylor 1910; Kerney 1966, 1999); the Romans went so far as importing, breeding, and rear- ing snails in “cochlearia” (Taylor 1900). The original geographic range of H. pomatia may have been central European, with its spread into western Europe a re- sult of ancient introductions by humans (Taylor 1910). Recently, introductions have been reported as far east as European Russia (Sysoev and Schileyko 2009; Egorov 2015). In the United States, perhaps the first es- tablished population was noted in Jackson, Michigan, in 1937 (Archer 1937; Pilsbry 1939), but H. pomatia is now also known from Wisconsin, New York, Mas- sachusetts, Indiana, Pennsylvania, Florida, and Cali- fornia (GBIF 2017; NatureServe 2019). The presence of this species in Canada has gone unreported in the literature, and we document here the occurrence of 7. pomatia in British Columbia and Ontario. Specimens were collected opportunistically by us or our contacts and are vouchered in the Mollusc Collection of the New Brunswick Museum, Saint John (NBM). Shell height (H) and diameter (or width; D) of unbroken, apparently mature specimens were meas- ured using a digital caliper, to the nearest 0.1 mm. Helix pomatia was first observed at Canatara Park in Sarnia, Ontario, on 10 May 2013 by J.K. (Figure la). Three live specimens and one empty shell were collected on 31 May 2013. Canatara is a multi-use municipal park of ~80 ha that is surrounded by urban lands. The south portion (~25 ha), where Roman Snails were found, was once a landfill site, which has re- generated over many decades. It consists of old field, thickets, and woodland with a high proportion of non-native invasive plant species. Roman Snails were found in dry meadow co-dominated by goldenrods (Solidago sp.) and Canada Thistle (Cirsium arvense (L.) Scopoli) with some Garlic Mustard (Alliaria peti- olata (M. Bieberstein) Cavara & Grande). The snails were often seen under debris, such as boards, or climbing on vegetation up to 30 cm above the ground. The non-native, European Grove Snail, Cepaea nemo- ralis (L., 1758), was abundant in Canatara Park. At Montrose, British Columbia (BC), S. Munch (pers. comm. 2015) had observed H. pomatia on his property since moving there in 2002. The habi- 156 ©The Ottawa Field-Naturalists’ Club FORSYTH AND K AMSTRA: ROMAN SNAIL IN CANADA Itsy FicureE 1. Roman Snail, Helix pomatia. a. Canatara Park, Sarnia, Ontario. b. Shells from Sarnia, Ontario (left; NBM 010198) and Montrose, British Columbia (right; NBM 010200). Photo: a. J. Kamstra. Photo: b. R. Forsyth. tat is an open area that includes lawn and tall uncut grass with small Trembling Aspen (Populus tremu- loides Michaux), spruce (Picea A. Dietrich sp.), and Western Red Cedar (Thuja plicata Donn ex D. Don). The maximum number of snails encountered at one time was 15-20 individuals in a 9 x 9 m area. We are also aware that H. pomatia has persisted for several years at Revelstoke, BC (H. Douglas pers. comm. 2014), but have little information about its presence there. No native or introduced land snail in Canada approaches the size of H. pomatia, which is one of the largest of the European Helix species (up to 50 mm; Kerney and Cameron 1979). The only other es- tablished, large helicid species in Canada is Brown Gardensnail, Cornu aspersum (Miller, 1774), more commonly known as Helix aspersa, but that species is easily recognized by its smaller shell, different pat- tern of shell pigmentation, and different sculpture, among other characters (Kerney and Cameron 1979). Helix lucorum L., 1758, which can grow to be larger than H. pomatia, might be confused with H. poma- tia, although it has markedly more prominent band- ing. Indeed, Burke (2013) figured a specimen of H. /ucorum that he misidentified as H. pomatia. Based on our own observations and correspond- ence with the discoverer of the Montrose popula- tion of H. pomatia, we believe that both this and the Sarnia population have persisted for several years (at least 13 years at Montrose) and that this species can be added to the growing list of introduced terrestrial molluscs in Canada. Variation in shell sizes suggests that different generations exist. Throughout its range in Europe, H. pomatia in- habits milder coastal areas as well as mountainous re- gions with more continental climates. Thus, the suc- 158 cessful establishment of this species in Canada is not a surprise. However, as climate change brings milder winters, in the future, we might expect increasingly more successful introductions of terrestrial molluscs in different parts of the country. Novel records of introduced terrestrial molluscs in Canada continue to be discovered, even in areas that are rather well ex- plored (e.g., Forsyth 1999, 2008; Reise et al. 2000; Forsyth et al. 2001, 2016; Maunder ef al. 2017). The discovery of H. pomatia at Montrose and Revelstoke adds a second introduced snail species known from the BC interior but not from the milder south coast region of BC. More surveys for terrestrial molluscs around populated centres, such as those of Forsyth (1999) who focussed on the Vancouver and Victoria regions, would be useful in the BC interior. Similar to BC, southern Ontario has been rather well ex- plored for terrestrial snails. The great extent of modi- fied habitats there allows for the foothold of many synanthropic, introduced species. The possible in- vasiveness of H. pomatia is not known. Apparently the species persists but had not spread far from Jackson, Michigan, even 80 years after it was first discovered (Atkinson 2019). At the Sarnia site, the species has not become appreciably more abundant in the five years since it was first discovered, 1n contrast to the highly invasive Cepaea nemoralis. Most non-native terrestrial snails and slugs are likely accidentally introduced with plants, soil, and debris, or movements of other materials. However, for H. pomatia, the source of the introductions is un- certain. It seems likely that someone raising Roman Snail deliberately or unintentionally released sev- eral individuals. Roman Snail is an edible species that has international commercial value and is sold for both food and the pet trade. It is listed for sale on various websites, such as My Happy Snails (https:// www.myhappysnails.com/), based in Ukraine, which will ship abroad to willing buyers. It is possible that Roman Snail was deliberately imported into Canada, although the importing of any species of Helix into Canada is prohibited (D. Mooi pers. comm. 2019). The Sarnia location is only about 150 km from where H. pomatia occurs in Michigan, but it seems unlikely that the Sarnia snails originated from that location, given the presence of an international border, a water barrier, and the distance. Voucher specimens Canada: Ontario: Lambton County: Sarnia: Cana- tara Park: ca. 43.0°N, 82.4°W, Jeg. James Kamstra, 31 May 2013 (NBM 010198, 1 specimen). British Co- lumbia: Kootenay-Boundary Regional District: Mont- rose, ca. 49.1°N, 117.6°W, /eg. Steven Munch, summer and fall 2015 (NBM 010200, 33 specimens; Figure 1b). British Columbia: Columbia-Shuswap Regional THE CANADIAN FIELD-NATURALIST Vol. 133 District: Revelstoke, 12 June 2014 (NBM 010199, 1 specimen). Canadian shells (Figure la) can be described as follows. The shell is large (H = 33.6—42.0 mm, D = 30.46-39.37 mm; mean H/D = 1.05, SD = 0.05, n = 27) and rather globular with a conical spire. It is pale grey-brown, with lighter and darker colabral streaks and, in general, two to five spiral bands, which are sometimes rather weakly marked or absent. There are ~4 convex whorls, with the last whorl descending in full-grown specimens. The periphery is rounded, medial on the last whorl. The protoconch is smooth. The teleoconch has irregular, low, somewhat riblet- like colabral ridges and spiral rows of weak granules. The aperture is large, subovate-rounded, and show- ing the external shell colour through the shell wall. The outer lip is scarcely thickened and narrowly out- wardly flared. The columellar lip is pinkish-brown, expanded, and almost sealing the umbilicus, which is a narrow slit. Acknowledgements We thank Steven Munch (Montrose, British Co- lumbia) for the donation of Montrose specimens, Michael Oldham (Natural Heritage Information Cen- tre, Peterborough, Ontario) for introducing us and encouraging us to write the manuscript, and Hume Douglas and Diana Mooy (Canadian Food Inspection Agency, Ottawa). We also acknowledge the New Brunswick Museum for its generous contribution to- wards the article publication fee. Literature Cited Archer, A.F. 1937. Helix pomatia Linné in Jackson, Michi- gan. Nautilus 51: 61-63. Atkinson, J.W. 2019. Exotic, invasive and pest snails and land slugs of Michigan. Michigan State University Snail Laboratory, Lansing, Michigan, USA. Accessed 13 March 2019. https://msu.edu/~atkinso9/pestsnailpage.htm. Bar, Z. 1977. Human consumption of land snails in Israel. Basteria 31: 53-58. Burke, T.E. 2013. Land Snails and Slugs of the Pacific Northwest. Oregon State University Press, Corvallis, Oregon, USA. Egorov, R. 2015. Helix pomatia Linnaeus, 1758: the history of its introduction and recent distribution in European Russia. Malacologica Bohemoslavia 14: 91-101. Fiorentino, V., G. Manganelli, F. Giusti, and V. Ket- maier. 2016. Recent expansion and relic survival: phy- logeography of the land snail genus Helix (Mollusca, Gastropoda) from south to north Europe. Molecular Phy- logenetics and Evolution 98: 358-372. https://doi.org/ 10.1016/j.ympev.2016.02.017 Forsyth, R.G. 1999. Distributions of nine new or little- known exotic land snails in British Columbia. Canadian Field-Naturalist 113: 559-568. Accessed 3 October 2019. https://biodiversitylibrary.org/page/34235429, 2019 Forsyth, R.G. 2008. First record of the European land snail Trochulus striolatus in British Columbia, Canada (Pul- monata: Hygromiidae). Festivus 40: 76-78. Forsyth, R.G., J.M.C. Hutchinson, and H. Reise. 2001. Aegopinella nitidula (Draparnaud, 1805) (Gastropoda: Zonitidae) in British Columbia—first confirmed North American record. American Malacological Bulletin 16: 65—69. Forsyth, R.G., J.E. Maunder, D.F. McAlpine, and R.G. Noseworthy. 2016. Distributional status of an intro- duced land snail Discus rotundatus (Rotund Disc, Mol- lusca: Discidae) in Canada. Canadian Field-Naturalist 130: 235-246. https://doi.org/10.22621/cfn.v13013.1887 GBIF (Global Biodiversity Information Facility). 2017. GBIF backbone taxonomy. Checklist dataset. Helix po- matia Linnaeus, 1758. Accessed 9 February 2019. https:// doi.org/10.15468/390mei Kerney, M.P. 1966. Snails and man in Britain. Journal of Conchology 26: 3-14. Kerney, M.P. 1999. Atlas of the Land and Freshwater Mol- luscs of Britain and Ireland. Harley Books, Colchester, Essex, United Kingdom. Kerney, M.P., and R.A.D. Cameron. 1979. A Field Guide to the Land Snails of Britain and North-west Europe. Collins, London, United Kingdom. Lubell, D. 2004. Prehistoric edible land snails in the circum- Mediterranean: the archaeological evidence. Pages 77— 98 in Petits animaux et sociétés humaines du complément alimentaire aux ressources utilitaires. X XIV’ rencontres internationales d’archéologie et d’histoire d’Antibes. Edited by J-J. Brugal and J. Desse. Editions APDCA, Antibes, France. Maunder, J.E., R.G. Noseworthy, J.M.C. Hutchinson, and H. Reise. 2017. Terrestrial molluscs of the Province of Newfoundland and Labrador, Canada. Part 1: Boett- gerillidae. Check List 13: 277-284. https://doi.org/10.155 60/13.4.277 Mumladze, L., D. Tarkhnishvili, and B.M. Pokryszko. 2008. A new species of the genus Helix from the Les- ser Caucasus (SW Georgia). Journal of Conchology 39: 483-487. FORSYTH AND K AMSTRA: ROMAN SNAIL IN CANADA 159 NatureServe. 2019. Helix pomatia— Linnaeus, 1758. Nature Serve Explorer, v. 7.1. Accessed 14 March 2019. http:// explorer.natureserve.org/servlet/NatureServe?searchName =Helix+pomatia. Neubert, E. 2011. Helix pomatia. In The IUCN Red List of Threatened Species. International Union for Conserva- tion of Nature and Natural Resources, Cambridge, Uni- ted Kingdom. e.T156519A4957463. https://doi.org/10.23 05/iucn.uk.2011-1 rlts.t156519a4957463.en Neubert, E. 2014. Revision of He/ix Linnaeus, 1758 in its eastern Mediterranean distribution area, and reassign- ment of Helix godetiana Kobelt, 1878 to Maltzanella Hesse, 1917 (Gastropoda, Pulmonata, Helicidae). Con- tributions to Natural History [Natural History Museum Bern] 26: 1—200. Pilsbry, H.A. 1939. Land Mollusca of North America (North of Mexico), Volume I, Part 1. Monograph 3. Aca- demy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania, USA. Reise, H., J.M.C. Hutchinson, R.G. Forsyth, and T.J. Forsyth. 2000. The ecology and rapid spread of the ter- restrial slug Boettgerilla pallens in Europe with refer- ence to its recent discovery in North America. Veliger 43: 313-318. Schileyko, A.A. 2006. Treatise on Recent terrestrial pul- monate molluscs, part 13. Helicidae, Pleurodontidae, Polygyridae, Ammonitellidae, Oreohelicidae, Thysano- phoridae. Ruthenica Supplement 2: 1765-1906. Sysoev, A., and A. Schileyko. 2009. Land Snails and Slugs of Russia and Adjacent Countries. Pensoft, Sofia, Bulgaria. Taylor, J.W. 1900. A Monograph of the Land and Fresh- water Mollusca of the British Isles. Structural and Ge- neral Volume. Part VI. Taylor Brothers, Leeds, United Kingdom. Taylor, J.W. 1910. A Monograph of the Land and Freshwater Mollusca of the British Isles. Zonitide. Endodontide. Helicide. Part XVII. Taylor Brothers, Leeds, United Kingdom. Received 30 October 2018 Accepted 30 July 2019 The Canadian Field-Naturalist First record of Paintedhand Mudbug (Lacunicambarus polychromatus) in Ontario and Canada and the significance of iNaturalist in making new discoveries CoLin D. JONES’*, MAEL G. GLON’, KAREN CEDAR®, STEVEN M. PAreRO‘, PAUL D. PRATT’, and THOMAS J. PRENEY® 'Natural Heritage Information Centre, Ontario Ministry of Natural Resources and Forestry, Science and Research Branch, 300 Water Street, Peterborough, Ontario K9J 3C7 Canada "Department of Evolution, Ecology, and Organismal Biology, Ohio State University, 318 West 12th Avenue, Columbus, Ohio 43210 USA 3Ojibway Nature Centre, 5200 Matchette Road, Windsor, Ontario N9C 4E8 Canada ‘University of Guelph, School of Environmental Sciences, 50 Stone Road East, Guelph, Ontario NIG 2W1 Canada “Corresponding author: colin.jones@ontario.ca Jones, C.D, M.G. Glon, K. Cedar, S.M. Paiero, P.D. Pratt, and T.J. Preney. 2019. First record of the Paintedhand Mudbug (Lacunicambarus polychromatus) in Ontario and Canada and the significance of iNaturalist in making new discov- eries. Canadian Field-Naturalist 133(2): 160-166. https://doi.org/10.22621/cfn.v133i2.2223 Abstract Paintedhand Mudbug (Lacunicambarus polychromatus (Thoma, Jezerinac & Simon 2005)) (Decapoda: Cambaridae) was recently discovered at three locations in Windsor, Ontario. These represent the first reports of this burrowing crayfish in Canada. iNaturalist, a nature app and website designed to record photo-based observations of plants and animals, was instrumental in facilitating this discovery. We discuss the importance of collaborative platforms, such as iNaturalist, for linking naturalists and citizen scientists to taxonomic experts around the globe. Key words: Cambaridae; crayfish; Decapoda; iNaturalist; Lacunicambarus polychromatus, new distribution record; Ontario; Paintedhand Mudbug Introduction Paintedhand Mudbug (Lacunicambarus polychro- matus) 1s a burrowing crayfish in the family Cam- baridae that was newly described in 2005 (Thoma et al. 2005). Until recently, it was included in the genus Cambarus Erichson, 1846, in the rejected subgenus Tubericambarus Jezerinac, 1993 (Crandall and De Grave 2017). Phylogenetic analyses of mitochondrial DNA (mtDNA) sequence data combined with morph- ological and ecological characteristics have, however, indicated that L. polychromatus and several closely re- lated species of burrowing crayfish are monophyletic and distinct from Cambarus (Glon et al. 2018). As a result, the subgenus Lacunicambarus Hobbs, 1969 has been resurrected and redescribed at the generic level to accommodate them (Glon ef al. 2018). Lacunicambarus polychromatus is known from throughout much of the North American midwest east of the Mississippi River (with the exception of a recent record from Missouri; Missouri Statewide Historical Crayfish Database 2019), as well as parts of Kentucky and Tennessee (see Figure 2 in Thoma et al. 2005). Throughout its range, L. polychromatus is commonly found in burrows in low-lying habi- tats close to the water table, including the banks and floodplains of lakes and rivers, roadside ditches, and wetlands (Thoma et a/. 2005). The recent recognition of L. polychromatus, stemming in part from histor- ical confusion with the closely related Devil Crayfish (Lacunicambarus diogenes), has led to a relative pau- city of correctly identified records of this species in museum collections and databases. The full extent of the range of L. polychromatus, therefore, remains un- determined, and no records of this species have been reported previously from Canada. Methods On 26 May 2017, members of the Committee on the Status of Species at Risk in Ontario (COSSARO) visited the Ojibway Prairie Provincial Nature Re- serve (42.263°N, 83.071°W) in Windsor, Ontario, along with several staff from the Ontario Ministry of Natural Resources and Forestry and the Ojibway Nature Centre, City of Windsor. During the visit, A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 160 ©The Ottawa Field-Naturalists’ Club 2019 a large adult burrowing crayfish was encountered above ground during the daytime. The individual was photographed and the photos (Figure 1) were uploaded to iNaturalist (see https://inaturalist.ca/ observations/6501788), a web-based application de- signed to capture and share photo-based records of plants and animals. At the time of the observation, the crayfish was identified as L. aff. diogenes (see Glon et al. 2018), a species known to occur in Ontario, Canada (Crocker and Barr 1968; Guiasu et al. 1996; Hamr 1998). Six months later, the iNaturalist record was discovered by M.G.G., an expert on the taxon- omy of Lacunicambarus currently working on revis- ing the genus, who suspected that it was more likely to be L. polychromatus. Additional photos were up- loaded to the record allowing for confirmation that the species was indeed L. polychromatus, a species not previously reported for Canada. On 15 May 2018, C.D.J. and M.G.G. returned to the original location to conduct additional surveys for the species and to collect a voucher specimen and tissue samples for DNA analysis. Burrows were JONES ET AL.: PAINTEDHAND MUDBUG IN CANADA 161 searched for in ditches and along watercourses, and a hand-pumping technique, which included pouring additional water into the burrows, was used to bring the crayfish to the surface. Surveys for burrowing crayfish were also conducted on the same day at four additional locations in the Windsor area (Figure 2, Table 1). When burrows were located, the tech- nique described above was used to extract crayfish. Voucher specimens were collected at each location where crayfish were found and have been deposited at the Royal Ontario Museum, Toronto, Ontario. A tis- sue sample was also taken from the Ojibway Prairie specimen for mtDNA sequencing. Results The surveys on 15 May 2018 at Ojibway Prairie Provincial Nature Reserve were successful in con- firming the presence of L. polychromatus. At this location, burrows were found along a shallow ditch running along the northern edge of the nature reserve not far from where the specimen was located in 2017. The nature reserve is 105 ha in size and includes a Ficure 1. Paintedhand Mudbug (Lacunicambarus polychromatus) recorded on 26 May 2017 at Ojibway Prairie Provincial Nature Reserve, Windsor, Ontario. Distinguishing characteristics of the species are indicated by lines: closed areola (a, c); deflected rostrum (bl); suborbital angle (b2); dorsal surface of the palm of the chela with mesial quarter to third studded with tubercles not forming distinct rows (d). Photos: Colin D. Jones. Todd Lane F - < o 6:3. fodiens she Lacunicambarus polychromatus %.. crayfish found FIGURE 2. The six survey sites in the Windsor, Ontario, area. Stars indicate the three locations where Paintedhand Mudbug (Lacunicambarus polychromatus) was detected. Circles indicate the two locations where Digger Crayfish (Creaserinus fodiens) was found. The x indicates a loca- tion where no crayfish burrows or crayfish were found. The hatched area represents the Ojibway Prairie Provincial Na- ture Reserve. combination of native tallgrass prairie and prairie/sa- vanna habitat. It is part of a complex of closely situ- ated natural areas located in the City of Windsor that together protect nearly 350 ha of native prairie, sa- vanna, and forest. A single specimen of L. polychromatus was col- lected, preserved in 95% ethanol and deposited at the Royal Ontario Museum (ROMIZ L5261). The GenBank accession numbers for 12S (MH878691), 16S (MH878723), and partial COl (MH882991) have been previously reported in Glon ef al. (2019). On the same day, L. polychromatus was also collected from a nearby site, Vince Marcotte Park (42.245°N, 83.071°W). Unlike the natural habitat of the nature reserve, the burrows at Vince Marcotte Park were located on the manicured lawn of the park bordering Turkey Creek. There were many bur- rows at this location and two specimens of L. poly- chromatus were collected and deposited at the Royal Ontario Museum (Table 1, Figure 2). On 15 May 2018, three additional sites were sur- veyed (Table 1). At one site, no crayfish burrows were THE CANADIAN FIELD-NATURALIST Vol. 133 located. At the other two sites, crayfish burrows were found and hand-pumped. The species present, how- ever, was Digger Crayfish (Creaserinus fodiens), which was first discovered in Ontario, Canada, in 1863 and is often found living in burrows adjacent to those of L. polychromatus in the United States (Cottle 1863; Hobbs 1974). Following the discovery that L. polychromatus occurs in the Windsor area (Table 1), an additional photo-based record was uploaded to iNaturalist from a third location adjacent to Ojibway Prairie. An in- dividual was found by Steve Marks above ground, at night, and backdated to 24 May 2016 (see https:// inaturalist.ca/observations/17855665). The location is a residential yard bordered by a shallow drainage ditch adjacent to a dry-moist old field. Crayfish bur- rows are periodically found in the manicured lawn and along the banks of the ditch at this location, but no attempts have been made to survey for crayfish. Discussion The Ojibway Prairie complex in extreme south- western Ontario supports a number of plant and ani- mal species that are restricted to this unique portion of the province, not to mention the entire country (Oldham 1992; Paiero et al. 2010; Pratt 2018). The discovery of L. polychromatus here represents the first record of the species in Canada. The closest known occurrences are in adjacent Michigan, where it has been known since it was first described (Thoma et al. 2005); thus, it is not entirely surprising that it was eventually recorded across the Detroit River in the Windsor area. Lacunicambarus polychromatus is almost certainly native to Canada, rather than be- ing a recent colonizer or introduction, and its pres- ence has likely gone undetected because of historical confusion about the taxonomy of Lacunicambarus. Additional surveys will be required throughout southwestern Ontario to delineate the full extent of the species’ range in Canada. We also suggest that preserved specimens ascribed to “C. diogenes” (i.e., L. aff. diogenes) be re-examined, particularly if they were collected before L. polychromatus was de- scribed in 2005, to determine if the latter species is more widely distributed in Ontario than the observa- tions documented herein led us to believe. Such in- formation will also be critical in making an accurate assessment of the species’ conservation status prov- incially and nationally. Lacunicambarus polychromatus can be differen- tiated from all other Canadian crayfishes by its closed areola (Figure la,c); strongly deflected rostrum (in lateral view; Figure 1bl); presence of a suborbital angle (Figure 1b2); Form I male gonopods with two terminal elements directed caudally at approxi- 163 po1d9][09 sualpof snuldasvadD JONES ET AL.: PAINTEDHAND MUDBUG IN CANADA 2019 OZEO6STI ISTTeINIeNI ouo ‘poduind pue payoojap =yied yeuoryeos901 SULIOp sInq\sIoywy v9cs T ZINOY Wow Tao DOW SMOLIN “YOIeIS poyosIe], “10g Yop opispeoy = 6L0€8 981 Cr “ysed preuey JAY 8107 ABW ST poyoayjoo (ysyAeig IO83Iq) sualpof snuldasDadD SETOSSTI STTeINIENI ouo ‘poduind pue payoojop JOSPUIM “BoIV S9CS'T ZINOU Wow Tao DOW SMOLIN “YOIBIS po}osIeL, durems popoom [$0 €8 9S CL [einyeN uapsey suLidg gio AR SI vole [eLysnpul poyoajop ysyAeso UI PpoYy [[euls SuLIOp eu Bu LTQCO°DOW JO SMOlINg ou ‘YyoIeas poyosIeL -10q YOUIp opispeoy 6S0'€8 ELT Th JOSPUIA “OAV [eWOUTIUOD gIOT AP SI LOEO6STI 3STTeINIeNI po1oa]]09 snyowosyodjod “T €9ZST ZINOY OM} ‘poduind pure poyoajop yoord SuLIap JOSPUI AA, c9CS'T ZNO Wow fao°DOW SMOLIN ‘Yves poyosse], -I0g ume] yed ueqiA ILO €8 SVC CY “YlWd OOIIRIY PUTA 8IOT APIA SI poyoo][oo snyoutosyodjod °T 68006STI 3SITernIeNI ouo ‘poduind pue payoojap ~— ates ssvisy]je} sATeU JOSPUIAA “OATOSOY OINIEN 19ST ZINOY Wow Tao DOW SMOLIN ‘YoIvas poyosse], SULJOpsIog YoUp MoT[eYs IL0'€8 €9T Tp [IOUT AOI o1eIg ARQ gIOT AR ST SIOqUIOU OUVSSOD + “TL “add “TNS°OM = poydessoi0yd snyouosyodjod °T JOSPUIM “SAIOSOY DINIEN 88Z10S9 IseINWNI “OOW fd) uo ‘yeJaplouy] ated ssess]][e} OATEN IL0'€8 €9T Tp [IOUT AOI otseIg AeMqHO L107 AP 97 poydeisoj0yd snjpwosyatjod *T Yyoup oseurvip SO9SS8LI ISITeINIeNT WS duo “[eJUaplou] SUlJapsog pred ueginqns 6S0 €8 C9T CV JOSPUIM “IS UOIQUIeT 9107 AB HZ ‘ou Boyeyeg «=—- AOUSodayy SIOAIOSGO sjiejoq yeyiqeH Saacior ae uol}v90'T ayeq ‘/SUOTILAIOSQO/eO 'IsTTeINJeUT//:sdyy “TY ey1 JO pus oy} 0} Jaquinu Zoyeyeo oy} Surppe Aq ouTpUO a[qissaooe aie pue Joquinu p10daI onbrun dy} 0} JajaI ISIPeINJENT JOJ sJoquinu soyeyed ‘(WOY) Woensnyy o1seyUG [eAoY oy} puv (Spode paseq-ojoYd JoJ) ISTPeINFNT YIOQ apnyout sarroysoday ‘A][NJssaoonsun JO} poyoress sem saroods ay] a1oYM SUOTIedO] SuIpNpoUr ‘epeuRD ‘oLIeIUG WoL] (snsoWOsYyIKjod snADqupsiUuNIVT) Sngpny pueypeiuleg Jo sp1ocdy *[ ATAV, 164 mately 90° relative to the gonopod shaft, the distal most of which (central projection) is markedly shorter than the proximal one (mesial process); and dorsal surface of the palm of the chela with mesial quar- ter to third studded with tubercles not forming dis- tinct rows (Figure 1d). Although colouration is sub- ject to variation and is not always a reliable character, L. polychromatus typically exhibits striking orange and red highlights along its rostral margins, chelae, and the dorsoposterior margins of its abdominal som- ites and blue or green shades across its body (Figure la—d). This crayfish is almost always collected from burrows, but it is occasionally encountered in open waters or even on land, particularly after heavy rain. Burrows with recent activity have fresh mud at the opening (Figure 3) often forming a chimney (Figure 4). The burrows themselves are not, however, diag- nostic to species, and there are several species of bur- rowing crayfish. iNaturalist was instrumental in facilitating the discovery of this species in Canada. Had it not been for iNaturalist, tts presence may have remained un- detected. As iNaturalist grows in popularity, more and more amateur naturalists and citizen scientists are uploading their photos and seeking confirmation of their identification. For example, in the 10 years since its inception in 2008 over 7 000 000 observa- tions have been contributed by 224 334 users (iNatur- alist 2019). In 2018 alone, the total number of obser- vations more than doubled to well over 15 000 000 and the number of users also more than doubled to an astonishing 501 308 (iNaturalist 2019; also see Martin 2018 for a summary of Canadian records). At the same time, more and more taxonomic experts are becoming involved in iNaturalist by offering their ex- pertise at providing or correcting identifications. This was certainly the case with the Ojibway Prairie rec- ord of L. polychromatus that would have been entered in field notes and potentially in databases as a record of L. aff. diogenes. In time, the photos may have been re-examined and re-determined as L. polychromatus, but iNaturalist has provided a platform that has greatly increased the ability of amateurs and experts to col- laborate in real time. This collaboration is greatly in- creasing our collective knowledge of the distribution and, in fact, the conservation status of species. There are other examples of this rapid increase in species discoveries. In Ontario alone, for example, at least 40 species of moths new to the province (at least 19 of which are new to Canada and one new to North America) have been discovered (M.V.B. Burrell pers. comm. 22 January 2019) through iNaturalist submis- sions since the recent publication of a comprehensive annotated Canadian Lepidoptera list (Pohl et a/. 2018). As species, such as Paintedhand Mudbug, are new- THE CANADIAN FIELD-NATURALIST Vol. 133 FiGurE 3. Fresh mud at the opening of a crayfish burrow in Vince Marcotte Park, Windsor, Ontario, Canada (42.245°N, 83.071°W), 15 May 2018. Photo: Colin D. Jones. a Figure 4. A Paintedhand Mudbug (Lacunicambarus poly- chromatus) chimney in Muhlenberg County, Kentucky, USA (37.313°N, 87.201°W), 11 June 2018. Photo: Mael G. Glon. ly discovered in Ontario, they are added to the prov- incial species list maintained by the Natural Heritage Information Centre (NHIC), Ontario Ministry of Na- tural Resources and Forestry, and are assigned a con- servation status rank following methods developed by NatureServe (Faber-Langendoen et al. 2012; Master et al. 2012). Paintedhand Mudbug has been assigned a rank of S1S2 (critically imperilled to imperilled). The NHIC typically compiles observations of species with a conservation status rank of S3 (vulnerable) or higher. These observations then inform areas of con- servation value (element occurrences) and are added to the provincial record where they are available to inform conservation, land-use, and natural resources management planning, policy, and legislation. iNaturalist is also greatly increasing the ability to detect the introduction of exotic and potentially inva- sive species. A new and potentially invasive vascu- lar plant species, Small-flowered Jewelweed (/mpa- tiens parviflora de Candolle), for example, has been 2019 recently documented in Ontario through iNaturalist (Oldham 2018). The growth and popularity of iNaturalist is not showing any signs of slowing down. With such growth, an increasing number of local, regional, and nation- al discoveries will be made, such as the discovery of L. polychromatus in Canada. Such discoveries will assist in advancing our collective knowledge of the distribution and conservation status of species. The NHIC has created a project on iNaturalist that users can join allowing their personal observations of prov- incially rare species to be considered for incorpora- tion into the provincial record (https://inaturalist.ca/ projects/nhic-rare-species-of-ontario). Author Contributions Writing — Original Draft: C.D.J. and M.G.G.; Wri- ting — Review and Editing: all; Conceptualization — C.D.J. and M.G.G.; Methods — M.G.G. and C.D.J.; In- vestigation — all. Acknowledgements We thank the Ontario Ministry of Natural Re- sources and Forestry (OMNRP) for providing a per- mit to conduct research in Ontario’s protected places and Jim Wigle, acting park superintendent for Ojib- way Prairie Provincial Nature Reserve for facilitating the surveys there. The City of Windsor provided per- mission to conduct surveys in city parks and protected places. Tim Haan of the Natural Heritage Information Centre, OMNRF, produced the map. Don Stacey and Sebastian Kvist accepted the specimens and have added them to the invertebrate collections at the Royal Ontario Museum, Toronto. The surveys during which these observations were made were supported by OMNRE and The Ohio State University. Mike Burrell of the Natural Heritage Information Centre provided a summary of moths new to Ontario via iNaturalist sub- missions. We also thank Don Sutherland, OMNRF, and Zac Loughman, West Liberty University, for their constructive comments on an earlier draft. The sur- veys and the collection of voucher specimens within provincially protected areas were conducted under a Letter of Authorization to Conduct Research in a Provincial Park or Conservation Reserve issued to the Natural Heritage Information Centre, Ontario Ministry of Natural Resources and Forestry. Literature Cited Cottle, T.J. 1863. On the two species of Astacus found in Up- per Canada. Canadian Journal of Industry, Science, and Art 45: 216-219. Crandall, K.A., and S. De Grave. 2017. An updated clas- sification of the freshwater crayfishes (Decapoda: Asta- cidea) of the world, with a complete species list. Journal of Crustacean Biology 37: 615—653. http://doi.org/dcw4 JONES ET AL.: PAINTEDHAND MUDBUG IN CANADA 165 Crocker, D.W., and D.W. Barr. 1968. Handbook of the Crayfishes of Ontario. Royal Ontario Museum life sci- ences miscellaneous publications, University of Toronto Press, Toronto, Ontario, Canada. https://doi.org/10.5962/ bhl.title.60758 Faber-Langendoen, D., J. Nichols, L. Master, K. Snow, A. Tomaino, R. Bittman, G. Hammerson, B. Heidel, L. Ramsay, A. Teucher, and B. Young. 2012. NatureServe conservation status assessments: methodology for as- signing ranks. NatureServe, Arlington, Virginia, USA. Glon, M.G., R.F. Thoma, M. Daly, and J.V. Freudenstein. 2019. Lacunicambarus chimera: a new species of bur- rowing crayfish (Decapoda: Cambaridae) from Illinois, Indiana, Kentucky, and Tennessee. Zootaxa 4544: 451— 478. https://doi.org/10.11646/zootaxa.4544.4.1 Glon, M.G., R.F. Thoma, C.A. Taylor, M. Daly, and J.V. Freudenstein. 2018. Molecular phylogenetic an- alysis of the devil crayfish group, with elevation of Lacunicambarus Hobbs, 1969 to generic rank and a re- description of the devil crayfish, Lacunicambarus dio- genes (Girard, 1852) comb. nov. (Decapoda: Astacoidea: Cambaridae). Journal of Crustacean Biology 38: 600— 613. https://doi.org/10.1093/jcbiol/ruy057 Guiasu, R.C., D.W. Barr, and D.W. Dunham. 1996. Distri- bution and status of crayfishes of the genera Cambarus and Fallicambarus (Decapoda: Cambaridae) in Ontario, Canada. Journal of Crustacean Biology 16: 373-383. https://doi.org/10.1163/193724096x00162 Hamr, P. 1998. Conservation status of Canadian freshwater crayfishes. World Wildlife Fund Canada and Canadian Nature Federation, Toronto, Ontario, Canada. Hobbs, H.H., Jr. 1974. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cam- baridae). Smithsonian Contributions to Zoology 166: 1— 161. https://dot.org/10.5479/si.00810282. 166 iNaturalist. 2019. iNaturalist.org year in review 2018. iNaturalist. Accessed 17 January 2019. https://www. inaturalist.org/stats/2018. Martin, A.E. 2018. iNaturalist Canada passes the 1 000 000 observation mark. Canadian Field-Naturalist 132: 434. https://doi.org/10.22621/cfn.v13214.2305 Master, L., D. Faber-Langendoen, R. Bittman. G.A. Hammerson, B. Heidel, L. Ramsay, K. Snow, A. Teu- cher, and A. Tomaino. 2012. NatureServe conservation status assessments: factors for evaluating species and ecosystem risk. NatureServe, Arlington, Virginia, USA. Missouri Department of Conservation. 2019. Missouri statewide historical crayfish database. Missouri Depart- ment of Conservation, Resource Science Division, Co- lumbia, Missouri, USA. Oldham, M.J. 1992. Rare vascular plants of Ojibway Prairie and vicinity, Windsor, Ontario. Unpublished re- port. Ontario Ministry of Natural Resources, Aylmer, Ontario, Canada. Oldham, M.J. 2018. Two new and potentially invasive vas- cular plant species recently documented in Ontario through citizen science and social media. Unpublished report. Ontario Ministry of Natural Resources, Peterbo- rough, Ontario, Canada. Paiero, S.M., S.A. Marshall, P.D. Pratt, and M. Buck. 2010. Insects of Ojibway Prairie, a southern Ontario tall- 166 grass prairie. Pages 199-225 in Arthropods of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats. Edited by J.D. Shorthouse and K.D. Floate. Biological Survey of Canada, Canada. https://doi org/10.3752/9780968932148.ch9 Pohl, G.R., J.-F. Landry, B.C. Schmidt, J.D. Lafontaine, J.T. Troubridge, A.D. Macaulay, E.J. van Nieuker- ken, J.R. DeWaard, J.J. Dombroskie, J. Klymko, V. Nazari, and K. Stead. 2018. Annotated checklist of the moths and butterflies (Lepidoptera) of Canada and Alaska. Pensoft Publishers, Sofia, Bulgaria. Pratt, P.D. 2018. Provincially rare vascular plants and wild- THE CANADIAN FIELD-NATURALIST Vol. 133 life of the Ojibway Prairie complex (version APR2018). Ojibway Nature Centre, Department of Parks, Windsor, Ontario. Accessed 22 January 2019. http://www.ojibway .ca/raresp.htm. Thoma, R.F., R.F. Jezerinac, and T.P. Simon. 2005. Cam- barus (Tubericambarus) polychromatus (Decapoda: Cambaridae), a new species of crayfish from the United States. Proceedings of the Biological Society of Wash- ington 118: 326-336. http://doi.org/dg6mcq Received 22 February 2019 Accepted 2 August 2019 The Canadian Field-Naturalist Note Duckling mortality at a river weir STEWART B. Roop"* and AMBER WILLCOCKS! ‘Department of Biological Sciences, University of Lethbridge, Lethbridge, Alberta T1K 3M4 Canada “Corresponding author: rood@uleth.ca Rood, S.B., and A. Willcocks. 2019. Duckling mortality at a river weir. Canadian Field-Naturalist 133(2): 167-171. https:// doi.org/10.22621/cfn.v13312.2159 Abstract River weirs are low-head dams that dissipate energy by creating hydraulic recirculation zones at their base. These recircu- lation zones are a major cause of human drownings and have been referred to as “drowning machines”. We observed an event that allowed us to add ducklings to the list of weir victims. As a Mallard (Anas platyrhynchos) hen and her brood floated over the Calgary weir, the mother flew safely over the hydraulic recirculation. The ducklings drifted into the recircu- lation and three quickly passed through; four were stalled, repeatedly recirculated, and died. We observed other regional weirs where adult birds commonly flew over the hazard. We did not observe any other waterfowl drifting into recircula- tion zones, and we found no prior report of this lethal hazard. Although mortality might be rare at each weir, with hundreds of thousands of low-head dams worldwide, the collective hazard could be substantial. Weirs can be designed to eliminate the lethal recirculation zone, and the apparent hazard to ducklings could provide another motivation to redesign or modify these common structures. Key words: Bow River; dams; mortality; waterfowl Human developments impose many lethal chal- lenges on wildlife. Avian mortality can follow colli- sions with windows, towers, power lines, airplanes, or wind turbines (Weir 1976; Avery et al. 1980). Al- terations to streams, ponds, and lakes impose addi- tional challenges for waterfowl, including the flood- ing of nests and increased predator access (Mauser et al. 1994; Flint and Grant 1997). We are investigating the ecological consequences of weirs and other river dams (Rood et al. 2003, 2010) and observed another, apparently unreported, cause of waterfowl mortality. Our observation was from the viewing platform at the weir across the Bow River in Calgary, Canada (Figure 1; 51°2.61'N, 114°0.85'W). This weir creates an elevated head pond upstream, which allows for off- stream water diversion for irrigation. The concrete weir produces a drop of about 2 m into the stilling pool, forming a hydraulic recirculation zone, which is also referred to as a “hydraulic jump” or “hole” (Figure 2; Makuk 1988; Bradshaw 2004). This re- circulation zone dissipates energy to reduce erosion of the weir base and downstream channel. Because of the recirculation, buoyant objects that float over the sill are often stalled in the trough, and are retained there or plunge down into the seam created by falling water. These objects flow slightly downstream, then resurface because of their buoyancy while still within the turbulent recirculation zone. This imposes a re- petitive and somewhat violent cycle, and the recircu- lation zone can thus be retentive, as revealed with sticks, logs, and a wheel in the trough of the Calgary weir (Figures | and 3). With the hydraulic recirculation, low-head dams will often stop swimmers or stall and capsize small boats, and then recirculate the victims. The lethal hazard of these “drowning machines” is well recog- nized, as human drownings at river weirs continue to represent about a fifth of recent North American river fatalities (Walbridge 2000; Bradshaw 2004). The Calgary weir has been especially lethal, with at least 10 fatalities from 1982 through 2007, includ- ing four double drownings (Makuk 1988; Canadian River Safety 2019). Largely because of the drowning hazard, this weir was modified in about 2010 to div- ide the single major drop into a sequence of smaller irregular drops, eliminating the retentive hydraulic recirculation zone and creating the safer Harvie Passage paddling park. While at that construction site on 4 June 2009, we observed a fatal wildlife incident. Near mid-day, a 167 ©The Ottawa Field-Naturalists’ Club Vol. 133 168 THE CANADIAN FIELD-NATURALIST ee ee Figure 1. The Calgary weir in June 2009 (discharge 109 m*/s), displaying the river water flowing over the weir and into the hazardous hydraulic recirculation zone, with trapped sticks and a wheel. The structure on the far side allows offstream diversion into an irrigation canal. This system was subsequently modified with the Harvie Passage to produce safer pad- dling channels. Photo: Stewart Rood. Water surface 2m Recirculation * Buoyancy Escape Aerated surface Stilling Pool 5m FiGuRE 2. A cross-section of the Calgary weir and hydraulic recirculation zone with scaling based on Makuk (1988) and Golder (2002). The positions of the submerged ducklings are based on flume simulations with objects of similar buoyancy and observed emergence patterns. Note the exaggerated vertical scale. Mallard (Anas platyrhynchos) hen and her brood of seven ducklings drifted slowly along the south bank (Figure 3). As water flowed over the weir, the duck- lings turned upstream and appeared to paddle vigor- ously. As the hen floated over the sill, she rose with a few wing strokes and glided downstream beyond the recirculation zone. Some of the ducklings turned to- ward her and paddled vigorously downstream. Paddling into the trough, three passed through fairly quickly, apparently aided by a break in the re- circulation because of a tumbling log in the trough. The other four ducklings were stalled. Each was plunged down, and we observed three of the four re- surface a few seconds later in the aerated surface, 3—4 m downstream. Despite paddling downstream, each was drawn back upstream into the trough and ROOD AND WILLCOCKS: RIVER WEIRS DROWN DUCKLINGS 169 Ficure 3. A Mallard (Anas platyrhynchos) hen with seven ducklings approaching the hydraulic recirculation zone at the Calgary weir, 4 June 2009. Three of these ducklings survived, while four died. The hen is just rising and subsequently flew over the recirculation area. Photo: Stewart Rood. again plunged under water. With each recirculation, the ducklings appeared to be weakened as they be- came less vigorous. We observed two ducklings re- circulating three times and one four times, but did not observe the fourth over the next few minutes. After about five more minutes, one duckling emerging in the water downstream from the hydraulic recircula- tion area, drifting passively on its side, apparently dead. We observed no further evidence of the remain- ing three ducklings over an additional 30 min. This incident demonstrates that a low-head dam can be lethal for ducklings. The mature hen was read- ily able to fly over the retentive hydraulic recircula- tion zone, but the ducklings were unable to fly and were, therefore, vulnerable. Of the seven ducklings, four apparently died, indicating that this powerful re- circulation was quite hazardous. This physical hazard from low-head dams may be very common; the design of these dams is fairly universal and consistently produces retentive and lethal hydraulic recirculation zones (Makuk 1988; Walbridge 2000; Bradshaw 2004). There are about 45 000 large dams world-wide (World Commission on Dams 2000) and low-head dams are probably about ten-fold more numerous (Chandler and Chapman 2003; Doyle et a/. 2003). With this widespread occur- rence and the apparent lethality for ducklings, we con- clude that weirs could provide a substantial hazard for juvenile waterfowl. To investigate whether this hazard is recognized in the field of ecology, we conducted Google Scholar and Web of Science literature searches with search term combinations including duck, duckling, mortality, death, hazard, dam, and weir. These revealed exten- sive reports on duckling mortality, including reviews and bibliographies, showing that duckling mortal- ity is considerable (Ringleman and Longcore 1982; Savard et al. 1991; Flint and Grand 1997), largely as a result of predation (Talent et a/. 1983; Mauser et al. 1994). Although many reports of hazards to water- fowl and other birds from various types of artificial structures exist (Weir 1976; Avery et al. 1980), we found no reference to the drowning hazard from river weirs. Wildlife biologists with Alberta Environment and Parks reported that they had never seen ducklings or goslings drifting over weirs, but were not surprised by the possibility, because waterfowl are abundant in head ponds above weirs in southern Alberta. News and social media websites were another source of information. A Google search (September 2019) on “ducklings and weirs” revealed YouTube, Facebook, and news media videos, photographs, and 170 reports of ducklings unable to follow the mother hen up and over weirs or stranded in troughs or drains below weirs. These ducklings were uninjured and were often assisted by observers or wildlife officers; for some repetitive cases, screens or other measures solved the stranding. We found no report of ducklings retained, injured, or killed in the hydraulic recircula- tion zone of a river weir. Over five years, we visited other river weirs, in- cluding the 1-m-high Lethbridge Northern Irrigation District weir (49°39.9'N, 113°36.1'W), the 1.5-m City of Lethbridge weir on the Oldman River (49°40.9'N, 112°51.4'W), and the 3-m Carseland weir on the Bow River (50°49.5'N, 113°26.6'W). These weirs also pro- duce retentive hydraulic recirculation zones and hu- mans have drowned at the Lethbridge and Carseland weirs (Canadian River Safety 2019). We undertook 50, 6-hour observer-days at these three weirs during June from 2010 through 2015. During these observations, water birds were com- mon near each of the weirs, including the piscivorous American White Pelican (Pelecanus erythrorhyn- chos) and Double-crested Cormorant (Phalarocorax auritus). In addition to Mallards, other ducks were abundant, including Common Merganser (Mergus merganser), Common Goldeneye (Bucephala clang- ula), occasional Wood Ducks (Aix sponsa), and other THE CANADIAN FIELD-NATURALIST Vol. 133 species. Canada Goose (Branta canadensis), includ- ing goslings, were also abundant, along with Ring- billed (Larus delawarensis) and other gulls. We com- monly observed adults drifting toward the weirs, and they would consistently fly before drifting into the hydraulic recirculation zone. Ducklings or gos- lings were common, but we did not see any others drift over the weirs; thus, it is probably uncommon for waterfowl to drift into recirculation zones. We have repeatedly observed pelicans and some other birds fishing right below and even in the recirculation zone during summer intervals when river flows are lower and the hydraulic power and hazard are prob- ably slight (Figure 4). Following these observations, we suspect that the hazard to waterfowl varies considerably across weirs and with different river flows. A small weir on a small stream would produce a modest hydraulic recircula- tion zone that would be less capable of stopping and recirculating ducklings. The hydraulic force and haz- ard would increase with increasing drop height and stream discharge. The Carseland weir hydraulic sys- tem was very loud and this may have signaled its presence from 100 m or more upstream. Waterfowl were abundant in the upstream pond, but remained well away from that drop. With the reduced haz- ard of smaller weirs and noise cues of larger weirs, Figure 4. The Lethbridge weir across the Oldman River in late summer when flows are low and the recirculation hazard is reduced. American White Pelicans (Pe/ecanus erythrorhynchos) are common, feeding on fish that are blocked by the weir from upstream passage. Photo: Stewart Rood. 2019 it is possible that intermediate-sized weirs, such as the Calgary weir, might provide the greatest risk to ducklings. The lack of prior reporting of duckling deaths in weir recirculation zones could indicate that this source of mortality is rare. Conversely, the lack of evidence may reflect ducklings’ inconspicuous na- ture. Ducklings are small and cryptic, and hens en- courage secretive behaviour (Mauser ef al. 1994). Also, reducing the likelihood of observation, hu- mans are discouraged from approaching river weirs by signage or fencing. Although stilling pools, the slow flowing zones downstream of weirs, are some- times favoured for fishing, this is less common during the interval of turbid water and high flow rates in late spring and early summer, when ducklings are unable to fly and thus more vulnerable. Although the extent of mortality is unknown, the prospective drowning hazard to waterfowl could pro- vide one more reason to avoid weir designs that cre- ate “drowning machines”. In addition, modifications to existing weirs to eliminate the hydraulic recircu- lation zone should reduce the risk for ducklings and possibly other wildlife, as well as humans. For ex- ample, the modifications to the Calgary weir to create the safer Harvie Passage might reduce future duck- ling mortality at this location where we observed the tragic drownings in 2009. Acknowledgements This work was supported by funding from Alberta Innovates and Alberta Environment and Parks (AEP). We extend thanks to ornithologist Andrew Hurly (University of Lethbridge) and ecohydrologist Andrew Paul (AEP) for valuable inputs and editors Dwayne Lepitzki and William Halliday and two re- viewers for many helpful recommendations. Literature Cited Avery, M.L., P.F. Springer, and N.S. Dailey. 1980. Avian mortality at man-made structures: an annotated bibli- ography (revised). United States Fish and Wildlife Ser- vice, Department of the Interior, Washington, DC, USA. FWS/OBS-80/54. Bradshaw, S. 2004. River Safety: A Floater’s Guide. Lyons Press, Guilford, Connecticut, USA. Canadian River Safety. 2019. Map of incidents in Alberta. Canadian River Safety Project, Canada. Accessed Sep- tember 2019. http://www.canadianriversafety.ca/w/core/ index.php?title=Province: Alberta. Chandler, J.A., and D. Chapman. 2003. Existing habitat ROOD AND WILLCOCKS: RIVER WEIRS DROWN DUCKLINGS 171 conditions of tributaries formerly used by anadromous fish (appendix 2.1-2). Technical report FERC no. 1971. Idaho Power Company, Boise, Idaho, USA. Doyle, M.W., E.H. Stanley, and J.M. Harbor. 2003. Channel adjustments following two dam removals in Wisconsin. Water Resources Research 39: 1011. https:// doi.org/10.1029/2002W ROO1714 Flint, P.L., and J.B. Grand. 1997. Survival of spectacled eider adult females and ducklings during brood rearing. Journal of Wildlife Management 61: 217—221. https:// doi.org/10.2307/3802430 Golder Associates. 2002. Pre-design of the preferred alter- natives to modify the Western headworks weir in Cal- gary. Golder Associates Ltd., Calgary, Alberta, Canada. Makuk, J.S. 1988. Multiple use of water resources: adapt- ing weirs for recreation. M.Env. Design thesis, Univer- sity of Calgary, Calgary, Alberta, Canada. Mauser, D.M., R.L. Jarvis, and D.S. Gilmer. 1994. Sur- vival of radio-marked mallard ducklings in northeast- ern California. Journal of Wildlife Management 58: 82— 87. https://doi.org/10.2307/3809552 Ringleman, J.K., and J.R. Longcore. 1982. Survival of black ducks during brood rearing. Journal of Wild- life Management 46: 622-628. https://doi.org/10.2307/ 3808552 Rood, S.B., J.H. Braatne, and L.A. Goater. 2010. Favor- able fragmentation: river reservoirs can impede down- stream expansion of riparian weeds. Ecological Appli- cations 20: 1664-1677. https://doi.org/10.1890/09-0063.1 Rood, S.B., C.R. Gourley, E.M. Ammon, L.G. Heki, J.R. Klotz, M.L. Morrison, D. Mosely, G.G. Scop- pettone, S. Swanson, and P.L. Wagner. 2003. Flows for floodplain forests: a successful riparian restoration. BioScience 53: 647—656. http://doi.org/c4tvrc Savard, J.-P.L., G.E.J. Smith, and J.N.M. Smith. 1991. Duckling mortality in Barrow’s Goldeneye and Buffle- head broods. Auk 108: 568-577. https://doi.org/10.2307/ 4088097 Talent, L.G., R.L. Jarvis, and G.L. Krapu. 1983. Survival of mallard broods in south-central North Dakota. Con- dor 85: 74-78. https://doi.org/10.2307/1367893 Walbridge, C. 2000. The American Canoe Association’s River Safety Report 1996-1999 (Second Edition). Mena- sha Ridge Press, Birmingham, Alabama, USA. Weir, R.D. 1976. Annotated bibliography of bird kills at man-made obstacles: a review of the state of the art and solutions. Department of Fisheries and the Environment, Canadian Wildlife Service, Ottawa, Ontario, Canada. World Commission on Dams. 2000. Dams and Develop- ment: a New Framework for Decision-making. Earth- scan Publications Ltd., London, United Kingdom. Received 22 November 2018 Accepted 17 October 2019 The Canadian Field-Naturalist Book Reviews Book Review Editor’s Note: The Canadian Field Naturalist is a peer-reviewed scientific journal publishing papers on ecology, behaviour, taxonomy, conservation, and other topics relevant to Canadian natural history. In line with this mandate, we review books with a Canadian connection, including those on any species (na- tive or non-native) that inhabits Canada, as well books covering topics of global relevance, including climate change, biodiversity, species extinction, habitat loss, evolution, and field research experiences. Currency Codes: CAD Canadian Dollars, USD United States Dollars, EUR Euros, AUD Australian Dollars, GBP British Pound. BOTANY Sedges of the Northern Forest: A Photographic Guide By Jerry Jenkins. 2019. Cornell University Press. 96 pages, 16.95 USD, Paper. Another regional guide to identification of sedges (Cy- peraceae) has been pub- lished, this time covering the species found in what has been referred to as the “Northern Forest”. This book—and it’s companion “Quick Guide”, a boxed set of two fold-out charts— joins several other excellent treatments of sedges from JERRY JENKINS various parts of North ANORTHERN FOREST ATLAS GUIDE America that have been published in recent years. The area covered by this book does not include much of the forest re- gion Canadians generally would think of as “north- ern’, which would be the boreal forest; rather, it cov- ers mainly what we think of as the Great Lakes-—St. Lawrence Forest Region in Ontario and Quebec, and the Acadian Forest Region in the Maritime Provinces (Rowe 1972). As the title indicates, this book is a photographic guide. It is laid out in such a way that comparisons among similar species can be made easily. The use of a black background behind the images assists greatly in highlighting the important features, such as shape of the perigynium, venation on the perigynium sur- face, perigynium beak shape, size, and orientation, overall inflorescence and individual spike morphol- ogy, scale shape, colour, and size, and various other features that are critical for identification. The great majority of the species found in the region are in- cluded with photographs, but there are a few spe- SEDGES OF THE NORTHERN FOREST GUIDE A PHOTOGRAPH cies for which only brief mention is made in the text that supplements the photographs; most of the spe- cies without photographs are rare, range-edge species within the region covered. The book begins with a brief introduction explain- ing its role and ways to use it. This is followed by an illustrated glossary, the illustrations being sim- plified caricatures or outlines of the features being described. Throughout the book, some simple sym- bols and abbreviations are used to assist in interpre- tation of structures, such as symbols for the sex of the individual flowers or spikelets, and the cross-sec- tional shape of the achenes. A helpful section on veg- etative features, arrangement of spikelets within the larger inflorescence, achene shapes, perianth bristles, etc. is followed by a quick guide to the genera found in the flora, the latter again accompanied by stylized illustrations. Then, the detailed treatment of genera and the sections and species within them (in Carex, for example) follows. All genera, and in the case of Carex, sections within the genus, are treated alpha- betically. Following this, a section called “Quick Guides to Carex” provides more details on morphol- ogy of spikelets, including arrangement of the sexes within them, perigynia, achenes, and, to a lesser ex- tent, leaf and sheath features. Short paragraphs sup- plement the photographs in this section and provide additional hints on identification including, where ap- plicable, habitat notes. More detail on each species is found later in the text, but this section provides a quick reference to help in narrowing down which group within Carex a specimen belongs. Almost half of the book is dedicated to photo- graphs of the most important features that aid in the 2 2019 identification of individual species. Generally, the photographs focus on inflorescence, spikelet, flower, and achene features, but occasionally also include vegetative features. Key features are noted with the photographs (e.g., solitary spikelet, stems smooth, base of bract with lobes), and there are also para- graphs for each genus, section, or group of similar species highlighting distinctive features or providing brief notes on habitat preferences. The quality of the images generally is very good and the notes are help- ful. In most cases, the photographs selected for inclu- sion convey a good representation of the characteris- tic appearance of the species. For difficult groups such as Carex section Ovales, a nice job has been done of grouping similar species together on the plates (e.g., species with perigynia >2 mm wide, species with perigynia covered by the scales). Another difficult group, the genus Eleocharis, has also been dealt with quite well, with good images and notes. One excep- tion to the representation of species with typical pho- tographs is the image of the upper part of the inflo- rescence of Carex molesta, which does not convey its typical appearance, at least to my eye. The spike- lets usually are more closely aggregated into a clus- ter. Another exception is the inflorescence of Carex lenticularis, which normally shows a striking pattern of pale green perigynia among bicoloured black and green scales within the spikelets. I did detect a number of typographical and for- matting errors in the text, such as an incorrect symbol beside the inflorescence of Carex exilis, lack of italics here and there, the odd missing letter in a word, du- plicated entries in the list of older names on p. 87, er- BooK REVIEWS 173 rors with the listing of Carex appalachica and Carex radiata in the list of older names, and so on, but in general the text and the plates are well put together. The author has included his own opinions on topics such as the flurry of nomenclatural changes that has occurred with Carex sectional names over the past few decades, the distinctness (or not) of certain spe- cies (e.g., Carex tincta), and similarities among un- related species, all of which add some colour to the text. One such comment that has me scratching my head, though, is his perception that Carex capillaris is somewhat similar to Carex eburnea. Nevertheless, the great majority of the text contributes well to as- sisting the user to identify sedges. Overall, this book is well conceived, and provides an excellent resource for field botanists who wish to get a better handle on the identification of species in this large and diverse family in the northeastern USA and southeastern Canada. Although the size is a bit cumbersome, | think that it could still be opened on a desk next to a specimen or adjacent to a dissecting scope and used effectively. I certainly recommend it field botanists and naturalists interested in improv- ing their knowledge of this important family of flow- ering plants. Literature Cited Rowe, E.C. 1972. Forest Regions of Canada. Canadian Fo- restry Service Publication No. 1300. Fisheries and En- vironment Canada, Canadian Forest Service, Ottawa, Ontario, Canada. WILLIAM J. CRINS Peterborough, ON, Canada ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 174 ORNITHOLOGY Gulls THE CANADIAN FIELD-NATURALIST Vol. 133 John C. Coulson. 2019. New Naturalist Library, HarperCollins Publishers. 495 pages, 108 charts and graphs, 35 maps, and 109 colour pictures, 135.99 CAD, Cloth. This book is about the gulls of Great Britain and Ireland and to a lesser extent those of Western Europe, but it includes many generalisa- ™ tions that apply to gulls ev- erywhere, gulls being a rel- atively uniform group of birds. A 42-page overview & chapter deals with the tax- onomy, biogeography, de- * mography, plumage varia- tion, and breeding biology within the Laridae. This is followed by comprehen- sive individual accounts for the nine species breed- ing regularly in Britain (about 25 pages each, but 64 on Black-legged Kittiwake [Rissa tridactyla]) and by chapters on rare species (17), on methods used to study gulls, on urban gulls (considered a big problem in Britain), and on conservation and management. There is a “selected bibliography” that is certainly sufficient to satisfy all but the most demanding reader (a full bibliography was considered too extensive to be included) and a few statistical appendices, as well as one dealing with the taxonomy of the Herring Gull (Larus argentatus) complex in Europe. Compared to a similar area of Canada, Britain is relatively rich in gulls: only five species breed in the Maritimes, for instance. In addition, there has been a great deal of research on British gulls, especially Herring Gull, Lesser Black-backed Gull (Larus fus- cus), and Black-legged Kittiwake. Consequently, the author has a lot of material to work with. Moreover, a substantial proportion of it was actually amassed by Coulson and his students (including Canadians John Chardine and Julie Porter) over a span of nearly Gulls John C. Coulson a \ 70 years. This is the summary of a lifetime’s work by a very industrious and distinguished (and long- lived) scientist! In his Foreword, Coulson mentions an early inter- est in banding. This seems to have continued because the book includes many maps of band recoveries and movements figure prominently in the species ac- counts. Where the author has researched the species extensively (Herring Gull, Black-legged Kittiwake) he gives many details from personal field experience. These anecdotes are among the most enlightening in the book. The methods chapter devotes several pages to banding as a research technique and includes a de- scription of how the author discovered and developed the use of Darvic to make colour-fast plastic bands. This material is still used in many avian studies world- wide and has been an important component of my own research on auks. For many years, the Canadian Wildlife Service sourced its colour bands from John Coulson’s technician at Durham University! The book, printed in China, is nicely produced on glossy paper with excellent maps, charts, and photo- graphs. John Coulson was a pioneer in the study of seabirds, especially long-term population studies. He was the source for many methods still current in the field. He writes with the authority appropriate to his seniority and lifetime of research. Yes, the ac- counts are a little parochial: we do know something about Herring Gulls and Black-legged Kittiwakes in North America, but Coulson makes no pretence that these are other than United Kingdom-based accounts. Despite its slightly narrow focus, Gulls is a must for anyone interested in any branch of Ornithology. ToNy GASTON Ottawa, ON, Canada ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 Ospreys: The Revival of a Global Raptor BooK REVIEWS 175 By Alan F. Poole. 2019. John Hopkins University Press. 205 pages, 39.95 USD, Cloth or E-book. Many of us are familiar with the catastrophic declines in Bald Eagle (Haliaeetus leu- cocephalus) and Peregrine Falcon (Falco peregrinus) " populations owing to DDT & and their subsequent recov- ery in at least parts of their ranges. But many may notbe & aware of the extent to which } Osprey (Pandion halieatus) were similarly affected. Alan ™= ia anil Poole is clearly an authority on the subject, ae studied Ospreys in New England for over 35 years and authoring numerous scientific publications on the subject. The title of this new offering is a bit of a misno- mer, however, as the book covers a lot more ground than just the recovery of the species. An initial Introduction sets the stage and “A Hawk that Fishes” (Chapter 2) describes the Osprey’s morphological adaptions, including differences among subspecies. The “Geography of Ospreys” (Chapter 3) summa- rizes the status of different Osprey populations across its global range, including Canada (citing several pro- vincial atlassing efforts). Subsequent chapters cover foraging ecology, nesting behaviour, and migration, with the final two chapters discussing threats and so- lutions and the future outlook for Osprey. With such a cosmopolitan species and the breadth of topics he touches upon, Poole has done an admirable job con- structing a cohesive narrative. Poole’s new book also brings into the discussion technological advances such as satellite telemetry, ge- olocators, and DNA evidence that have helped further our understanding of Ospreys over the last 30 years. Clearly willing to share the credit, he includes side- bar profiles on other Osprey researchers—remind- ing us how it takes a global village to understand this wide-ranging raptor. He also mentions where our un- derstanding of Osprey is lacking, such as the period OSPREYS THE REVIVAL OF A GLOBAL RAPTOR when young Osprey are learning to forage and dur- ing their first migration. I would like to have learned what factors exclude Osprey from breeding in tropi- cal South America and Africa, given they overwin- ter there and reside year-round in Australia—but per- haps that will have to wait for Poole’s next book. Scientific chops notwithstanding, Poole’s very readable prose makes it accessible for the layper- son despite the quantity of information presented. For example, when discussing Osprey learning to fly, he remarks “As any human pilot will tell you — fly- ing is easy; it’s the landings that are tough” (p. 104) and mentions how human mothers with a houseful of rowdy teenagers might sympathize with female Ospreys at the nest. The book also benefits from over 100 excellent, decent-sized photographs and illustra- tions of Ospreys and their habitats, covering all man- ner of behaviour. Well laid-out maps showing Osprey movement patterns (from telemetry data) and several charts add to the fantastic visuals. On a very minor note, it would have been informative to have a den- sity map of breeding Osprey in North America, but perhaps there are insufficient data. More Canadian content (and calling Nunavut a territory rather than a province on p. 44) would also have been welcome, but I suppose one can only ask so much with a global rap- tor. I do wish however, that all species names be cap- italized or none; for example, Osprey is capitalized but Black Caiman and Agami Heron are not (p. 133). My first childhood encounter with Ospreys was watching one catch a fish in a northern Ontario lake, only to see it lost to a marauding Bald Eagle. Since then, despite being a professional biologist, I have learned woefully little about this remarkable rap- tor. Ospreys: The Revival of a Global Raptor has greatly helped me address such gross negligence. I thoroughly enjoyed reading it and would highly rec- ommend it to both novice naturalist and experienced birder alike. ROBERT F. FOSTER Northern Bioscience, Thunder Bay, ON, Canada ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 176 ZOOLOGY THE CANADIAN FIELD-NATURALIST Vol. 133 The New Beachcomber’s Guide to the Pacific Northwest By J. Duane Sept. 2019. Harbour Publishing. 432 pages and 880 photographs, 32.95 CAD, Paper. A Field Guide to Marine Life of the Protected Waters of the Salish Sea By Rick M. Harbo. 2019. Harbour Publishing. Pamphlet, 7.95 CAD, Folding Guide. A Field Guide to Marine Life of the Outer Coasts of the Salish Sea and Beyond By Rick M. Harbo. 2019. Harbour Publishing. Pamphlet, 7.95 CAD, Folding Guide. My recent move from On- tario to British Columbia (BC) introduced me to a number of new environs, each with their own suite of organisms. Mountains, gla- ciers, and temperate rain- forests were all new to me. Perhaps most exciting of all faa were my first encounters with shoreline and inter- tidal life on the Pacific ! ——— Ocean. Being more acquainted with the Carolinian Forest and Great Lakes shores, I was unable to iden- tify most of what I saw. This similar situation must present itself to many who visit Canada’s Pacific Coast or the United States’ Pacific Northwest for the first time. A series of new field guides, all by the same BC- based publisher, aim to provide a quick introduction to the species found on the shorelines and beaches of BC and the Pacific Northwest. Two different for- mats are employed. The first is a full field guide: The New Beachcomber’s Guide to the Pacific Northwest by J. Duane Sept. This compact but hefty softcover book gives a detailed introduction to life in the in- tertidal zone. A brief opening chapter explains tides, shoreline zones, and microhabitats within the coastal ecosystem. What follows is 300+ pages of accounts of the most common invertebrate, algae, lichen, and plant species to be found on the beach. Each species is given one or two full colour photographs, a com- mon name, scientific name, and brief outline of range, physical description, and life history notes. Species accounts proceed taxonomically with sections on sponges; anemones, hydroids, and jellies; comb jel- lies; worms; molluscs; lampshells; arthropods; bryo- zoans; echinoderms; tunicates; fish; seaweeds; plants; and lichens. No keys or identification guides are pro- vided, so the book is best used by flipping through to find a photo that looks most like the strange crea- et Tg: Buane-Sept... ture you just found under a rock. For easier compar- ison, full colour plates of a range of limpets, snails, bivalves, and urchins are provided separately. This book is big, certainly bigger than a back pocket, but will likely provide greater chance of identification for those that must know about everything they find. Parallel to, but separate from, this approach are the two pamphlets written by Rick Harbo: 4 Field Guide to Marine Life of the Protected Waters of the Salish Sea and A Field Guide to Marine Life of the Outer Coasts of the Salish Sea and Beyond. These two works take a similar approach to quick identifi- cation of intertidal life, but are definitely designed to be thrown into any pocket. In order to accommodate a drastic reduction in size, a more targetted region and fewer species are included. Both guides focus on the Salish Sea, the region roughly encompassing the marine region east of Vancouver Island, west of the lower BC mainland, and stretching down into Puget Sound in Washington State. The two guides roughly divide this region into an inner protected region and an outer coastal region. Understandably, there is some overlap in species. Like Sept’s book, species accounts are presented taxonomically. Sticking with the condensed format however, fewer biological de- tails are given for each species. Despite these restric- tions, each species still gets a full colour photograph and over 75 species are included in each pamphlet. All three of these publications can be valuable tools for those venturing out on the Pacific Coast. The cost is very reasonable and durable construction en- sures that they should survive more than a few trips to the shore. The only difference is the relative size and biological depth that each provides. Certainly, any one of these works would be a valuable purchase for those, like me, who want to go to the shore and be able to identify most of the fascinating lifeforms that they will encounter. JOEL F. GIBSON Curator, Entomology, Royal BC Museum, Victoria, BC ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 Bats: An Illustrated Guide to All Species BooK REVIEWS 177 By Marianne Taylor. Photography by Merlin D. Tuttle. 2018. Smithsonian Books. 400 pages, 39.95 CAD, Cloth. Part lavishly illustrated desk- top field guide, part coffee table book, this stunning hardcover is a triumph of production quality. From the cover through to the in- dex, this book has the look and feel of a text three times its price. Not only do the colour saturation and satisfying weight of paper stock do justice to the in- credible images, the book is intuitively organized and rendered in exquisite detail. Starting with a well written and accessible 50- page introduction section that offers an overview of everything from evolution, diversity, and biology to behaviour and bat-human relationships, the bulk of this book is in the species descriptions, organized by suborder, family, and species. As advertised, it cov- ers the 1384 currently recognized bat species of the world, following the 2018 bat taxonomic review by Simmons and Cirranello (2019). Each species is rep- resented by at least a portrait, as well as its distribu- tion (details and map), physical description, and a few interesting facts. The length, weight, and IUCN sta- tus of each species is also provided. Although lacking a species key and not designed as a true field guide, the level of detail in the fam- ily and genus descriptions is more reminiscent of a textbook than a popular science entry. While Taylor largely avoids jargon throughout, the long-form de- scriptions interspersed between the species accounts do have a noticeably different tone. Casual readers will likely enjoy the more accessible Introduction section and enjoy the book as a curio for its beauti- ful photography and interesting facts. There is cer- tainly enough here for those with a keener interest, but they will have to seek out further reading on their own as the book provides no citations for the infor- mation presented. The highlight of this book is certainly the photo- graphy, much of which captures night-time scenes. The bat portraits are all the more impressive for the way they capture the likeness of their subject. It should be said that this is a good resource to win over the bat-fearing folk among us—aside from a handful of very unusual looking adaptations, bats are by and large endearing creatures and Tuttle captures them with remarkable skill. Taking charismatic photos that show a bat’s natural expression (i.e., avoiding squint- ing eyes from a nocturnal animal exposed to flash) is not an easy task, but the night-time photography, and especially the action shots, in this book are sim- ply unparalleled. If the author’s and photographer’s objectives were to create a spectacular photo inventory of bat species, prefaced by interesting front matter and interspersed with highly detailed family and genus descriptions, they have met and exceeded their goals. The care, time, and effort put into this book are obvious. Dr. Tuttle has studied and photographed bats for 60 years, and his photo collection combined with Taylor’s clear writing are what make this book possible. The result is a gorgeous, informative catalogue of species which absolutely deserves a place on your bookshelf. Literature Cited Simmons, N.B., and A.L. Cirranello. 2019. Bat Species of the world: a taxonomic and geographic database. Accessed 2 October 2019. http://www.batnames.org. HEATHER A. CRAY Halifax, NS, Canada ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 178 THE CANADIAN FIELD-NATURALIST Vol. 133 Mammal Tracks and Sign: A Guide to North American Species. Second Edition By Mark Elbroch with contributions by Casey McFarland. 2019. Globe Pequot / Stackpole Books. 680 pages, 49.95 USD, Paper, 47.50 USD, E-book. The first edition of this mag- nificent guide was published 16 years ago. As Mark EI- broch explains in his Pref- ace, the second edition is less personal—though it re- tains personal stories as introductions to the chap- ters—and more rigorous in terms of added depth of de- tail. It contains less about the “tools of the trade” (p. v), because basic infor- mation is now readily available through increasing online resources. Some adjustments in species in- clusion were made, primarily removing any south of Mexico. Surprisingly, the second edition is shorter than the first (by just over 100 pages), a result of being tightened up, with white space at a premium and pho- tos sometimes smaller. In sum, if you have the first, you wouldn’t be amiss to acquire this edition as well. Senior author Elbroch and contributor Casey McFarland have extensive experience tracking an- imals. They are both certified Senior Trackers un- der the CyberTracker Conservation program, started in South Africa by biologist David Lieberman, who was inspired to develop CyberTracker freeware fol- lowing his experiences with Kalahari Desert bush- men, whose lives depend on their ability. Elbroch is involved as well with Pathera, a big cat conservation organization, as Director of its Puma Program. His 10 books have been well received, the first edition of this guide being one of several of his books to win a National Outdoor Book Award. The first chapter provides a brief and succinct in- troduction to this prodigiously illustrated and com- prehensive guide. “Tracking is ... field ecology”, the authors note, that “grounds us in the natural history of a place” and provides “a natural bridge to science” (p. 2). Extensive, detailed knowledge is required to track successfully, but, they caution, one must al- ways keep in mind the dangers of drawing conclu- sions without it. A hundred pages are required in Chapter 2 to cover “Mammal Tracks and Track Patterns”. Feet make tracks, Elbroch reminds us, and the first section of this chapter follows its opening story with descrip- tions and sketches of the morphology of mammalian feet. But the tracks these feet produce have their own, obverse morphology, and eight questions are pre- sented that relate the tracks to the feet that created Mammal Tracks & Sign A Guide to North American Species. SECOND EDITION inte alr ints by Casey McFarland them. Track morphology can be affected by many things, including substrate (sand, soil, snow), sexual dimorphism (helpful in sexing individual animals), and the animal’s age. The many types of locomotion (gaits, hops, gallops, etc.) are covered in detail, with diagrams and illustrations of each, before the discus- sion turns to interpretation of tracks. This is an imag- inative, speculative process that requires the tracker “to build a working hypothesis and test it” against later observations of the animal’s behaviour as evi- denced in its tracks: “we either toss out our original hypothesis and create a new one, or continue to re- fine and support it” (p. 45). This lengthy chapter con- cludes with a “Reference guide to mammal tracks”, 46 pages of plates of life-size tracks, each example— as throughout the book’s illustrations—keyed to the long, final section of species accounts. Seven subsequent chapters follow this format of text plus grouped illustrations, the latter having their own colour-coded page edges making these sections easy to find. In the course of their movements and other activities, mammals leave behind plenty of sign which offers rich and diverse clues to who they are and what they were doing. Thus, the emphasis shifts from tracks to sign, beginning with a discussion in Chapter 3 of the “Runs, Paths, and Eskers” created by the movements of animals. (Also known as trail castings, eskers are the backfilling materials depos- ited in snow tunnels created by the smallest mammals [p. 113].) Animals eat and then excrete, the subject of the fourth chapter. Scat is the primary topic, be- cause urine, being hard to find, remains understud- ied generally. It can have its uses, however—Elbroch has an amusing account of tracking a Bobcat by sniff- ing the urine it had sprayed on “scent posts”, mov- ing and sniffing from post to likely post to figure out its movements (p. 445). The short Chapter 5, “Nests, Lodges, and Other Constructions”, describes the of- ten difficult to identify shelters animals “construct ... from materials they collect and manipulate” (p. 178); the shelters are organized under the likely places to find them. The next three chapters cover sign left on the ground, beginning with discussion in Chapter 6 of “diverse signs” (p. 192) created in the course of bedding down, rooting around for food, taking a dust bath, or making below-ground dens. Signs are also left “on Fungi, Herbaceous Plants, and Cacti’ (Chapter 7) and “on Trees and Shrubs” (Chapter 8). While these are generally the result of feeding (Chapter 7), they can also be produced by climbing, stripping off bark, browsing on twigs, and taking advantage of tree cav- 2019 ities (Chapter 8). Fourteen pages of colour plates are required to detail the “Sign on mast crops” alone. The penultimate chapter is not for the squeamish—it deals with “Interpreting Prey Remains” in gruesome, full-coloured detail. A suite of forensic skills is re- quired when dealing with remains, one of the most difficult areas of tracking to decipher. This chapter is full of questions to consider and authoritative ref- erences to consult; techniques and data to aid with interpretation are offered, including, for example, a chart of the spread between upper and lower incisors of various mammals. The topics of the preceding chapters are the basis for the rich data presented on each species described in Chapter 10, “Species Accounts”. The Revised Checklist of North American Mammals North of Mex- ico, 2014 (Bradley et al. 2014) provides the taxonomic orders for 261 pages of individual species accounts. This is the most technical part, with data presented in small print, including measurements of front and hind tracks, trail strides of various kinds, and other measurable elements, all organized along the lines of the chapter headings. Sketches and photographs abound. The species themselves are named but not described—the evidence they leave behind is the constant topic. The trails data sections are frequently followed by notes describing that species’ movements under varying conditions. By now, the reader can be forgiven for conclud- ing—with good reason—that using tracks and sign is a more demanding and complex means of mam- mal identification than looking at the mammal itself. To summarize the authors, it’s a matter of detec- tive work, close observation, detailed and intricate knowledge, plus a healthy dose of humility—the ca- pacity to accept and admit one’s inevitable mistakes. Becoming a good tracker takes time, experience, and patience plus the helpful analyses presented in this BooK REVIEWS 179 excellent guide. It’s on the heavy side for packing in the field, but it’s well organized and indexed, so use- ful whether in the field or in the study. Anyone inter- ested in the topic will learn from this book. It opens up a new universe for exploring the natural world for those coming fresh to the topic and no doubt contains much for those with experience. As the authors note early on, it’s not only easy to get it wrong, but costly in terms of natural history. This guide will help. A final note: Stackpole Books is to be congratu- lated on its publication of another fine field guide in its tracks and sign series. This one on mammals joins guides on birds (co-authored by Elbroch; Elbroch et al. 2001), reptiles and amphibians (Tkaczyk 2015), and insects (Eiseman ef a/. 2010), all similar in form and topics covered. Literature Cited Bradley, R.D., L.K. Ammerman, R.J. Baker, L.C. Brad- ley, J.A. Cook, R.C. Dowler, C. Jones, D.J. Schmidly, F.B. Stangl, Jr., R.A. Van Den Bussche, and B. Wiirsig. 2014. Revised Checklist of North American Mammals North of Mexico, 2014. Museum of Texas Tech Univer- sity, Lubbock, Texas, USA. Accessed 7 November 2019. http://www.nsrl.ttu.edu/publications/opapers/ops/ OP327.pdf. Eiseman, C., and N. Charney. 2010. Tracks & Sign of Insects and Other Invertebrates: a Guide to North American Species. Stackpole Books, Mechanicsburg, Pennsylvania, USA. Elbroch, M., E. Marks, and C.D. Boretos. 2001. Bird Tracks & Sign: a Guide to North American Species. Stackpole Books, Mechanicsburg, Pennsylvania, USA. Tkaczyk, F.A. 2015. Tracks & Sign of Reptiles & Am- phibians: a Guide to North American Species. Stackpole Books, Mechanicsburg, Pennsylvania, USA. BARRY COTTAM Ottawa, ON, and Corraville, PEI ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 180 THE CANADIAN FIELD-NATURALIST Vol. 133 The Rise of Wolf 8: Witnessing the Triumph of Yellowstone’s Underdog By Rick McIntyre. 2019. Greystone Books. 304 pages, 34.95 CAN, 26.95 USD, Cloth. Rick McIntyre has spent over 40 years observing § wolves in national parks, including Denali, Glacier, and Yellowstone. The Rise of Wolf § is part one of a planned trilogy of his nearly 25 years of observ- | ing Yellowstone wolves. § What is significant about this book is that the author is the source for this story. For the past 20 plus years, many people have written about McIntyre, his col- leagues, and the Yellowstone wolves through inter- views and television documentaries. But now we get to hear from the man himself. And the book doesn’t disappoint. The profiles of individual wolves found in this book are unparalleled for a carnivore, espe- cially here in North America where most carnivores are elusive, usually being active away from people, in forested areas, and at night. As Doug Smith says in the Afterword, “Rick has become a global ambassa- dor for wolves ... what you have just read 1s one of the most personal and in-depth insights into wolves ever produced” (p. 267). I agree! The only resource that I can compare it to is Jane Goodall’s The Chimpanzees of Gombe (1986, Belknap Press), which describes the lives of individual chimpanzees in unprecedented de- tail. The same is true of McIntyre’s observations of wolves; his knowledge of individuals provides a re- markable real-life narrative of wild wolves. Hoping just to see a wolf during the first sum- mer (1995) that they were reintroduced to the park (pp. 19-20), McIntyre observed a pack for hours on his first day there and went on to observe more wild wolves than any other human in history, as docu- mented in the Afterword of this book (pp. 265-267). He has had over 100 000 wolf sightings as of January 2019 (p. 267), aided by radio-telemetry, spotting scopes, and a cadre of wildlife watchers and profes- sionals assisting him. He once went out an indefat- igable 6175 consecutive straight days into the park, saw wolves 892 days in a row, and has compiled a journal of over 12 000 pages (pp. 265-266)! Over the years, he has been a critical resource for thousands of people in observing wild wolves which is important because observing carnivores is a major economic driver in many regions (Way and Bruskotter 2012), with people specifically visiting regions where car- nivores are protected to have “peak life experiences” (McIntyre 2016). Rick was an important source for my own book, My Yellowstone Experience (2013, White Cottage Publishing), so it is probably not sur- prising that I was revelling in the opportunity to re- view this book. After a brief background describing his childhood and earlier park service jobs, mostly up in Alaska at Denali and in Texas at Big Bend, the book starts in 1994 in Yellowstone where McIntyre was the coun- try’s first “Wolf Interpreter” for the park service. There he primed about 25 000 park visitors that sum- mer about the return of wolves to Yellowstone (p. 10). The next few chapters discuss the reintroduc- tion project—some of the human actors involved as well as the early wolves, brought from Alberta (near Jasper National Park) to Yellowstone. We learn that Wolf 8 of the Crystal Creek pack was the smallest male brought to Yellowstone, a Coyote-like 72-pound (32.6-kg), grey coloured male (p. 15) called “the little guy” (p. 29). He was frequently picked on by his three larger brothers while they were held in an acclima- tion pen to reduce their homing instincts (Chapter 3). The most important story in the book, and argu- ably one of the most important events in the annals of Yellowstone wolf reintroduction, was Wolf 8 leaving his birth pack as a 1.5-year-old to join up with Wolf 9 and her eight pups of the Rose Creek pack (Chapter 6). Wolf 9’s mate #10 had been illegally killed out- side the park in April 1995, so 9 and her pups were brought back to the park soon after and kept in an ac- climation pen until October. Just before the planned release of the wolves, 8 showed up and greeted and fed two of the pups who had escaped the pen during a summer storm. Park employees immediately released the rest of the wolves and the ten wolves formed a co- hesive pack led by step-father wolf 8 who protected his new pack (e.g., pp. 48-50) and regularly had to go out and hunt prey animals much bigger and stronger than he was (pp. 248-249). Wolf 8 proved his worth by eventually fathering 54 pups over his life (p. 232). McIntyre moved full-time to Yellowstone in 1999 to immerse himself in the wolves’ world (p. 182). Thus, The Rise of Wolf § is highly personalized as it is about individual wolves, similar to Wild Wolves We Have Known which details the lives of famous wolves studied around the globe (Way 2016). But two things stand out in MclIntyre’s book: 1) it is about individ- ual wolves studied in one area; and 2) studying indi- viduals is becoming more and more frequent, despite the stereotype that biologists shouldn’t ascribe names and emotions to their study subjects. Thus, this book normalizes the importance of individual—not just populations of—animals. We clearly see that wolves 2019 have individual personalities, are social, and love to play—all characteristics of higher-order animals like humans. We learn that play is vitally important for bonding within packs as well as preparing them for real world circumstances; McIntyre provides a list and vivid descriptions of these games throughout the book (e.g., pp. 210, 227, 228, 242). The culmination of the book was “The Battle of Specimen Ridge” (Chapter 27) between Wolf 8’s Rose Creek pack and his step-son 21’s Druid Peak pack. McIntyre’s Shakespearean description of the encoun- ter is absolutely riveting. He noted that 8’s determina- tion to battle the much larger 21, who was undefeated in battle situations with other wolves (p. 263), was the bravest thing he had ever seen, “a fight that 21 could not lose and one 8 could not win” (p. 257). Wolves 8 and 21 were never known to have killed other wolves and there are repeated accounts throughout the book of them letting rivals go (p. 262). This is exceptional because they were both alpha wolves for years and wolves killing each other is the most common cause of death where people don’t kill them (p. 179). Wolf 8 was battle-worn and had many debilitating inju- ries (pp. 247-248). Yet he soldiered on, fueled solely by willpower (p. 249), until June 2000 when he died from either an Elk’s kick to his head or drowning soon afterward (p. 260). Dying in combat, McIntyre noted, was an honourable ending to his life—it was a good death (p. 261). At the beginning of the book McIntyre noted (p. xix) that all the elements of a great tale are present, in- cluding warfare, betrayal, murder, bravery, compas- sion, empathy, and loyalty, yet a literary genius such as Dickens or Shakespeare was not available to write BooK REVIEWS 181 this story. I disagree. I believe that McIntyre himself was the perfect person to write this wonderful ac- count. He combines his extraordinary level of obser- vation of wolves with great storytelling (p. 265). For fans of Yellowstone or wolves, this book is priceless, with a historic feel to it that is palpable. I wholeheartedly recommend it and believe that read- ers interested in nature, carnivores, and individ- ual animals will be fascinated to learn about the wolves described here and in forthcoming volumes of McIntyre’s planned trilogy. McIntyre (2016) has de- scribed viewing Yellowstone wolves as a “peak life experience”. Reading this book and gaining the in- sights into Yellowstone wolves that McIntyre pro- vided in unparalleled detail was a ‘peak reading ex- perience’ for me! I eagerly await the next edition of the Yellowstone wolf saga. Literature Cited McIntyre, R. 2016. A peak life experience: watching wolves in Yellowstone National Park. Yellowstone Science 24: 44-46. Way, J.G. 2016. [Book Review] Wild wolves we have known: stories of wolf biologists’ favorite wolves. Ca- nadian Field-Naturalist 130: 85-87. https://doi.org/10. 22621 /cfn.v13011.1800 Way, J.G., and J.T. Bruskotter. 2012. Additional consid- erations for gray wolf management after their removal from Endangered Species Act protections. Journal of Wildlife Management 76: 457—461. https://doi.org/10. 1002/jwmg.262 JONATHAN (JON) Way Eastern Coyote/Coywolf Research, Osterville, MA, USA ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). OTHER To Speak for the Trees: My Life’s Journey from Ancient Celtic Wisdom to a Healing Vision of the Forest By Diana Beresford-Kroeger. 2019. Random House Canada. 289 pages, 32.00 CAD, Paper. Diana Beresford-Kroeger is ge aname that has become in- = creasingly well-known in recent years. Awareness and || acceptance of her work has |} been a long time coming. | Variously described as a classical botanist, medical biochemist, and protector Bags: of forests, Fellow of The Royal Canadian Geograph- §& ical Society, and—in a de- [az lightful poke at the conservative elements that re- sist her message—“an enemy of the people” (https:// www.insideottawavalley.com/news-story/5463236- diana-beresford-kroeger-is-now-an-enemy-of-the- public/; accessed 10 September 2019), she is at once grounded in local activities and at home in world-wide travels through various media. She has published some 300 scientific papers, but the public has come to know her through her previous half-dozen books (or more, depending how you count the editions), pub- lished over the past 20 years, and more recently the film Call of the Forest, now streaming on the Internet 182 (https://www.youtube.com/watch? v=I0asL2LfPFk). Not bad for someone who echews email and social media as taking too much time from her real work (D. Beresford-Kroeger pers. comm. 13 August 2019)! Trees have been at the centre of her explorations and writings, and she combines in her study and ad- vocacy two broad and apparently disparate streams, the scientific fields of botany and biochemistry and the esoteric realm of ancient Celtic beliefs and prac- tices regarding nature. Now, in her latest book, she relates the personal story behind this double exper- tise. Zo Speak for the Trees doesn’t begin with the Ancient Celtic Wisdom noted in the subtitle; rather, it begins with an account of the difficult life of a mis- placed soul whose journey began in trauma and trag- edy. These hard beginnings contain the roots of the dual streams noted above. Her father’s family came from a long lineage of wealth and status, her moth- er’s from an even longer lineage dating back to the Celtic/Druidic world, based in Lisheens Valley, County Cork, Ireland. The former was the source of the trauma; tragedy—the deaths of her parents within months of each other when Diana was 12—propelled her initially into the world of science as shared with an uncle who took her in. The serendipitous discovery of her capacious intellect, revealed through her pho- tographic memory, lifted her into deeper studies, at university but also while under the care of her Celtic relations with whom she spent her summers. That ag- ing community decided she would be the ideal expo- nent of their traditional knowledge and proceeded to teach her everything they knew. Thus, her increas- ingly happier world came to be divided, productively, between science and indigenous Celtic lore. The first six chapters cover all this and more. In Chapter 7, “But Where are the Trees?”, the book moves on to its central substance, her love of, and life-long relationship with, trees and her increasing awareness of their absolute essentiality to the sur- vival of other forms of life on earth. The title ques- tion for this chapter relates to her realization that the ancient forests of Ireland had largely disappeared. Thus began her quest to understand what happened and why, a quest that ultimately took her around the world, inspired a great deal of original research, and led to the insight—and she is marvellous at gain- ing insights—that if the trees disappear, we disap- pear, the dependency is that tight (p. 114). This fun- damental principle is at the core of a life of activism and the development of her “global bioplan” (p. 159) for addressing climate change, the gravest single problem faced by humanity today. Her research re- sults coalesced into this practical plan, one she lives, THE CANADIAN FIELD-NATURALIST Vol. 133 with her husband, Christian Kroeger, on 160 acres (64.75 hectares) of land they call Carriglaith, near Merrickville, south of Ottawa. Chapter 11, “My Own Work in My Own Way”, recounts the story of devel- oping this land into what amounts to a gene bank of trees hardy enough to live in northern climes as tem- peratures rise. The paragraph above compresses the second half of Part One of the book. As her work progressed, she came to understand that she “could serve as a bridge between these two worlds, the ancient and the sci- entific” (p. 98). Her account is bravely personal, for she is frank about how the suppression she experi- enced growing up and into early adulthood continued through misogynistic workplaces and the reluctance of the scientific community to accept the esoteric and spiritual elements her research revealed. She took strength from these experiences, however, coming to realize that her background was essential to her abil- ity to think for herself, to refuse to suffer fools gladly, to stand firm for her growing beliefs in the power of trees, and to bring this into form through her actions, from the land she and Christian manage to promot- ing her hard-won knowledge through the books and documentary noted above, to advising many groups and governments. Examples of her efforts are de- scribed in the final chapters of Part One, which con- cludes with her “Philanthropy of the Mind”, the com- bination of her “scientific knowledge and ... energy” (p. 76) that drives her and, she hopes, will inspire all of us to act. In Part Two, she returns to her roots to provide us with “one more gift ...: my annotation of the Ogham script, ... the first alphabet of Europe, in which ev- ery letter is named for a tree or an important com- panion plant of trees” (p. 189). There are 20 letters, each receiving an account that mingles poetically scientific knowledge of the trees with ancient Celtic lore, starting with A/Ailm, Pine, and ending with Z/ Straif, Blackthorn. Old uses, now largely lost, medic- inal properties, and notes on the alphabet itself are melded into each of these brief notes. This is a courageous, highly readable book, yield- ing insights of value and leaving us—field natural- ists, researchers, nature lovers—with much to think about. The discoveries Diane Beresford-Kroeger has made are at last entering the mainstream, as a source of knowledge and inspiration for the increasing num- bers of people concerned with the physical and future state of the planet. BARRY COTTAM Ottawa, ON, and Corraville, PEI ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 The Overstory: A Novel BooK REVIEWS 183 By Richard Powers. 2018. W.W. Norton. 512 pages, 27.95 USD, Paper. Readers of The Canadian Field-Naturalist might rea- sonably wonder why a nov- el would be reviewed in its pages. A precedent of sorts was set several issues ago with our first review—at least on my watch as Book Review Editor—of a book of poetry, Alice Major’s Welcome to the Anthropo- cene (Citron 2017). But a novel? This isn’t just any novel, however—it’s a brilliant evocation of the life and times of trees as understood, misunderstood, used and abused, by the one species on Earth that supposes itself to be dominant—Homo sapiens, you and I and all the other 8 billion or so humans alive to- day. The entire book, from its title on out, is struc- tured along the lines of and about trees. The title is a play on words, referring at a literal level to the over- arching canopy of trees in a forest, the collective crowns of the tallest trees, while signalling the au- thor’s ambition to tell as fully as possible the richly complex story of trees as sentient beings and their in- teractions with humans. This Powers does brilliantly, in language that flows with an unusually high degree of authen- ticity. The novel has four sections: Roots, Trunk, Crown, and Seeds. The first is the only one with sub- titled chapters, nine of them, named for the charac- ters introduced in each. I was a bit puzzled at first to read separate stories of such a disparate and seem- ingly disconnected group of people. Each however had some kind of connection to trees and these early chapters set the stage for Powers’ interweaving, in various combinations, the stories of these people in ways that allow him to address his panoramic pur- pose. We humans take multifaceted approaches to- ward trees, whether as sources of food and medicines, shelter and fuel, beauty and meditative healing, or the many lines of work they provide, from scientific stud- ies to logging and milling up construction materials. Powers addresses our approaches in ways that reveal our inherent, eternal conflicts arising from our need to preserve ourselves—read lifestyles, economies— and our yearning for connectivity and transcendence. Powers does not resolve these conflicts, however. While reading his book, it’s possible to hate and fear Richard Powers the destruction our ‘normal’ lumbering practices are inflicting on the forests while having sympathy for the people whose jobs are at stake if those practices cease, to engage emotionally with the activists go- ing to increasing lengths to end the destruction while gasping at some of their decisions. The world of sci- ence is also revealed as conflicted, especially in the characterization of ‘maverick’ versus conventional science through the career of Patricia Westerford and her supportive husband Dennis, based on the career pathway of the very real Diane Beresford-Kroeger and her husband Christian Kroeger. Powers has done his homework—he has a reputation as a prodigiously inquisitive researcher—yet weaves his learning into a highly imagined fictional form. The author’s deliberate unwillingness to mission- ize, to leave the reader pondering these dilemmas, is the strength and power of the book, and reason enough, I think, to review it in these pages. Powers claims his own life was changed as a result of re- searching and writing this, his twelfth novel. In a re- cent interview, he declared that “Trees are among the very largest, longest-lived, most successful, and most collaboratively social forms of life on the planet. They live, all at once, in the sky, on the surface, and under the ground ... We will learn, as Thoreau says, to resign ourselves to the influence of the earth, or we will disappear” (Brady 2018). The clarity of his personal position notwithstanding, he has managed to resist judgement and condemnation while creating a credible and complex picture of human activities in relation to trees. A number of excellent novels deal- ing with nature and natural history have come out in recent years, and this is one of the best, shortlisted in 2018 for the Man Booker Prize, winner in 2019 of the Pulitzer Prize in Fiction. Powers sends readers a mes- sage worth the time spent to listen to it. Literature Cited Citron, M. 2017. [Book Review] Welcome to the Anthro- pocene. Canadian Field-Naturalist 131: 372-373. https:// doi.org/10.22621/cfn.v13114.2087 Brady, A. 2018. Richard Powers: writing “The Overstory’ quite literally changed my life. Chicago Review of Books. Accessed 7 September 2019. https://chireviewof books. com/2018/04/18/overstory-richard-powers-interview/. BARRY COTTAM Ottawa, ON, and Corraville, PEI ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 184 NEw TITLES Prepared by Barry Cottam THE CANADIAN FIELD-NATURALIST Vol. 133 Please note: Only books marked *¢ or * have been received from publishers. All other titles are listed as books of potential interest to subscribers. Please send notice of new books—or copies for review—to the Book Review Editor. +Available for review *Assigned Currency Codes: CAD Canadian Dollars, AUD Australian Dollars, USD United States Dollars, EUR Euros, GBP British Pound. BOTANY The Life of Plants: A Metaphysics of Mixture. By Emanuele Coccia. Translated by Dylan J. Montanari. 2018. Polity. 166 pages, 22.95 USD, Paper. Plantes de l’enclave argileuse Barlow-Ojibway — 2 Québec 2018. Par Pierre Martineau. 2018. Editions P. Martineau. 218 pages, 40.00 CAD, Cloth. Free PDF available at http://cikwanikaci.ca/volume/index.html or http://depositum.uqat.ca/785/1/MartineauPierre. pdf. Darwin’s Most Wonderful Plants: Darwin’s Bo- tany Today. By Kenneth Thompson. 2019. Profile Books. 256 pages, 8.99 GBP, Paper. Joseph Banks’ Florilegium: Botanical Treasures from Cook’s First Voyage. By Mel Gooding, David Mabberley, and Joe Studholme. 2019. Thames and Hudson. 320 pages and 180 illustrations, 45.00 USD, Cloth. CONSERVATION & ECOLOGY +The Coastal Everglades: The Dynamics of Social-Ecological Transformation in the South Florida Landscape. Long-Term Ecological Research Network Series. Edited by Daniel L. Childers, Evelyn Gaiser, and Laura A. Ogden. 2019. Oxford University Press. 300 pages, 79.95 CAD, Cloth. Also available as an E-book. Community Ecology. Second Edition. By Gary G. Mittelbach and Brian J. McGill. 2019. Oxford Uni- versity Press. 432 pages, 50.00 CAD, Cloth. Also available as an E-book. Working with Nature: Saving and Using the World’s Wild Places. By Jeremy Purseglove. 2019. Profile Books. 288 pages, 32.99 AUD, Paper. Green Growth That Works: Natural Capital Poli- cy and Finance Mechanisms from Around the World. Edited by Lisa Mandle, Zhiyum Ouyang, James Salzman, and Gretchen C. Daily. 2019. Island Press. 368 pages, 39.95 USD, Paper. The Plant Messiah: Adventures in Search of the World’s Rarest Species. By Carlos Magdalena. 2018. Penguin. 240 pages, 9.99 GBP, Paper or E-book. Wild Capital: Nature’s Economic and Ecological Wealth. By Barbara K. Jones. 2019. University Press of Florida. 304 pages, 60.00 USD, Cloth. After Geoengineering: Climate Tragedy, Repair, and Restoration. By Holly Jean Buck. 2019. Verso Books. 2019. 288 pages, 33.95 CAD, Cloth, 13.99 CAD, E-book. How Did We Get into This Mess? Politics, Equality, Nature. By George Monbiot. 2017. Verso Books. 352 pages, 32.99 CAD, Cloth, 22.95 CAD, Paper, 13.99 CAD, E-book. International Wildlife Management: Conservation Challenges in a Changing World. Edited by John L. Koprowski and Paul R. Krausman. 2019. Johns Hopkins University Press. 248 pages, 74.95 USD, Cloth or E-book. The Anthropocene Disruption. By Robert William Sandford. 2019. Rocky Mountain Books (RMB). 168 pages, 20.00 CAD, Cloth. Rivers of the Anthropocene. Edited by Jason M. Kelly, Philip Scarpino, Helen Berry, James Syvitski, and Michel Meybeck. 2017. University of California Press. 242 pages, 34.95 USD, Paper. Free PDF avail- able at https://doi.org/10.1525/luminos.43 The Shock of the Anthropocene: The Earth, Histo- ry and Us. By Christophe Bonneuil and Jean-Baptiste Fressoz. Translated by David Fernbach. 2017. Verso Books. 320 pages, 25.95 CAD, Paper, 13.99 CAD, E-book. Changing Tides: An Ecologist’s Journey to Make Peace with the Anthropocene. By Alejandro Frid. 2019. New Society Publishers. 208 pages, 19.99 CAD / USD, Paper, 13.99 CAD / USD, E-book. Towards Zero Waste: How to Live a Circular Life. By Féidhlim Harty. 2019. Permanent Publications. 176 pages, 17.95 USD, Paper. ENTOMOLOGY Dance of the Dung Beetles: Their Role in Our Changing World. By Marcus Byrne and Helen Lunn. 2019. Wits University Press. 240 pages, 35.00 USD, Paper. 2019 Forests and Insect Conservation in Australia. By Tim R. New. 2019. Springer International Publishing. 291 pages, 169.99 USD, Cloth or Paper, 129.00, E-book. Gods, Wasps and Stranglers: The Secret History and Redemptive Future of Fig Trees. By Mike Shanahan. 2018. Chelsea Green Books. 208 pages, 14.95 USD, Paper. Moths: A Complete Guide to Biology and Behavior. By David Lees and Alberto Zilli. 2019. Smithsonian Institution Scholarly Press. 208 pages, 24.95 USD, Paper. ORNITHOLOGY +Feed the Birds: Attract and Identify 196 Common North American Birds. By Chris Earley. 2019. Fire- fly Books. 296 pages, 29.95 CAD, Paper. A Chorus of Cranes: The Cranes of North America and the World. By Paul A. Johnsgard. Photographs by Thomas D. Mangelsen. 2015. University Press of Colorado. 208 pages, 29.95 USD, Paper, 23.95 USD, E-book. Great Plains Birds. Discover the Great Plains Series. By Larkin Powell. 2019. Bison Books, University of Nebraska Press. 244 pages, 16.95 USD, Paper or E-book. A Season on the Wind: Inside the World of Spring Migration. By Kenn Kaufman. 2019. Houghton Mif- flin Harcourt. 288 pages, 26.00 USD, Cloth, 256 pages, 14.99 USD, E-book. Close to Birds: An Intimate Look at Our Feathered Friends. By Roine Magnusson, Mats Ottoson, and Asa Ottoson. 2019. Shambhala Publications. 272 pages, 39.95 USD, Cloth. The Art of the Bird: The History of Ornithological Art Through Forty Artists. By Roger J. Lederer. 2019. UCP. 224 pages and 200 colour plates, 35.00 USD, Cloth, 21.00 USD, E-book Oceanic Birds of the World: A Photo Guide. By Steve N.G. Howell and Kirk Zufelt. 2019. Princeton University Press. 360 pages, 368 plates with 2200 co- lour photos, and 114 colour distribution maps, 35.00 USD, Paper. Peterson Reference Guide to Sparrows of North America. By Rick Wright. 2019. Houghton Mifflin Harcourt. 443 pages, 35.00 USD, Cloth. Ornithology: Foundation, Analysis, and Appli- cation. Edited by Michael L. Morrison, Amanda D. Rodewald, Gary Voelker, Melanie R. Colon, and Jonathan F. Prather. 2018. Johns Hopkins University Press. 1016 pages and 936 illustrations, 110.00 USD, Cloth. Also available as an E-book. NEw TITLES 185 Henry Dresser and Victorian Ornithology: Birds, Books and Business. By Henry A. McGhie. 2017. Manchester University Press. 368 pages. 25.00 GBP, Cloth, 30.00 GBP, E-book. RSPB Spotlight Ospreys. By Tim Mackrill. 2019. Bloomsbury Wildlife. 128 pages and 200 colour pho- tos, 12.99 GBP, Paper. Also available as an E-book. A Sparrow’s Life’s as Sweet as Ours: In Praise of Birds and Seasons. By Carry Akroyd and John McEwen. 2019. Bloomsbury. 128 pages, 12.99 GBP, Cloth. To the Ends of the Earth: Ireland’s Place in Bird Mi- gration. By Anthony McGeehan. 2018. Collins Press, Dublin. 248 pages, 24.99 GBP, Cloth. ZOOLOGY +Mammals of Prince Edward Island and Adjacent Marine Waters. By Rosemary Curley, Donald F. McAlpine, Dan McAskill, Kim Riehl, and Pierre- Yves Daoust. 2019. Island Studies Press. 354 pages, 49.95 CAD, Paper. The Hidden World of the Fox. By Adele Brand. 2019. William Collins. 216 pages, 12.99 GBP, Cloth. Beavers: Boreal Ecosystem Engineers. By Carol A. Johnston. 2017. Springer International Publishing. 311 pages, 199.99 CAD, Cloth or Paper, 149.00 CAD, E-book. Common Spiders of North America. By Richard A. Bradley. Illustrations by Steve Buchanan. 2019. University of California Press. 288 pages, 34.95 USD, Paper. Also available as an E-book. Cloth edi- tion published in 2012. Principles of Animal Behavior. Fourth Edition. By Lee Alan Dugatkin. 2019. University of Chicago Press. 576 pages and 529 colour plates, 95.00 USD, Paper or E-book. *Yellowstone Cougars: Ecology Before and During Wolf Restoration. By Toni K. Ruth, Polly C. Buotte, and Maurice G. Hornocker. 2019. University Press of Colorado. 336 pages, 65.00 USD, Cloth, 53.00 USD, E-book. Tunas and Billfishes of the World. By Bruce B. Collette and John Graves. Illustrations by Val Kells. 2019. Johns Hopkins University Press. 352 pages, 241 colour illustrations, and 61 maps, 75.00 USD, Cloth. Freshwater Mollusks of the World: A Distribution Atlas. Edited by Charles Lydeard and Kevin S. Cummings. 2019. Johns Hopkins University Press. 256 pages, 125.00 USD, Cloth or E-book. 186 OTHER Algonquin Wild: A Naturalist’s Journey Through the Seasons. By Michael Runtz. 2019. Fitzhenry & Whiteside. 233 pages, 45.00 CAD, Paper. Beyond the Trees: A Journey Alone Across Ca- nada’s Arctic. By Adam Shoalts. 2019. Allen Lane. 288 pages, 32.95 CAD, Cloth, 15.99 CAD, E-book. Coral Whisperers: Scientists on the Brink. Critical Environments: Nature, Science, and Politics Series. By Irus Braverman. 2018. University of California Press. 344 pages, 85.00 USD, Cloth, 26.95 USD, Paper or E-book. Dinosaurs Rediscovered: The Scientific Revolution in Paleontology. By Michael J. Benton. 2019. Thames & Hudson. 352 pages, 34.95 USD, Cloth. Also avail- able as an E-book. Extinction: A Very Short Introduction. By Paul B. Wignall. 2019. Oxford University Press. 144 pages, 11.95 CAD, Paper. Also available as an E-book. Falter: Has the Human Game Begun to Play Itself Out? By Bill McKibben. 2019. Henry Holt and Co. 304 pages, 28.00 USD, Cloth, 17.00 USD, Paper, 2.99 USD, E-book. Firestorm: How Wildfire Will Shape Our Future. By Edward Struzik. 2019. Island Press. 248 pages, 24.95 CAD, Paper. Cloth and E-book editions pub- lished in 2017. Floating Coast: An Environmental History of the Bering Strait. By Bathsheba Demuth. 2019. W.W. Norton. 416 pages, 27.95 USD, Cloth. Forests in our World: How the Climate Affects Woodlands. By Gunther Willinger. 2019. teNeues Publishing UK Ltd. 240 pages, 90.00 CAD, Cloth. How to Catch a Mole: Wisdom from a Life Lived in Nature. By Marc Hamer. 2019. Greystone Books. 208 pages, 29.95 CAD, Cloth or E-book. Invasive Aliens: The Plants and Animals From THE CANADIAN FIELD-NATURALIST Vol. 133 Over There That Are Over Here. By Dan Eatherley. 2019. William Collins. 336 pages, 16.99 GBP, Cloth, 9.99 GBP, E-book. Journeys in the Wild: The Secret Life of a Camera- man. By Gavin Thurston. Foreword by Sir David Attenborough. 2019. Orion Books. 441 pages, 16.99 GBP, Cloth or E-book. The Natural History of The Bahamas: A Field Guide. By Dave Currie, Joseph M. Wunderle, Jr., Ethan Freid, David N. Ewert, and D. Jean Lodge. 2019. Cornell University Press. 464 pages, 34.95 USD, Paper, 17.95 USD, E-book. Naturalist: A Graphic Adaptation. By Edward O. Wilson. Adapted by Jim Ottaviani. Illustrations by C.M. Butzer. 2019. Island Press. 208 pages, 31.95 USD, Cloth. Nature’s Calendar: A Year in the Life of a Wildlife Sanctuary. By Colin Rees. 2019. Johns Hopkins Uni- versity Press. 360 pages, 32.95 USD, Cloth or E-book. In Nature’s Realm: Early Naturalists Explore Van- couver Island. By Michael Layland. 2019. Touch- Wood Editions. 288 pages, 40.00 CAD, Cloth. Neptune’s Laboratory: Fantasy, Fear, and Science at Sea. By Antony Adler. 2019. Harvard University Press. 256 pages, 39.95 USD, Cloth. *North Pole: Nature and Culture. By Michael Bravo. 2019. Reaktion Books. 256 pages, 24.95 USD, Paper. Rain Comin’ Down: Water, Memory and Identity in a Changed World. By Robert William Sandford. 2019. Rocky Mountain Books (RMB). 360 pages, 22.00 CAD, Paper. Rainforest: Dispatches from Earth’s Most Vital Frontlines. By Tony Juniper. 2019. Island Press. 456 pages, 22.00 USD, Paper, 21.99 USD, E-book. Turning the Boat for Home: A Life Writing about Nature. By Richard Mabey. 2019. Chatto & Windus. 272 pages, 18.99 GBP, Cloth or E-book. The Canadian Field-Naturalist News and Comment Upcoming Meetings and Workshops The Committee on the Status of Endangered Wildlife in Canada The next Wildlife Species Assessment Meeting of COSEWIC will be held 24—29 November 2019 at the Lord Elgin Hotel, Ottawa, Ontario. See how COSEWIC assigns the status to Canadian wildlife spe- cies, the first step in protection and recovery under the federal Species at Risk Act. Please contact ec.cosepac- cosewic.ec@canada.ca for the procedure to attend as an observer at least one week before the meeting be- gins. More information about COSEWIC is available at http://cosewic.ca. The Society for Integrative & Comparative Biology Annual Meeting The Society for Integrative & Comparative Biology Annual Meeting to be held 3—7 January 2020 at the JW Marriott Austin, Austin, Texas. Registration is cur- rently open. More information is available at https:// burkclients.com/sicb/meetings/2020/site/, Science, Practice & Art of Restoring Native Ecosystems Conference The Science, Practice & Art of Restoring Native Eco- systems Conference to be held 17-18 January 2020 at the Kellogg Center, East Lansing, Michigan. Re- Midwest Fish & Wildlife Conference The 80th Midwest Fish & Wildlife Conference to be held 22-29 January 2020 at the BOS Center, Springfield, Illinois. The theme of the conference is: ‘Bringing Science Back to the Forefront of Re- Forests Ontario Annual Conference The annual conference of Forests Ontario to be held 14 February 2020 at the Nottawasaga Inn, Alliston, Ontario. The theme of the conference is: ‘We the gistration is currently open. More information is avail- able at https://conference.stewardshipnetwork.org/. source Management’. Registration is currently open. More information is available at http://www.midwest fw.org/. Forest’. Registration is currently open. More infor- mation is available at https://www.forestsontario.ca/ community/annual-conference/. Society for Range Management’s Annual Meeting, Technical Training and Trade Show The Society for Range Management’s 73rd Annual Meeting, Technical Training and Trade Show to be held 16—20 February 2020 at the Sheraton Denver Downtown Hotel, Denver, Colorado. The theme of Wetland Science Conference The Wetland Science Conference, a program of the Wisconsin Wetlands Association, to be held 18—20 February 2020 at the Osthoff Resort, Elkhart Lake, the conference is: ‘Transformation & Translation’. Registration is currently open. More information is available at http://www.srm2020.org/. Wisconsin. More information is available at https:// conference.wisconsinwetlands.org/. 187 188 THE CANADIAN FIELD-NATURALIST Vol. 133 Sedges of the Northern Forest: downloadable digital resource (pdf) A photographic “atlas” of the sedges of the northern forest was posted online (Version 1.3, 357 pages) in early 2019 by Jerry Jenkins at http://northernforest atlas.org/, The guide contains over 1400 high-reso- lution digital images of 223 Cyperaceae species oc- curring in the northeast (including both forest and wetland species). These are composite macro images that are “stacked” to obtain clarity throughout the en- tire depth of field. The result is a collection of crisp, magnified, almost 3D images that carefully illustrate the distinguishing characters of each species—inflo- rescences, perigynia, achenes, ligules, and even an- thers. The stacked images themselves are a marvel, showing even familiar species anew, with rich detail and subtle beauty. In itself, this free downloadable resource is not a sedge guide, although it accompanies a Cornell publi- cation that has recently become available (also called Sedges of the Northern Forest; 2019, Cornell Uni- versity Press, see review on pp. 172-173 of this issue of The Canadian Field-Naturalist). It is intended to be used together with its accompanying book, or with any of the other excellent field guides and floras re- cently released (e.g., Field Guide to Wisconsin Sedges [Hipp 2008]; Field Manual of Michigan Flora [Voss and Reznicek 2012]; Sedges of Maine [Arsenault et al. 2013]). While it has no keys, schematic diagrams of each genus or section (in the case of Carex) highlight main characters. Helpful field marks and key differ- ences with similar species are described and often il- lustrated. For example, a brilliant comparative page of scaled photos of achenes of the challenging Ovales group is sure to be bookmarked on my copy—think of Peterson’s famous watercolour plate of “Confusing Fall Warblers”, but for Ovales sedges! This is a beautiful, portable, searchable resource. As apdf, it can easily be transferred to a phone or tab- let, and taken into the field. When used with regional floras, it will aid both beginners and advanced profes- sionals in the study of this ecologically important, di- verse, yet often difficult group. Literature Cited Arsenault, M., G.H. Mittlehauser, D. Cameron, A.C. Dibble, A. Haines, S. Rooney, and J.E. Weber. 2013. Sedges of Maine—A Field Guide to Cyperaceae. Uni- versity of Maine Press, Orono, Maine, USA. Hipp, A.L. 2008. Field Guide to Wisconsin Sedges: an In- troduction to the Genus Carex (Cyperaceae). University of Wisconsin Press, Madison, Wisconsin, USA. Voss, E.G., and A.A. Reznicek. 2012. Field Manual of Michigan Flora. University of Michigan Press, Ann Ar- bor, Michigan, USA. HOLiLy BICKERTON Ottawa, ON, Canada TABLE OF CONTENTS (concluded) Volume 133, Number 2 Book Reviews BoTAny: Sedges of the Northern Forest: A Photographic Guide ORNITHOLOGY: Gulls—Ospreys: The Revival of a Global Raptor ZooLoGy: The New Beachcomber’s Guide to the Pacific Northwest—A Field Guide to Marine Life of the Protected Waters of the Salish Sea—A Field Guide to Marine Life of the Outer Coasts of the Salish Sea and Beyond—Bats: An Illustrated Guide to All Species—Mammal Tracks and Sign: A Guide to North American Species. Second Edition—The Rise of Wolf 8: Witnessing the Triumph of Yellowstone’s Un- derdog OTHER: To Speak for the Trees: My Life’s Journey from Ancient Celtic Wisdom to a Healing Vision of the Forest—The Overstory: A Novel NEw TITLES News and Comment Upcoming Meetings and Workshops The Committee on the Status of Endangered Wildlife in Canada—The Society for Integrative & Com- parative Biology Annual Meeting—Science, Practice & Art of Restoring Native Ecosystems Confer- ence—Midwest Fish & Wildlife Conference—Forests Ontario Annual Conference—Society for Range Management’s Annual Meeting, Technical Training and Trade Show—Wetland Science Conference Sedges of the Northern Forest: downloadable digital resource (pdf) Mailing date of the previous issue 133(1): 15 October 2019 2019 172 174 176 181 184 187 188 THE CANADIAN FIELD-NATURALIST Volume 133, Number 2 Roadkill of Eastern Newts (Notophthalmus viridescens) in a protected area in Quebec DavID C. SEBURN, ELENA KREUZBERG, and LEAH VIAU First records of Finescale Dace (Chrosomus neogaeus) in Newfoundland and Labrador, Canada DONALD G. KEEFE, ROBERT C. PERRY, and GREGORY R. MCCRACKEN Albinism in Orange-footed Sea Cucumber (Cucumaria frondosa) in Newfoundland EMALINE M. MONTGOMERY, TIFFANY SMALL, JEAN-FRANCOIS HAMEL, and ANNIE MERCIER Wintercresses (Barbarea W.T. Aiton, Brassicaceae) of the Canadian Maritimes COLIN J. CHAPMAN, C. SEAN BLANEY, and Davip M. MAZEROLLE Summer movements of a radio-tagged Hoary Bat (Lasiurus cinereus) captured in southwestern Ontario DEREK MORNINGSTAR and AL SANDILANDS A reconnaissance survey for Collared Pika (Ochotona collaris) in northern Yukon SYDNEY G. CANNINGS, THOMAS S. JUNG, JEFFREY H. SKEVINGTON, ISABELLE DucLos, and SALEEM DAR Occurrence of the rare marine littoral millipede, Thalassisobates littoralis (Diplopoda: Nemato- somatidae), in Canada DONALD F. MCALPINE A practical technique for preserving specimens of duckmeal, Wolffia (Araceae) DANIEL F. BRUNTON Sighting rates and prey of Minke Whales (Balaenoptera acutorostrata) and other cetaceans off Cormorant Island, British Columbia JARED R. TOWERS, CHRISTIE J. MCMILLAN, and REBECCA S. PIERCEY Use of salmon (Oncorhynchus spp.) by Brown Bears (Ursus arctos) in an Arctic, interior, montane environment MATHEW S. SoruM, KYLE Joy, and MATTHEW D. CAMERON Roman Snail, Helix pomatia (Mollusca: Helicidae), in Canada ROBERT G. ForSYTH and JAMES KAMSTRA First record of Paintedhand Mudbug (Lacunicambarus polychromatus) in Ontario and Canada and the significance of iNaturalist in making new discoveries COLIN D. JONES, MAEL G. GLON, KAREN CEDAR, STEVEN M. PAIERO, PauL D. Pratt, and THomas J. PRENEY Duckling mortality at a river weir STEWART B. Roop and AMBER WILLCOCKS 2019 101 105 113 118 125 130 136 139 144 151 156 160 167 (continued inside back cover) ISSN 0008-3550 The CANADIAN FIELD-NATURALIST A JOURNAL OF FIELD BIOLOGY AND ECOLOGY Promoting the study and conservation of northern biodiversity since 1880 Volume 133, Number3 ¢ July—September 2019 Ottawa Field-Naturalists’ Club Club des naturalistes d’Ottawa The Ottawa Field-Naturalists’ Club FOUNDED 1863 (CURRENT INCORPORATION 1879) Patron Her Excellency the Right Honourable Julie Payette, C.C., C.M.M., C.O.M., C.Q., C.D. Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encour- age investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Honorary Members Ronald E. Bedford Michael D. Cadman J. Bruce Falls Robert E. Lee Allan H. Reddoch Charles D. Bird Paul M. Catling Peter W. Hall John Mcneill Joyce M. Reddoch Fenja Brodo Francis R. Cook Christine Hanrahan Theodore Mosquin Dan Strickland Irwin M. Brodo Bruce Di Labio C. Stuart Houston Robert W. Nero John B. Theberge Daniel F. Brunton Anthony J. Erskine Ross A. Layberry E. Franklin Pope Sheila Thomson 2019 Board of Directors President: Diane Lepage Annie Bélair Edward Farnworth Dwayne Lepitzki Henry Steger 1st Vice-President: Jakob Mueller Fenja Brodo Catherine Hessian —_ Bev McBride Ken Young Recording Secretary: Elizabeth Moore Robert Cermak Anouk Hoedeman Gordon Robertson Eleanor Zurbrigg Treasurer: Ann Mackenzie Owen Clarkin Diane Kitching Jeff Saarela To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2, or e-mail: ofnc@ofnc.ca. For information on club activities, go to www.ofnc.ca. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this jour- nal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Website: www.canadianfieldnaturalist.ca/index.php/cfn Editor-in-Chief: Dr. Dwayne Lepitzki Assistant Editor: Dr. Amanda Martin Copy Editors: Sandra Garland and Dr. John Wilmshurst Layout: Robert Forsyth Book Review Editor: Dr. Barry Cottam Online Journal Manager: Dr. Bill Halliday Subscription Manager: Eleanor Zurbrigg Author Charges Manager: Ken Young Associate Editors: Dr. Ron Brooks Dr. Jennifer R. Foote Dr. Donald F. McAlpine _ Dr. Jeffery M. Saarela Dr. Carolyn Callaghan Dr. Graham Forbes Dr. Garth Mowat David C. Seburn Dr. Paul M. Catling Thomas S. Jung Dr. Marty Obbard Dr. Jeffrey H. Skevington Dr. Frangois Chapleau Chair, Publications Committee: Dr. Jeffery M. Saarela All manuscripts intended for publication—except Book Reviews—should be submitted through the online submission system at the CFN website: http://www.canadianfieldnaturalist.ca/index. php/cfn/user. Click the “New Submission” link on the right side of the webpage and follow the prompts. Authors must register for a cfn account at http://www. canadianfieldnaturalist.ca/index.php/cfn/user/ register in order to submit a manuscript. Please contact the Online Journal Manager (info@canadianfieldnaturalist.ca) if you have any questions or issues with the online submission process. In only rare, exceptional circumstances will submissions other than online be considered and, in these cases, authors must contact the Editor-in-Chief (editor@canadianfieldnaturalist.ca) prior to submission. In- structions for Authors are found at https://www.canadianfieldnaturalist.ca/public/journals/1/CFN-author-instructions20Sept2019._ pdf. Book-review correspondence, including arranging for delivery of review copies of books, should be sent to the Book Review Editor by e-mail: bookrevieweditor@canadianfieldnaturalist.ca. Subscriptions and Membership: Subscription rates for individuals are $40 (online only), $50 (print only), or $60 (print + online). Libraries and other institutions may subscribe for $120 (online only or print only) or $180 (print + online). All foreign print subscrib- ers and members (including USA) must add $10 to cover postage. The Ottawa Field-Naturalists’ Club annual membership fee of $40 (individual), $45 (family), or $20 (student) includes an online subscription to The Canadian Field-Naturalist. Members can receive printed issues of CFN for an additional $30 per volume (four issues). For further details, see http://ofnc.ca/membership-and-donations. The club’s regional journal, 7rail & Landscape, covers the Ottawa District and regional Club events and field trips. It is mailed to all club members. It is available to libraries at $40 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be sent to subscriptions@canadianfieldnaturalist.ca or mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O., Ottawa, ON, K1Z 1A2 Canada. Canada Post Publications Mail Agreement number 40012317. Return postage guaranteed. The Thomas H. Manning fund, a special fund of the OFNC, established in 2000 from the bequest of northern biologist Thomas H. Manning (1911-1998), provides financial assistance for the publication of papers in the CFN by independent (non-institutional) authors, with particular priority given to those addressing Arctic and boreal issues. Qualifying authors should make their application for assistance from the fund at the time of their initial submission. Cover: Rocky Mountainsnail (Oreohelix strigosa), one of the 44 terrestrial gastropod taxa found during extensive surveys in the north- ern Columbia River basin, southcentral British Columbia, 2007-2015. See the article in this issue by Ovaska et al. pages 221— 234. Photo: K. Ovaska, September 2007. The Canadian Field-Naturalist Note First recorded co-occurrence of Valvata lewisi Currier, 1868 and Valvata lewisi ontariensis Baker, 1931 (Gastropoda: Valvatidae) from Alberta, Canada, with notes on morphometric and genetic variability ROBERT P. HINCHLIFFE’”, CHERYL TEBBY', and TYLER P. Coss! ‘Royal Alberta Museum, 9810 — 103a Avenue, Edmonton, Alberta T5J 0G2 Canada “Corresponding author: robert. hinchliffe@gov.ab.ca Hinchliffe, R.P., C. Tebby, and T.P. Cobb. 2019. First recorded co-occurrence of Valvata lewisi Currier, 1868 and Valvata lewisi ontariensis Baker, 1931 (Gastropoda: Valvatidae) from Alberta, Canada, with notes on morphometric and genetic variability. Canadian Field-Naturalist 133(3): 189-192. https://doi.org/10.22621/cfn.v13313.2237 Abstract Sympatric populations of Loosely Coiled Valve Snail (Valvata lewisi ontariensis Baker, 1931) and Fringed Valvata (Valvata lewisi Currier, 1868) are documented from Alberta, Canada, for the first time. Both forms have been identified concurrently in aquatic invertebrate survey samples collected from three wetlands in northeastern Alberta by the Alberta Biodiversity Monitoring Institute. Molecular analysis (DNA barcodes) indicates that there is no genetic distinction between V. lewisi (sensu stricto) and V. lewisi var. ontariensis. Morphometric measurements show that the degree of open coiling, the char- acter that defines V. /ewisi var. ontariensis, is highly variable in Alberta specimens. Our findings confirm that V. /ewisi var. ontariensis 1s a phenotypic morph of V. /ewisi. Key words: Distribution; range extension; Alberta; Valvatidae; Valvata; Valvata lewisi;, Valvata lewisi ontariensis;, Alberta Biodiversity Monitoring Institute; ABMI Loosely Coiled Valve Snail (Valvata lewisi ontari- ensis Baker, 1931; common name from Clarke 1981) is a tiny, enigmatic freshwater gastropod that has rarely been collected since first being described by Frank Collins Baker in 1931. Originally thought to be confined to a few locations in western Ontario, Canada (Baker 1931; Clarke 1973; Figure 1), similar specimens have since been collected as Pleistocene fossils or empty shells in Manitoba, Canada (Clarke 1973), and Minnesota, USA (Bright 1981), with the only other confirmed record of living specimens from the Cottonwood Lake Study area in North Dakota, USA (Hanson ef al. 2002; Figure 1). Here, we report on the first sympatric collections of Valvata lewisi (sensu stricto) and V. lewisi var. ontariensis from Alberta, Canada, and provide notes on morphometric and genetic variability. In contrast to the distribution of V. /ewisi var. on- tariensis, Valvata lewisi Currier, 1868 (Fringed Val- vata, according to Turgeon et al. 1998) is commonly found across the prairie, parkland, and boreal regions of Canada from Newfoundland to British Columbia, north into the Yukon and Alaska, and south into the northern United States (Clarke 1981; NatureServe 2017; Figure 1). Valvata lewisi is a small freshwater snail that seldom exceeds 5 mm in diameter and has a depressed spire, multi-spiral operculum, and bi-pec- tinate gill (Clarke 1973; Burch 1982). Shell sculpting consists of fine striations on the first one and a half to two whorls, which develop on subsequent whorls into axial lamellae that are usually elevated and blade- like, but may be reduced to coarse collabral threads (Clarke 1973). In comparison, V. /ewisi var. ontari- ensis exhibits the same characteristics, but, unlike V. lewisi (sens. str.) where the body whorl directly con- tacts the preceding whorl, V. /ewisi var. ontariensis exhibits open coiling in which the last one to one and a half whorls are separated (Baker 1931; Figure 2). Valvata lewisi var. ontariensis has been detected at five wetlands in Alberta, Canada, through the on- going activities of the Alberta Biodiversity Mon- itoring Institute (ABMI). The ABMI collects bio- logical information on a wide range of terrestrial and aquatic organisms across the province using 189 ©The Ottawa Field-Naturalists’ Club 190 THE CANADIAN FIELD-NATURALIST Vol. 133 W302 (2016) W390 (2007, 2015)@ Fort McMurray e W633 (2015) OGW-732-1 (2013, 2015) @ Ficure I. a. Alberta Biodiversity Monitoring Institute wetland site locations where specimens of Valvata lewisi vat. ontari- ensis have been documented in Alberta (large solid circles, collection years in parentheses). Inset map b. shows known range of Valvata lewisi (sensu stricto) in North America (shaded area) with historical collection records of V. /ewisi var. ontariensis (solid circles). standardized, publicly available methods (e.g., ABMI 2015, 2018). During routine taxonomic analysis of aquatic invertebrate samples collected by the ABMI in 2007, we detected 12 V. lewisi var. ontarien- sis specimens in samples obtained from a perma- nent wetland in the northeastern part of the prov- ince (ABMI site W390; 57.26899°N, 110.72157°W; Figure 1). Following this initial detection, several additional V. /ewisi var. ontariensis specimens were subsequently recovered from ABMI samples col- lected from the initial detection site and four addition- al wetlands in the same region—W152 (58.78107°N, 110.86238°W), W302 (57.80382°N, 110.65305°W), W633 (55.97079°N, 112.23922°W), and OGW-732-1 (55.25222°N, 110.91161°W; Figure 1)—for a total col- lection of 75 specimens. This sampling effort also 2019 HINCHLIFFE ET AL.: VALVATA LEWISI VAR. ONTARIENSIS IN ALBERTA 19] FiGurE 2. Dorsal and ventral views of Valvata lewisi (top) and Valvata lewisi var. ontariensis (bottom). Photos: Robert P. Hinchliffe. revealed the co-occurrence of V. /ewisi var. ontarien- sis and V. lewisi (sens. str.) at sites W390, W633, and OGW-732-1. Voucher specimens have been preserved in 80% ethanol and deposited in the invertebrate zo- ology collection at the Royal Alberta Museum in Edmonton, Alberta, Canada (ABMI.A.91, ABMI. A.5396, ABMI.A.11900, ABMI.A.15382, ABMI.A. 18222, and ABMI.A.30673). Morphometric analyses of V. /ewisi var. ontarien- sis revealed considerable variation in the degree of open coiling in Alberta specimens. Measurements showed a clear gradation in the ratio of open-coiled gap to aperture diameter in specimens from both W390 (0.04:1 to 0.22:1, mean 0.10:1, 7 = 16) and OGW- 732-1 (0.02:1 to 0.41:1, mean 0.13:1, 2 = 18), a pattern also noted, although not directly measured, by Baker (1931) when examining western Ontario specimens. Our examination of intra- and inter-specific vari- ability associated with the DNA barcode markers cytochrome c oxidase 1 (COl1) and internal tran- scribed spacer 2 (ITS2) for V. lewisi (sens. str.) and V. lewisi var. ontariensis specimens from sites W390 and OGW-732-1 indicated no genetic distinction between the two morphs. For CO1, the mean interspecific vari- ation (+ SD) was 0.17% + 0.13 with intraspecific gen- etic distances at 0.16% + 0.12 for V. lewisi (sens. str.; n = 12) and 0.16% + 0.14 for V. lewisi var. ontarien- sis (n = 28). For ITS2, the mean interspecific varia- tion ( SD) was 0.07% + 0.14 and the intraspecific genetic distance of V. /ewisi (sens. str., n = 10) was 0.03% + 0.07 and of V. /ewisi var. ontariensis (n = 23) was 0.11% + 0.17. DNA barcoding was conducted by the Canadian Centre for DNA Barcoding in Guelph, Ontario, Canada. Genetic sequences have been sub- mitted to GenBank (CO1l: MK721872 to MK721913, ITS2: MK721934 to MK721969). Our detections of V. /ewisi var. ontariensis consti- tute the first record of this morph in Alberta, Canada, and the first explicitly documented instances of co- occurrence of V. /ewisi var. ontariensis with V. lewisi (sens. str.). Clarke (1973: 229) noted that loosely coiled specimens seemed to “occur also (rarely) in some apparently normal populations”, but neglected to provide references or observational evidence for this statement. Furthermore, he suggested that the specimens he examined were uniform as all one morph or the other. Other published reports on this species do not make any mention of co-occurrence of the two morphs. The taxonomic status of V. /ewisi var. ontarien- sis aS a valid subspecies has historically been uncer- tain (see Baker 1931; Clarke 1973, 1981; Burch 1982). 192 The working definition of a subspecies is two or more populations of the same species from separate geo- graphic locations with one or more distinguishing characters (Mayr 1942, 1982). The initial Ontario col- lections identified by Baker (1931) and later expanded on by Clarke (1973) were originally thought to be an isolated and distinct population of the open-coiled morph. Our concurrent collections of V. /ewisi (sens. str.) and V. lewisi var. ontariensis clearly show that the two morphs can occur in the same water body. This, in addition to the lack of a CO1 or ITS2 barcode gap between the two morphs, supports the conclu- sion that V. /ewisi ontariensis is a phenotypic morph of V. lewisi. Despite the broad and common distribution of V. lewisi (sens. str.), the open-coiled morph has, thus far, been collected in only three isolated regions of North America. Given the widely spaced and seem- ingly isolated locations where V. /ewisi var. ontari- ensis has been collected, it is possible that the open- coiled morph is the result of some unknown and possibly localized environmental factor. However, it is also possible that V. /ewisi var. ontariensis is more common than collection records indicate and is sim- ply difficult to detect during routine aquatic inverte- brate surveys because of its small size and propen- sity to burrow into the upper layer of soft substrates (R.P.H. pers. obs.). Targetted sampling in other re- gions is needed to more fully understand the com- plete distribution of V. /ewisi var. ontariensis. We recommend that future collections of open-coiled V. lewisi specimens be identified and labeled as V. lewisi var. ontariensis to allow for better tracking of the localities where this morph occurs and per- haps yield additional clues as to the possible source of open coiling. Author Contributions Writing — Original draft: R.P.H.; Writing —-Review & editing: R.PH., C.T, and T.P.C.; Visualization: R.P.H.; Conceptualization: R.P.H. and T.P.C.; Investi- gation: R.P.H. and C.T.; Resources: C.T.; Formal an- alysis: R.P.H. Acknowledgements We thank Dr. Matthias Buck at the Royal Alberta Museum for assistance with the molecular analy- sis and valuable editorial suggestions on earlier ver- sions of the manuscript. We also thank the Alberta THE CANADIAN FIELD-NATURALIST Vol. 133 Biodiversity Monitoring Institute for providing additional data. This research was supported by the Alberta Biodiversity Monitoring Institute and the Royal Alberta Museum. Literature Cited ABMI (Alberta Biodiversity Monitoring Institute). 2015. Processing aquatic invertebrates (10017) version 2015-07-23. ABMI, Edmonton, Alberta, Canada. Ac- cessed 12 November 2018. https://www.abmi.ca/home/ publications/351-400/396. ABMI (Alberta Biodiversity Monitoring Institute). 2018. Wetland field data collection protocols (abridged version) 2018-05-07. ABMI, Edmonton, Alberta, Can- ada. Accessed 12 November 2018. https://www.abmi.ca/ home/publications/501-550/510. Baker, F.C. 1931. Description of a new variety of Valvata /ewisi Currier. Nautilus 44: 119-121. Bright, R.C. 1981. A new record of Valvata sincera ontari- ensis F.C. Baker from Minnesota. Nautilus 95: 20. Burch, J.B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. United States Environmental Pro- tection Agency, Washington, DC, USA. EPA/600/3-82- 026 (NTIS PB82207168). Accessed 25 May 2018. https:// cfpub.epa.gov/si/si_public_record_Report.cfm?Lab= ORD&dirEntryID=32218. Clarke, A.H. 1973. The freshwater molluscs of the Cana- dian Interior Basin. Malacologia 13: 1-509. Clarke, A.H. 1981. The Freshwater Molluscs of Canada. National Museum of Natural Science, Ottawa, Ontario, Canada. Hanson, B.A., N.H. Euliss, Jr., and D.M. Mushet. 2002. First records of loosely coiled valve snail in North Dakota. Prairie Naturalist 34: 63—65. Mayr, E. 1942. Systematics and the Origin of Species, from the Viewpoint of a Zoologist. Columbia University Press, New York, New York, USA. Mayr, E. 1982. Of what use are subspecies? Commentary. Auk 99: 593-595. NatureServe. 2017. Valvata species report. Accessed 27 April 2017. http://explorer.natureserve.org/servlet/Nature Serve?searchSciOrCommonName=Valvata. Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P. Mikkelsen, R.J. Neves, C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. Common and Scientific Names of Aquatic Inver- tebrates from the United States and Canada: Mollusks. Second Edition. Special Publication 26. American Fish- eries Society, Bethesda, Maryland. Received 2 April 2019 Accepted 24 December 2019 The Canadian Field-Naturalist Note Spotless burnsi pattern in Northern Leopard Frog (Lithobates pipiens) in Maine Scott B. LINDEMANN"*, Davip E. PUTNAM’, MALCOLM L. HUNTER, JR.', and TREVOR B. PERSONS? 'Department of Wildlife, Fisheries, and Conservation Biology, University of Maine, 5755 Nutting Hall, Room 210, Orono, Maine 04469 USA "University of Maine at Presque Isle, 214 South Hall, Presque Isle, Maine 04769 USA 3206 Bigelow Hill Road, Norridgewock, Maine 04957 USA “Corresponding author: lindemann.scott@gmail.com Lindeman, S.B., D.E. Putnam, M.L. Hunter, Jr., and T.B. Persons. 2019. Spotless burnsi pattern in Northern Leopard Frog (Lithobates pipiens) in Maine. Canadian Field-Naturalist 133(3): 193-195. https://doi.org/10.22621/cfn.v13313.2283 Abstract We document the spotless “burnsi” morph in Northern Leopard Frog (Lithobates pipiens) in Maine. Key words: Northern Leopard Frog; Lithobates pipiens, amphibian; pattern variant; Maine The “burnsi” mutation in Northern Leopard Frog (Lithobates pipiens) results in loss of the frog’s char- acteristic spots from the back, and sometimes also from the dorsal surface of the legs (McKinnell et al. 2005). Herpetologists have studied this muta- tion since the early 20th century, and Moore (1942) demonstrated that the burnsi mutation allele is dom- inant over the wild-type allele. More recently, this mutation has been used to study the effects of gen- etic bottlenecks in this species of conservation con- cern (McKinnell et a/. 2005). This mutation is re- ported most frequently in central Minnesota and the surrounding area, where it occurs in 4.0-7.1% of L. pipiens, although it has also been documented rarely outside this region (Merrell 1965; Brown and Funk 1977; McKinnell et al. 2005; Rogers and Peacock 2012). On 3 June 2018, S.B.L. and D.E.P. discovered a burnsi-type L. pipiens (Figure 1) along the bank of the north branch of Presque Isle Stream (46.641949°N, 68.177440°W) on Scopan Maine Public Reserved Land, T1l1 R4 WELS township, Aroostook County, Maine, USA. The frog was sitting in grass along a stream channel lined with Speckled Alder (A/nus in- cana subsp. rugosa (Du Roi) R.T. Clausen), which was further surrounded by scrub-shrub wetland and mixed coniferous—deciduous forest. The frog was identified as L. pipiens by the gold colouration of the dorsolateral fold, lack of colouration on the groin, white venter, and green dorsum. Four wild-type con- specifics were also found at the same site (Figure 1). Burnsi-type L. pipiens have been collected from only one other locality in Maine, as determined from a review of the Maine Amphibian and Reptile Atlas Project (MARAP 2019) database, which is main- tained by the Maine Department of Inland Fisheries and Wildlife. MARAP contains specimen records from most major North American herpetology collec- tions, as well as most smaller regional ones. In addi- tion, MARAP includes observations from the cit- izen science iNaturalist Web site (www. inaturalist. org). Four specimens at the American Museum of Natural History (AMNH 51343—6) were collected in 1940 in Woodland (i.e., Baileyville, located 175 km south-southeast of the June 2018 collection), Washing- ton County; these were briefly noted by Merrell (1965), but he did not provide catalog numbers or specific locality data. In addition, the Museum of Comparative Zoology houses a series of specimens (MCZ 25541—50) collected on the same date and from the same locality as the AMNH specimens, and the MCZ catalog ledger notes: “Of the 36,000 frogs col- lected in three seasons, about 4% were unspotted, but in other respects wholly typical like the true pipiens occurring at the spot, intergrades between them were present also”. 193 ©The Ottawa Field-Naturalists’ Club 194 ey + THE CANADIAN FIELD-NATURALIST Vol. 133 FicureE 1. Burnsi-type (a) and wild-type (b) Northern Leopard Frogs (Lithobates pipiens) from Presque Isle Stream, Aroostook County, Maine, USA. Photos: Scott B. Lindemann. Some of the specimens at both AMNH and MCZ retain spotting on the hind limbs similar to ours, while others are completely unspotted. Assuming the MCZ catalog ledger note is accurate, the burnsi mutation was apparently, at least at this location in Maine, as common as reported in central Minnesota and sur- rounding areas. We have not surveyed the Woodland area for Leopard Frogs, and the MCZ catalog ledger also states that “The pond has since been destroyed by peat cutting”. Aside from this series of specimens, the Presque Isle Stream individual is the only example of unspotted L. pipiens we are aware of from Maine. Acknowledgements We thank David Dickey and David Kizirian (American Museum of Natural History) and Jose Rosado (Museum of Comparative Zoology) for pro- viding photographs and information on specimens in their care. Our appreciation is extended to Malcolm Hunter, Aram Calhoun, and Kristine Hoffmann of the University of Maine and to Phillip deMayna- dier, Derek Yorks, and Jason Czapiga of the Maine Department of Inland Fisheries and Wildlife for pro- ject guidance. Funding support was provided by the Maine Department of Inland Fisheries and Wildlife (Endangered and Nongame Wildlife Fund and United States Fish and Wildlife Service State Wildlife Grant), the Department of Wildlife, Fisheries, and Conservation Biology at the University of Maine, Eastern Maine Conservation Initiative, and the University of Maine Graduate Student Government. Fieldwork was performed under a Maine Department of Inland Fisheries and Wildlife scientific collec- tion permit (#2018-184) issued to S.B.L. Animal use protocols were approved by the Institutional Animal Care and Use Committee of the University of Maine (#A2018-03-01). Publication #3695 of the Maine Agricultural and Forest Experiment Station. This project was supported by the United States Department of Agriculture, National Institute of Food and Agriculture, Hatch Project Number MEO- 21705 through the Maine Agricultural and Forest Experiment Station. 2019 LINDEMANN ET AL.: SPOTLESS BURNSI NORTHERN LEOPARD FROG 195 Literature Cited Merrell, D.J. 1965. The distribution of the dominant burnsi Brown, L.E., and R.S. Funk. 1977. Absence of dorsal spot- gene in the leopard frog, Rana pipiens. Evolution 19: ting in two species of leopard frogs (Anura: Ranidae). 69-85. https://doi.org/10.2307/2406296 Herpetologica 33: 290-293. Moore, J.A. 1942. An embryological and genetical study of MARAP (Maine Amphibian and Reptile Atlas Project). Rana burnsi Weed. Genetics 27: 408-416. 2019. Data files. Maine Department of Inland Fisheries Rogers, S.D., and M.M. Peacock. 2012. The disappearing and Wildlife, Augusta, Maine, USA. Northern Leopard Frog (Lithobates pipiens): conserva- McKinnell, R.G., D.M. Hoppe, and B.K. McKinnell. tion genetics and implications for remnant populations 2005. Monitoring pigment pattern morphs of Northern in western Nevada. Ecology and Evolution 2: 2040-— Leopard Frogs. Pages 328-337 in Amphibian Declines. 2056. https://doi.org/10.1002/ece3.308 Edited by M. Lannoo. University of California Press, Berkeley and Los Angeles, California, USA. https://doi. | Received 16 May 2019 org/10.1525/california/9780520235922.001.0001 Accepted 19 December 2019 The Canadian Field-Naturalist Note Axanthism in Green Frogs (Lithobates clamitans) and an American Bullfrog (Lithobates catesbeianus) in Maine Scott B. LINDEMANN’, AIDAN M. O’ BRIEN!, TREVOR B. PERSONS’, and PHILLIP G. DEMAYNADIER? 'Department of Wildlife, Fisheries, and Conservation Biology, University of Maine, 5755 Nutting Hall, Room 210, Orono, Maine 04469 USA 7206 Bigelow Hill Road, Norridgewock, Maine 04957 USA 3Maine Department of Inland Fisheries and Wildlife, 650 State Street, Bangor, Maine 04401 USA *Corresponding author: lindemann.scott@gmail.com Lindemann, S.B., A.M. O’Brien, T.B. Persons, and P.G. DeMaynadier. 2019. Axanthism in Green Frogs (Lithobates clam- itans) and an American Bullfrog (Lithobates catesbeianus) in Maine. Canadian Field-Naturalist 133(3): 196-198. https://doi.org/10.22621/cfn.v13313.2285 Abstract We document eight cases of axanthism in Green Frogs (Lithobates clamitans) and one case in an American Bullfrog (Lithobates catesbeianus) in Maine. Although this mutation has been previously reported for both species, this 1s the first confirmed documentation of “blue” L. clamitans and L. catesbeianus from Maine. Key words: Green Frog; Lithobates clamitans; American Bullfrog; Lithobates catesbeianus; amphibian; blue colour vari- ant; axanthism; Maine Although “blue” frogs have been documented since 1885 (Haller 1885; also cited in Berns and Uhler 1966), Jablonski et al. (2014) note that axanthism is one of the least known colour aberrations in anur- ans. Axanthism results from the absence or altera- tion of xanthophores, the dermal chromatophores responsible for red and yellow pigmentation (Berns and Narayen 1970). Normally, these xanthophores contain pteridines and carotinoids, which cause the underlying blue iridophores to appear green; in their absence, the skin appears blue (Berns and Narayan 1970). Berns and Uhler (1966) noted that blue Green Frogs (Lithobates clamitans) and Northern Leopard Frogs (Lithobates pipiens) have been recorded from northeastern United States (mentioning Maine spe- cifically) and southeastern Canada, although they did not state which species were found in which state or province. Of 111 blue frogs they examined from throughout eastern North America, 100 were L. clamitans, 10 were L. pipiens, and one was an American Bullfrog (Lithobates catesbeianus, from Kentucky). Dodd (2013) cited reports of blue L. clam- itans from Massachusetts, Delaware, and Virginia, but not Maine. Desroches and Rodrigue (2004) illus- trated a blue L. clamitans, although not stated, it was presumably from Quebec. Dodd (2013) included a photograph of a blue L. catesbeianus, but did not give its locality, and Gilhen and Russell (2015) reported three blue L. catesbeianus from Nova Scotia. On 9 June 2018, S.B.L., James A. Elliott, and A.M.O. found an axanthic adult male L. clamitans (Figure 1) in a small pool with emergent vegetation in an ~10-year-old clearcut in coniferous forest, TS R11 WELS township, Piscataquis County, Maine, USA (46.116659°N, 69.211416°W). Roughly 20 addi- tional wild-type conspecifics were also found at the same location. Axanthic L. clamitans (all adults) have also been photo-documented from the follow- ing localities in Maine: Washington, Knox County, 11 August 2010; Wiscasset, Lincoln County, 4 October 2013; Buxton, York County, mid-June 2017; Phillips, Franklin County, 16 August 2017; Raymond, Cumberland County, 16 September 2017; and Bethel, Oxford County, 15 July 2018 (MARAP 2019). Although these were coloured similarly to the indi- vidual in Figure | (1.e., metallic greenish-blue over the entire dorsum), an additional one, from Hurd’s Pond, Swanville, Waldo County (44.476658°N, 196 ©The Ottawa Field-Naturalists’ Club 2019 @ o ; ~~ — a eo. 7 — asd Figure 1. Axanthic adult male Green Frog (Lithobates cla- mitans) from T5 R11 WELS township, Piscataquis County, Maine, USA, 9 June 2018. Photo: Scott B. Lindemann. 69.032297°W ) was piebald blue-green (Figure 2). In contrast to L. clamitans, only a single axan- thic L. catesbeianus has been documented from Maine (MARAP 2019). A subadult (or small adult) female (Figure 3) was photographed at Headquarters Pond, Cobscook Bay State Park, Edmunds Township, LINDEMANN ET AL.: AXANTHISM IN GREEN FROGS AND BULLFROGS IN MAINE 197 Washington County (44.849505°N, 67.167045°W ) by Owen and Raymond Brown on 19 June 2011. To our knowledge, these represent the first con- firmed records of axanthic L. clamitans and L. cates- beianus from Maine. We do not know the true in- cidence of axanthic frogs in Maine. The MARAP database contains 773 records of L. clamitans and 445 of L. catesbeianus, but these records are not the result of systematic surveys, and the resulting ratios of axanthic frogs to normally pigmented ones (1:97 for L. clamitans and 1:445 for L. catesbeianus) are undoubtedly overestimates, as axanthic individuals are much more likely to be reported. Based on data from midwestern supply houses, Berns and Uhler (1966) estimated that out of a sample of roughly two million frogs, axanthics (mostly L. clamitans) oc- curred at a frequency of about 1:29 000, although in some local areas the rate was as high as 1:318 (22 of 7000). Our data support Berns and Uhler’s (1966) finding that axanthism appears to be most common in L. clamitans. Like albinism, axanthism is presum- ably a heritable trait (Bechtel 1995). Whether there is a potential selective advantage or disadvantage to axanthism is unknown, but its rarity suggests that it is likely neutral or even potentially disadvantageous. Further documentation of axanthic specimens, such as those reported here, is conducive to an improved understanding of taxonomic and geographic patterns in this interesting colour aberration. FiGuRE 2. Piebald axanthic adult male Green Frog (Lithobates clamitans) from Swanville, Waldo County, Maine, USA, 27 June 2016. Photo: Trevor B. Persons. 198 THE CANADIAN FIELD-NATURALIST Vol. 133 FicgureE 3. Axanthic American Bullfrog (Lithobates catesbeianus) from Cobscook Bay State Park, Edmunds Township, Washington County, Maine, USA, 19 June 2011. Photo: Raymond Brown. Acknowledgements Our appreciation is extended to Malcolm Hunter, Aram Calhoun, and Kristine Hoffmann, of the Uni- versity of Maine, and Derek Yorks and Jason Czapiga, of the Maine Department of Inland Fisheries and Wildlife, for project guidance. Funding support was provided by the Maine Department of Inland Fisheries and Wildlife (Endangered and Nongame Wildlife Fund), the United States Fish and Wildlife Service (State Wildlife Grant), the Department of Wildlife, Fisheries, and Conservation Biology at the University of Maine, Eastern Maine Conservation Initiative, and the University of Maine Graduate Stu- dent Government. Fieldwork was performed under a Maine Department of Inland Fisheries and Wildlife scientific collection permit (#2018-184) issued to S.B.L. Animal use protocols were approved by the Institutional Animal Care and Use Committee of the University of Maine (#A2018-03-01). Literature Cited Bechtel, H.B. 1995. Reptile and Amphibian Variants: Colors, Patterns, and Scales. Krieger Publishing, Malabar, Flor- ida, USA. Berns, M.W., and K.S. Narayan. 1970. An histochem- ical and ultrastructural analysis of the dermal chro- matophores of the variant ranid blue frog. Journal of Morphology 132: 169-180. https://doi.org/10.1002/jmor. 1051320205 Berns, M.W., and L.D. Uhler. 1966. Blue frogs of the genus Rana. Herpetologica 22: 181-183. Desroches, J., et D. Rodrigue. 2004. Amphibiens et rep- tiles du Québec et des Maritimes. Editions Michel Quintin, Waterloo, Quebec, Canada. Dodd, Jr., C.K. 2013. Frogs of the United States and Canada, Volume 2. Johns Hopkins University Press, Baltimore, Maryland, USA. https://doi.org/10.1353/boo0k.25108 Gilhen, J., and R.W. Russell. 2015. Three records of rare blue American Bullfrogs, Lithobates catesbeianus, in Nova Scotia, Canada. Canadian Field-Naturalist 129: 395-398. https://doi.org/10.22621/cfn.v12914.1762 Haller, B. 1885. Uber das blaue Hocheitskleid des Gras- frosches. Zoologischer Anzeiger 8: 611—617. Jablonski, D., A. Alena, P. Vicek, and D. Jandzik. 2014. Axanthism in amphibians: a review and the first record in the widespread toad of the Bufotes viridis complex (Anura: Bufonidae). Belgian Journal of Zoology 144: 93-101. MARAP (Maine Amphibian and Reptile Atlas Project). 2019. Data files. Maine Department of Inland Fisheries and Wildlife, Augusta, Maine, USA. Received 15 May 2019 Accepted 19 December 2019 The Canadian Field-Naturalist Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt new to Atlantic Canada SEAN R. HAUGHIAN!:* and THOMAS H. Netty? ‘Botany & Mycology Section, Department of Natural Sciences, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada *Biology Department, Saint Mary’s University, 923 Robie Street, Halifax, Nova Scotia B3H 3C3 Canada 3Mersey Tobeatic Research Institute, 9 Mount Merritt Road, P.O. Box 215, Kempt, Queens County, Nova Scotia BOT 1BO Canada *Corresponding author: sean.haughian@smu.ca Haughian, S.R., and T.H. Neily. 2019. Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt new to Atlantic Canada. Canadian Field-Naturalist 133(3): 199-205. https://doi.org/10.22621/cfn.v13313.2052 Abstract Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt is reported for the first time in Atlantic Canada. It was found on the base of a large Eastern White Cedar (Thuja occidentalis) in a swamp in Nova Scotia. The specimen was examined using light microscopy, diagnosed using standard keys, and compared with reference specimens, including two European collections from the New Brunswick Museum, two North American collections annotated by R.M. Schuster, and the only material that may have been previously collected in Canada, by T. Drummond. We speculate on the original loca- tion of Drummond’s collection, and the implications of this finding for conservation. Key words: Liverwort; Nova Scotia; hepatic; Lejeuneaceae Introduction Harpalejeunea molleri (Stephani) Grolle (Lejeune- aceae) is a rare leafy liverwort (Note: liverworts typically do not have common names) with a dis- junct global distribution, primarily around the North Atlantic, with European and North American populations recognized as subspecies. In Europe, Harpalejeunea molleri subsp. molleri has been col- lected on the west coast of Norway and in the United Kingdom, Ireland, and Spain (GBIF 2018), as well as Finland, Italy, Madeira, the Azores, the Canary Islands, and Corsica (Hodgetts 2015). The North American subspecies, Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt, is known pri- marily from the Appalachian Mountain Range and Atlantic Coastal Plain in the southeastern United States, where it has been collected in Alabama, Georgia, Kentucky, and North and South Carolina (Schuster 1980; Consortium of North American Bryo- phyte Herbaria 2017), as well as Florida, Mississippi, Tennessee, and Virginia (Breil 1970). A single speci- men is thought to have been collected from Canada by Thomas Drummond in the early 19th century, but the collection location is ambiguous, and no other specimens are known to have been collected in Canada since then. Two recent collections are also re- ported from Brazil, without subspecific designation (GBIF 2018). The correct name for H. molleri and its infraspecific taxa has historically been a source of confusion. Schu- ster (1980) used the name Harpalejeunea ovata (Dickson) Schiffner, and, consequently, much of the material in North American herbaria has been ac- cessioned under that name. However, Grolle (1989) demonstrated that this name is a synonym of Douinia ovata (Dickson) H. Buch (Scapaniaceae) and that H. molleri is the correct name for the taxon, as recog- nized recently by European authorities (Paton 1999; Damsholt and Pagh 2002). Nevertheless, the former taxonomic confusion continues to impede accurate delineation of the species’ distribution because many herbarium records have not been revised to reflect current taxonomy. In North America, H. molleri subsp. integra has been found in old growth swamps or riparian areas with relatively open forest canopies, most commonly as an epiphyte on the base of hardwood trees (Breil 1970; Schuster 1980) and in crevices on sediment- ary rock (Consortium of North American Bryophyte Herbaria 2017). It is often in mixed species col- onies (Breil 1970), and common liverwort associ- ates in herbarium records include Frullania asagray- A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 199 ©The Ottawa Field-Naturalists’ Club 200 ana Montagneagne, Lejeunea lamacerina (Stephani) Schiffner, Lejeunea ruthii (A. Evans) R.M. Schuster, Lejeunea ulicina (Taylor) Gottsche, Lindenberg & Nees, and Radula obconica Sullivant (Consortium of North American Bryophyte Herbaria 2017). Methods The collection site was a mixedwood swamp near Hectanooga, Digby County, Nova Scotia (~44.082°N, 66.056°W). Geologically, this part of Digby County is underlain by the Church Point Formation, which is composed primarily of grey to green, fine- to medium-grained metasiltstone and metasandstone, with rare shale deposits (White and Horne 2012). Soils are stony in places and poorly drained, being a mix of peat, sandy loam, and loam-till derived from slate (Hilchey et al. 1962). The habitat is a rich swamp forest, dominated by Eastern White Cedar (Thuja oc- cidentalis L.), Red Maple (Acer rubrum L.), Balsam Fir (Abies balsamea (L.) Miller), and Yellow Birch (Betula alleghaniensis Britton). Hummock and hol- low microtopography characterizes the ground layer, with hummocks dominated by sedges (Carex spp.) and Cinnamon Fern (Osmundastrum cinnamomeum (L.) C. Presl) and hollows dominated by Sphagnum spp. and standing water. The shrub layer is patchy, with Common Winterberry (//ex verticillata (L.) A. Gray) and Grey Alder (A/nus incana (L.) Moench). Collections of H. molleri subsp. integra were made opportunistically during searches for Frullania selwyniana Pearson, also rare in the province. Approximately 40 mature cedar trees were visually inspected during this search; eight were found to host visible mixed-species colonies of leafy liverworts. A mixed-species collection was made from each of the eight host trees and later examined using stereo- microscopy. Liverworts were identified using stan- dard taxonomic keys (Schuster 1980; Paton 1999), Two of these collections were found to contain H. molleri subsp. integra. The larger of the two col- lections was then compared with reference material from three sources: (1) two exsiccatae from the herb- arium at the New Brunswick Museum (NBM), (2) two recent collections from the United States that were annotated by liverwort authority R.M. Schuster at the Field Museum (F), and (3) the original (sup- posedly) Canadian collection by T. Drummond, held by the New York Botanical Garden. Neither S.R.H. nor T.H.N. has since had the opportunity to return to this location to assess the population size or health of the colony. In this paper, we provide a brief description of the morphology of the specimen that was deposited at the NBM as evidence for our subspecific designation. THE CANADIAN FIELD-NATURALIST Vol. 133 We also highlight noteworthy aspects of the historical collections for the sake of comparison. Results Harpalejeunea molleri subsp. integra was col- lected from two cedar trees in the Hectanooga Cedar Swamp. One of these collections was accessioned at the NBM, while the other is held in the private herb- arium of T.H.N. Common species in these colonies included Frullania asagrayana Montagne, Frullania oakesiana Austin, Ptilidium pulcherrimum (Weber) Vainio, and Radula complanata (L.) Dumortier, while rarer species included F! se/wyniana, Lejeunea cavifolia (Ehrhart) Lindberg, and Lejeunea ulicina (Taylor) Gottsche, Lindenberg & Nees. Both of the (mixed-species) colonies, in which H. molleri subsp. integra was detected, were ~40 cm? on the bases of large Eastern White Cedars (~25 cm diameter at breast height). Harpalejeunea molleri subsp. integra occupied only a small fraction (<<10%) of the colonies and the subsequent collected material, but was dis- tinct from the other species present, being obviously greener than F. selwyniana, larger than L. ulicina, and with more acutely angled leaves than L. cavifolia. Shoot and colony architecture of H. molleri subsp. integra in the collected material corresponded to a previously published description (Schuster 1980). The collective, multi-species colony structure for our sample was that of a loose “smooth mat”, al- though each individual species exhibited a thread- like growth form (sensu Bates 1998). Shoots of H. molleri subsp. integra were 0.4—0.6 mm wide (trans- verse axis, including leaves) and displayed a dichot- omous irregular lateral branching pattern. Stem postical cortical cells were 13-15 um wide on ma- ture shoots. Leaves were two-ranked, spreading, and complicate-bilobed with alternate insertions along the stem (Figure 1). Antical leaf lobes were comma shaped and longer than broad (1.1—1.2 length to width ratio); proximal margins overlapped the stem above the transverse insertion (Figure 2), and distal margins were acute tipped, typically tapering to a single cell, or occasionally two cells and often curved toward the substrate (Figure 3). The smaller, postical lobe (lob- ule) attached to the stem along the entire length of its proximal margin and folded under the larger, antical lobes, forming a rounded keel along the anterior leaf margin (Figure 3); the angle between the distal edge of the keel and the free antical lobe ranged from 90° to 120°, and the joint was often strongly indented (Figure 2). The distal tips of most lobules bore a slightly elongated, tooth-like cell, located proximal to the distal margin of the keel; this cell projected away from the stem and was ~1.5—2 times the length of a median lobule cell (Figure 3). Immediately proximal 2019 Figure 1. Postical view of Harpalejeunea molleri subsp. integra shoot (Neily 1629, New Brunswick Museum). Photo: Sean Haughian. FiGure 2. Antical view of Harpalejeunea molleri subsp. integra shoot, from newly collected material (Neily 1629, New Brunswick Museum). Photo: Sean Haughian. to this tooth-like cell, some lobules also had a clavate, hyaline papilla (not shown). Underleaves were 0.12— 0.16 mm across, shallowly bilobed, and widely diver- gent; each lobe was four cells wide at the base and rounded at the apex (Figure 4). The specimen had no obvious reproductive structures. The two collections from North America (F) were consistent with Schuster’s (1980) descriptions of H. molleri subsp. integra. They exhibited stem postical cortical cells 13-19 um in width, bilobed underleaves with four cells at the base of each lobe, and strongly indented leaf margins where the distal terminus of the lobule’s keel attached to the antical leaf lobe. The two collections from Spain and Portugal (NBM) had characters consistent with Schuster’s (1980) and Paton’s (1999) descriptions of H. molleri subsp. molleri. Compared with the USA material, they had consistently wider postical cortical cells of 19-23 um, more weakly indented joints (forming angles of ~90—135°) between the lobule and leaf lobe, and slightly more variable underleaf lobe widths (4—7 cells). Drummond’s collection was somewhat transi- tional between the European and the North American collections examined; the leaf lobe—lobule joints were HAUGHIAN AND NEILY: LIVERWORT NEW TO ATLANTIC CANADA 201 FiGure 3. Postical view of Harpalejeunea molleri subsp. integra shoot, showing lobules, underleaves, and antical lobe tips, from newly collected material (Neily 1629, New Brunswick Museum). Photo: Sean Haughian. Figure 4. Postical view of Harpalejeunea molleri subsp. integra shoot, showing underleaves and cortical stem cells, from newly collected material (Nei/y 1629, New Brunswick Museum). Photo: Sean Haughian. strongly indented on mature stems, and the under- leaf lobes were mostly 4 (-6) cells across. However, the postical cortical cells of the stem were wider (19— 24 um) than is typical for H. molleri subsp. integra. Associated taxa in this packet included Diplophyllum albicans (L.) Dumortier and Frullania tamarisci (L.) Dumortier. The only writing on the packet was the former Latin name of the species (“Lejeunea ovata’) and the vague place-name, “British North America”. 202 Discussion This is the first report of H. molleri subsp. inte- gra in Atlantic Canada, and the first reliable report of the species in Canada. The apparent disjunction of this occurrence from other known localities in North America suggests that the population is a relic of a previously more contiguous North American distri- bution, that it is a recent colonist from the southeast, or that the species is present between the new sites and the ones further south but unrecorded. We think the latter is unlikely given the search effort for mosses and liverworts in much of the northeastern United States and the uniqueness of the Nova Scotia habitat. Alternatively, H. molleri subsp. integra may be a dispersal-limited disjunct of Nova Scotia’s Atlantic Coastal Plain flora. This species is only rarely fertile, even in locations where it is more common and abun- dant (Breil 1970; Schuster 1980). Consequently, re- productive propagules are unlikely to have colonized any new habitats in recent years. Moreover, other spe- cies that are associated with this type of habitat (both vascular plants and epiphytes) are known to be as- sociated with the Atlantic Coastal Plain, for which southwestern Nova Scotia forms a natural northern disjunction (Sweeny and Ogilvie 1993). Regardless, the combination of potential dispersal limitation with habitat and substrate associations, makes H. molleri subsp. integra an exceptional rarity, even among flora of the Atlantic Coastal Plain. The first record in Canada? Although our find was exceptional, it may not be the first detection of this species in Canada; a single collection of H. molleri was supposedly made by T. Drummond in the early 19th century and is held by the New York Botanical Garden. The location origin- ally listed in the digital record of the specimen was “British Columbia” (Consortium of North American Bryophyte Herbaria 2018), but the writing on the packet says “British North America’, a vague term, which, at the time the collection was made (ca. 1830), could have referred to all of the British territories north of the United States (Nicholson 2006) or primarily those west of Upper Canada, which was both the of- ficial name and a more commonly used descriptor for material collected by Drummond from what is now southern Ontario (Consortium of North American Bryophyte Herbaria 2018). If the specimen was in- deed from Ontario, Drummond would probably have made this collection at the beginning of his exped- ition in 1825 (the only time he visited Ontario), which began in the Niagara area, and proceeded toward Lake Superior, and then on to the Rocky Mountains via the Saskatchewan River route (Bird 1967). It is possible that he considered much of northwestern Ontario to be outside of Upper Canada sensu stricto. THE CANADIAN FIELD-NATURALIST Vol. 133 However, we have reasons to doubt that this col- lection was from Canada. First, the collection was part of William Mitten’s herbarium, which was both extensive and somewhat poorly organized and anno- tated (Thiers 1983), casting a general doubt on the ac- curacy of packet labels. Second, the associated taxa in Drummond’s col- lection (D. albicans and F. tamarisci) are, in Canada, primarily known from either the British Columbia coast or the Atlantic provinces, where Drummond did not collect; other supposed early records of D. albicans (Macoun 1902) probably represent Diplo- Phyllum taxifolium (Wahlenberg) Dumortier (Ley and Crowe 1999), Third, most other Drummond bryophyte collec- tions from the Ontario region list a specific area (e.g., “Lake Superior” or “Niagara Falls”; Consortium of North American Bryophyte Herbaria 2018), many of which would have been considered part of Upper Canada, rather than “British North America”. Fourth, while lands around Lake Superior are known to harbour some rare taxa associated with cedar swamps (e.g., COSEWIC 2019), neither Drum- mond’s own records nor those of others who have studied Drummond’s work (Bird 1967) suggest that he sampled extensively in cedar swamps of Ontario. Fifth, the Drummond H. molleri collection could be from another location entirely: the specimen is somewhat morphologically ambiguous, with stem cortical cells suggestive of the European subspecies, perhaps from the United Kingdom (UK), and other morphological aspects suggestive of H. molleri subsp. integra, perhaps from the southeastern USA. Drummond is known to have sampled bryophytes extensively in the UK before his work in North America, as exemplified in his two-volume Musci Scotici (Geiser 1937), and to have travelled widely throughout the southeastern USA in the 1830s, amass- ing thousands of specimens, including exsiccatae en- titled Musci Americani and Musci Louisiana, which were posthumously released by Hooker and Wilson (Hooker 1840; Short 1841; Geiser 1937). The associ- ated taxa in his H. molleri collection do not provide definitive guidance on alternative localities: in North America, D. albicans and F. tamarisci are known pri- marily from the Pacific Northwest or from Atlantic Canada and the Appalachian range of the USA, but have also been recorded in the UK. Nevertheless, we believe the collection was more likely to have been from the southeastern USA. Although hepatics were a minority in all of Drummond’s collections and are not fully enumerated in any documents we could lo- cate, Evans (1902) reports that Drummond’s “Mosses of the Southern States” contains Jungermannia serpyllifolia. Although this name was later consid- 2019 ered a synonym of L. cavifolia (Evans 1902), at the time Drummond was collecting, H. molleri subsp. integra was known as J. serpyllifolia subsp. ovata (Grolle 1989), and the omission of such a subspecific designation could have been easily overlooked by later handlers of this material. Even if the specimen to which Evans (1902) referred was, indeed, L. cavi- folia, it suggests that Drummond collected in the right type of habitat to have also recovered H. molleri. Significance and conservation Although it may be the first Canadian record, our Nova Scotian collection of H. molleri subsp. integra was not entirely unexpected: the rich swamp forests of southwest Nova Scotia harbour several rare spe- cies that are unknown elsewhere (e.g., Neily and Anderson 2010) or are otherwise restricted to the southern Appalachians or Atlantic Coastal Plain of North America (Wisheu and Keddy 1989; Sweeny and Ogilvie 1993). The other liverworts found in the colony with H. molleri subsp. integra are themselves rare or uncommon in Atlantic Canada, having been reported only a handful of times in Nova Scotia (R. Newell pers. comm. 31 May 2017). The Hectanooga Cedar Swamp, in which our specimens were collected, has been viewed as rare and exceptional in Nova Scotia for several decades (Ogilvie 1984), but its ecological importance has only been recognized more recently. In addition to an absence of historical disturbance in large parts, with some trees nearly 200 years old (Nova Scotia Department of Environment 2013a), the swamp har- bours the largest number of naturally occurring Eastern White Cedar in mainland Nova Scotia (Nova Scotia Nature Trust 2010). The swamp also harbours many rare and at-risk species of lichens (COSEWIC 2009, 2010, 2015, 2016), birds (COSEWIC 2007, 2008), and trees, including Eastern White Cedar (Newell 2005). The Hectanooga Cedar Swamp is, therefore, of considerable value for biodiversity con- servation and scientific research. Historically, much of the Hectanooga Cedar Swamp was privately owned, but large parts are now scheduled to be protected by a provincial Nature Reserve. In 2010, the Nova Scotia Nature Trust pur- chased 75 ha of this land, and later transferred owner- ship of it to the provincial government with the pro- tection of a conservation easement. These lands, combined with an adjacent area of Crown land to the north, are proposed as the Hectanooga Cedar Swamp Nature Reserve, including both important swamp forest and some mature mixed hardwood forest to reduce the negative edge influence (Nova Scotia Department of Environment 2013b). On the other hand, logging activities between 2008 and 2012 had already removed a substantial area of adjacent old- HAUGHIAN AND NEILY: LIVERWORT NEW TO ATLANTIC CANADA 203 growth forest, and several roads run along the edges of the proposed reserve (S.R.H. and T.H.N. pers. obs.). As such, the reserve may yet suffer from nega- tive edge influence, exacerbated by its small size (124 ha), fragmented configuration (divided into three sec- tions), and elongate shape. Such forested wetlands may be declining in Nova Scotia, and these declines may be exacerbated in the future in a warming cli- mate (Newell 2005; Lemieux 2010). We recommend enhancing protections for such unique hotspots of biodiversity by promptly conferring legal protected status upon them wherever possible, by adding addi- tional parcels to make the reserves contiguous, and by increasing reserve sizes to increase protection from adjacent industrial activities. Vouchers examined Harpalejeunea molleri subsp. integra (R.M. Schust.) Damsh—CANADA, NOVA SCOTIA: Digby Co., Hectanooga Cedar Swamp, 44.082°N, 66.056°W, 17 May 2017, 7: Neily 1629 (NBM BH- 2739); ibidem:. 44.083°N, 66.052°W, 17 May 2017, 7: Neily 1654 (personal collection of T.H.N., Digby Co.); U.S.A., TENNESSEE: Pickett Co., rocky slopes W of Hwy 154 near Scott Co. line, Pickett State Forest, 17 April 1991, P.G. Davison 1613 (F-C0074242, as H. ovata subsp. integra), SOUTH CAROLINA: Oconee, gorge of Whitewater River, 0.3-—0.4 mi. (0.5—0.6 km) below Lower Falls, ca. 3 mi. (4.8 km) above Jocassee, 24 August 1958, R.M. Schuster 40899a (F-C0578334, as H. ovata subsp. integra); BRITISH NORTH AMERICA: ca. 1825-1835 (en- tered as 1906), 7. Drummond s.n. (NY00265235, as Lejeunea ovata). Harpalejeunea molleri (Steph.) Grolle subsp. molleri—SPAIN: 1927, P. Allorge, Exsiccata Bryo- theca Iberica No. 11 (NBM BH-00858, as H. ovata); PORTUGAL: 1937, P. Allorge, Exsiccata Bryophyta Azorica No. 37 (NBM BH-00519, as H. ovata). Author Contributions Conceptualization — S.R.H. and T.H.N.; Investiga- tion (specimen discovery & identification) — T.H.N.; Investigation (specimen verifications & comparisons) — S.R.H.; Investigation (nomenclatural & historical research) — S.R.H.; Methods — S.R.H. and T.H.N.; Visualization (photography) — S.R.H.; Writing (ori- ginal draft preparation) — S.R.H.; Writing (review & editing) — S.R.H. and T.H.N. Acknowledgements This work was self-funded by the authors. We wish to thank Stephen Clayden and Kendra Driscoll for assistance in accessing specimens at the New Brunswick Museum, Ruth Newell for assistance with searching for Nova Scotian liverwort records at the 204 Acadia University Herbarium, Barbara Thiers for assistance in locating the Drummond record at the New York Botanical Garden, the Field Museum and New York Botanical Garden Herbaria for loans of material, and three anonymous reviewers, Jennifer Doubt, Jeff Saarela, and Dwayne Lepitzki for their insightful feedback on our draft manuscript. Literature Cited Bates, J.W. 1998. Is “life-form” a useful concept in bryo- phyte ecology? Oikos 82: 223-237. https://doi.org/10. 2307/3546962 Bird, C.D. 1967. The mosses collected by Thomas Drum- mond in Western Canada, 1825-1827. Bryologist 70: 262-266. https://doi.org/10.2307/3240954 Breil, D.A. 1970. Liverworts of the Mid-Gulf Coastal Plain. Bryologist 73: 409-491. https://do1.org/10.2307/3241490 Consortium of North American Bryophyte Herbaria. 2017. Harpalejeunea ovata (Hooker) Schiffner subsp. integra R.M. Schuster. Accessed 23 November 2017. http://bryophyteportal.org/portal/collections/list.php. Consortium of North American Bryophyte Herbaria. 2018. CNABH search results for: Country = Canada; Collector = Drummond; Dates = 1820 to 1908. Accessed 20 September 2018. http://bryophyteportal.org/portal/collections/list.php ?db=all&country=Canada&collector=Drummond&event datel=1820&eventdate2=1908&page=1. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2007. COSEWIC assessment and status report on the Olive-sided Flycatcher Contopus cooperi in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2008. COSEWIC assessment and status report on the Canada Warbler Wilsonia canaden- sis. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2009. COSEWIC assessment and status report on the Vole Ears Erioderma mollissimum in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2010. COSEWIC assessment and status report on the Blue Felt Lichen Degelia plumbea in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2015. COSEWIC assessment and status report on the Black-foam Lichen Anzia colpodes in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2016. COSEWIC assessment and status report on the Wrinkled Shingle Lichen Pannaria lurida in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2019. COSEWIC assessment and status report on the White-rimmed Shingle Lichen Fus- copannaria leucosticta in Canada. COSEWIC, Ottawa, Ontario, Canada. Damsholt, K., and A. Pagh. 2002. Illustrated Flora of Nordic Liverworts and Hornworts. Nordic Bryological Society, Lund, Sweden. THE CANADIAN FIELD-NATURALIST Vol. 133 Evans, A.W. 1902. The Lejeuneae of the United States and Canada. Memoirs of the Torrey Botanical Club 8: 113-183. GBIF (Global Biodiversity Information Facility). 2018. Harpalejeunea molleri (Steph.) Grolle. GBIF Backbone Taxonomy, GBIF Secretariat, Copenhagen, Denmark. Checklist dataset. Accessed 22 September 2018. https:// doi.org/10.15468/390mei Geiser, S.W. 1937. Thomas Drummond. Naturalists on the Frontier. Southern Methodist University, Dallas, Texas, USA. Grolle, R. 1989. A technically new lectotypification of Har- palejeunea (Hepaticae). Taxon 38: 88-90. https://do1. org/10.2307/1220901 Hilchey, J.D., D.B. Cann, and J.I. MacDougall. 1962. Soil survey of Digby County, Nova Scotia. Nova Scotia Soil Survey, Truro, Nova Scotia, Canada. Hodgetts, N. 2015. Checklist and country status of European bryophytes—towards a new Red List for Europe. Irish Wildlife Manuals, Department of Arts, Heritage, and the Gaeltacht, Ireland, Dublin, Ireland. Hooker, W. 1840. Announcement: Thomas Drummond’s Mosses and Hepaticae of Louisiana. Journal of Botany 1: 438. Lemieux, M.J. 2010. A management plan for native oc- currences of eastern white cedar (Thuja occidentalis L.) in Nova Scotia, Canada. Nova Scotia Department of Natural Resources, Truro, Nova Scotia, Canada. Ac- cessed 26 February 2018. https://novascotia.ca/natr/ wildlife/biodiversity/pdf/Management_Plan_EWC_NS_ July_2010_MJL.pdf. Ley, L.M., and J.M. Crowe. 1999. An Enthusiast’s Guide to the Liverworts and Hornworts of Ontario. Lakehead University, Thunder Bay, Ontario, Canada. Macoun, J. 1902. Catalogue of Canadian Plants. VII— Lichenes and Hepaticae. Government Printing Bureau, Ottawa, Ontario, Canada. Accessed 26 February 2018. https://www. biodiversitylibrary.org/item/144111. Neily, T., and F. Anderson. 2010. Leptogium hibernicum Mitch. ex P.M. Jorg. discovered in North America. Lichenologist 42: 629-630. https://doi.org/10.1017/S00 24282910000162 Newell, R.E. 2005. Provincial (Nova Scotia) status report on Northern White Cedar Thuja occidentalis. Nova Scotia Department of Natural Resources, Truro, Nova Scotia, Canada. Accessed 26 February 2018. https://novascotia. ca/natr/wildlife/biodiversity/pdf/statusreports/Status ReportEasternCedar.pdf. Nicholson, N.L. 2006. British North America. The Can- adian Encyclopedia. Online. Historica Canada, To- ronto, Ontario, Canada. Accessed 11 January 2020. https://www.thecanadianencyclopedia.ca/en/article/ british-north-america. Nova Scotia Department of Environment. 2013a.Ourparks and protected areas: a plan for Nova Scotia. Nova Scotia Department of Environment, Halifax, Nova Scotia, Can- ada. Accessed 26 February 2018. https://novascotia.ca/ parksandprotectedareas/pdf/Parks-Protected-Plan.pdf. Nova Scotia Department of Environment. 2013b. Hecta- nooga Cedar Swamp Nature Reserve. Nova Scotia 2019 Department of Environment, Halifax, Nova Scotia. Accessed 26 February 2018. https://www.novascotia.ca/ nse/protectedareas/consult/AreaHandler.ashx?id=518 &type=html. Nova Scotia Nature Trust. 2010. Nature trust saves homes for imperiled species. News release. Nova Scotia Nature Trust, Halifax, Nova Scotia, Canada. Ogilvie, R. 1984. Important ecological sites in Nova Scotia. Curatorial report 49. Nova Scotia Museum, Halifax, Nova Scotia, Canada. Paton, J.A. 1999. The Liverwort Flora of the British Isles. Brill Academic Publishers, Leiden, Netherlands. Schuster, R.M. 1980. The Hepaticae and Anthocerotae of North America, East of the 100th Meridian. Volume IV. Columbia University Press, New York, New York, USA. Short, C.W. 1841. A sketch of the progress in botany in Western North America. Journal of Botany 3: 97-124. Sweeny, S., and R. Ogilvie. 1993. The conservation of coastal plain flora in Nova Scotia. Maine Naturalist 1: HAUGHIAN AND NEILY: LIVERWORT NEW TO ATLANTIC CANADA 205 131-144. https://doi.org/10.2307/3858236 Thiers, B.M. 1983. Index to the genera and species of hepa- ticae described by William Mitten. Brittonia 35: 271— 300. https://doi.org/10.2307/2806026 White, C.E., and R.J. Horne. 2012. Bedrock geology map of the Meteghan area, NTS sheet 21B/01, Digby and Yarmouth counties, Nova Scotia. Nova Scotia Department of Natural Resources, Mineral Resource Branch, Halifax, Nova Scotia, Canada. Accessed 26 February 2018. https://novascotia.ca/natr/meb/data/mg/ ofm/pdf/ofm_2012-086_d127_dp.pdf. Wisheu, I.C., and P.A. Keddy. 1989. The conservation and management of a threatened coastal plain plant commun- ity in eastern North America (Nova Scotia, Canada). Biological Conservation 48: 229-238. https://do1.org/10. 1016/0006-3207(89)90120-1 Received 26 February 2018 Accepted 17 December 2019 The Canadian Field-Naturalist Lichens and allied fungi of Sandbar Lake Provincial Park, Ontario Hanna R. Dorval" and RICHARD TROY MCMULLIN? 'Lakehead University, Natural Resources Management, 955 Oliver Road, Thunder Bay, Ontario P7B 5E1 Canada Canadian Museum of Nature, Research and Collections, P.O. Box 3443 Stn. D, Ottawa, Ontario K1P 6P4 Canada “Corresponding author: hdorval@lakeheadu.ca Dorval, H.R., and R.T. McMullin. 2019. Lichens and allied fungi of Sandbar Lake Provincial Park, Ontario. Canadian Field- Naturalist 133(3): 206-215. https://do1.org/10.22621/cfn.v13313.2209 Abstract Sandbar Lake Provincial Park (Sandbar Lake) covers 8053 ha in the boreal forest in northwestern Ontario. Within the park boundary are natural forests representative of those in the region, as well as forests that are heavily disturbed from resource extraction activities, which are prevalent in northwestern Ontario. The lichen biota in this part of the boreal forest is known to be rich and abundant, but lichen diversity is also known to be negatively impacted by disturbances (e.g., timber harvest- ing, mining, and climate change). Therefore, lichens can be used to monitor the effects of these disturbances, but baseline data are required. Here, we present the results of the first detailed inventory of the lichens and allied fungi of Sandbar Lake. We report 139 species in 69 genera from 16 sites that represent all macrohabitats present in the park. Seven species have a provincial conservation status rank from S1 to S3 (critically imperilled to vulnerable), and one species, Arthrosporum populorum, has previously been collected only once in northwestern Ontario. Our results fill biogeographic gaps for many species and allow lichens to be used as biomonitors during further study at Sandbar Lake. We show that Sandbar Lake has important conservation value, and our data provide an opportunity for further study in an area with no previous research on lichens. Key words: Sandbar Lake Provincial Park; lichens; fungi; boreal forest; Great Lakes—St. Lawrence forest; conservation; biogeography; bioindicators; protected areas Introduction Provincial parks in Ontario are designed to main- tain and preserve natural and cultural integrity while allowing for recreational and educational op- portunities for the public and scientific commun- ities (Ontario 2015). They are regulated under the Provincial Parks and Conservation Reserves Act and, since 1954, have been managed by Ontario Parks, a branch of the Ontario Ministry of Natural Resources and Forestry. Between 1920 and 1954, they fell under the Department of Lands and Forests. The first prov- incial parks in Ontario were created from land that was considered unsuitable for agriculture and settle- ment. However, now parks are also established to pro- vide opportunities for outdoor recreation and the re- sulting economic benefits. Parks allow the public and researchers to gain knowledge of the natural herit- age of Ontario; they also protect the biodiversity, eco- systems, and provincially significant elements within their boundaries (Ontario 2006). Currently, more than 7 420 816 ha in Ontario have been incorporated into 335 provincial parks, accounting for 7% of the land area in the province (Ontario 2017). Sandbar Lake Provincial Park was established in 1970 (Ontario Parks 2012). It was initially classi- fied as a recreation park but was changed to its cur- rent classification as a natural environment park in 1986 (Ontario Parks 2012). This designation dictates that the management goals include maintenance of ecosystem representativeness and natural and cul- tural heritage, while allowing for recreational, edu- cational, and research activities (Ontario Parks 2012). The park area includes Sandbar Lake, sand beaches, and conifer-dominated forests. Since it was estab- lished, two additions to the park have doubled its size to over 8000 ha (Ontario Parks 2012). Sandbar Lake Provincial Park is in a region northwest of Lake Superior that is known for rich lichen diversity (Crowe 1994; Ahti and Crowe 1995) and as a hotspot for lichen diversity in North America (Brodo et al. 2001). However, the lichens of the Sandbar Lake area have not been documented previously. The history of lichen collecting in Ontario has been summarized by McMullin and Lewis (2013). Of the 1083 species known from the province (McMullin et al. 2015, 2018), at least 455 occur in the region northwest of Lake Superior (Crowe 1994; Ahti and Crowe 1995). Despite the known diversity, the only A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 206 ©The Ottawa Field-Naturalists’ Club 2019 focussed studies within this region of Ontario have been in Slate Islands National Park (C. Wetmore un- publ. data accessed through the Consortium of North American Lichen Herbaria [CNALH]) and Voyageur National Park in Minnesota, USA (Wetmore 1981). The aim of our study was to document the lichens and allied fungi in Sandbar Lake Provincial Park. Our objectives were to examine all major ecosystems in the park, create a checklist of lichens and allied fungi traditionally treated with lichens, compare our results with those from two other provincial parks in Ontario, and provide provincial conservation status ranks for each species. Our results will provide an in- creased understanding of the biodiversity in the park and a baseline that can be used to detect changes to the lichen community from disturbances, such as cli- mate change, acid rain, or land use changes in or near the park (McMullin et al. 2017). Study Area Sandbar Lake Provincial Park is located in north- western Ontario, ~4 km northeast of the town of Ignace and ~250 km northwest of Thunder Bay (Figure 1), along Highway 599. This protected area covers 8053 ha. When the park was established in 1970, its area was 3157 ha. In 1986, 1926 ha were added and, in 2003, Ontario’s Living Legacy Land Use Strategy facilitated a second addition of 3720 ha on the north side of the park (Ontario Parks 2012). The latter comprises mainly wetland generated through paludification, where organic matter ac- cumulation, especially Sphagnum spp., contributes to increased soil moisture (Lavoie et al. 2005; Ontario Parks 2012). This process results in reduced soil tem- perature, a reduction and overall change in tree and vascular plant cover, and shifts in nutrient avail- ability, microbial activity, and decomposition rates (Lavoie et al. 2005). This wetland is designated a nature reserve zone, which is a classification used to enhance protection of features represented within provincial parks that are considered unique within the region and the province (Ontario Parks 2012). Minimal to no development is permitted within these zones, but research may be permitted. Sandbar Lake Provincial Park contains five nature reserve zones, all of which were examined for lichens during this study. Natural environment zones are areas intended for limited development to generate and maintain low-intensity recreational ac- tivities. Development permitted in these zones in- cludes the provision of signage for trail navigation and the maintenance of interpretive facilities. Two natural environment zones covering 6872 ha are present in Sandbar Lake Provincial Park (Ontario Parks 2012). Historical zones were also established in the park to DORVAL AND MCMULLIN: SANDBAR LAKE PARK LICHENS AND ALLIED FUNGI 207 mark and protect culturally and historically signifi- cant areas, including those that have historically been occupied by humans and human-made structures (Ontario Parks 2012). Sandbar Lake Provincial Park has four historical zones, each delineated based on human occupation during the Laurel Period (2200-— 1600 years B.P.) and the Blackduck Period (1200 years B.P. to European contact; Ontario Parks 2012). All nature reserve, natural environment, and histor- ical zones within Sandbar Lake Provincial Park are illustrated in Figure 1. The park provides opportunities for outdoor re- creation, including hiking, canoeing, and seasonal camping at 75 campsites. All campsites and hiking trails are located in the campground area in the south- eastern corner of the park adjacent to Highway 599. Use of the campground area and recreational fish- ing in Sandbar Lake accounts for the vast majority of park use by visitors; the remainder of the park is vis- ited only infrequently. Sandbar Lake Provincial Park is located within the transition zone between the boreal and Great Lakes— St. Lawrence forests (Ontario Parks 2012). The park comprises mostly conifer-dominated forests; how- ever, mixed-wood forests, wetlands, exposed bedrock outcrops, and outwash plain ecosystem types are also represented (Ontario Parks 2012). Timber harvesting, sporadic fires, and windthrow events, which occurred mainly during the early 20th century, are largely re- sponsible for shaping the vegetation communities currently in the park (Ontario Parks 2012). In recent decades, timber harvesting and mining operations have surrounded the park boundary (Ontario Parks 2012). Methods Sampling Fieldwork was conducted in the fall of 2017. Collections were made throughout the park in all major ecosystems, nature reserves, and natural eco- system zones over 12 days. Floristic habitat sampling, completed through the intensive study of large areas, was used to evaluate species presence (Newmaster et al, 2005). This sampling technique was used at 16 sites, shown with corresponding geographic coordin- ates and habitat descriptions in Table 1. As many microhabitats as possible were examined in each site: e.g., a variety of tree species, rocks, forest floor. Most sites were visited on only one occasion; how- ever, sites I and II, Campground and Red Pine (Pinus resinosa Aiton) forest, were visited more than once. All collections have been deposited at the National Herbarium of Canada (CANL) at the Canadian Museum of Nature Natural Heritage Campus in Gatineau, Quebec. 208 THE CANADIAN FIELD-NATURALIST Vol. 133 iz Park boundary = Development zone ZZ. Historical zone FicurE 1. Sandbar Lake Provincial Park, showing designated zones and sampling sites. Identification sary, an ultraviolet light chamber was used for addition- All specimens were identified using standard tech- al chemical examination. Thin-layer chromatography niques outlined by Brodo et a/. (2001), including the __ was also used in further chemical analysis, following use of microscopy and chemical spot tests. Whenneces- Orange and White (2001) using solvents A, B', and C. 2019 TABLE 1. Location and description of collection sites. Site Location Latitude ((N) Longitude (°W) I Campground 49. 46517 91.55536 II Red Pine forest 49.46503 91.55407 III Beach forest 49.47220 91.54784 IV Mixed-wood forest* 49.48950 91.53708 V Jack Pine forest* 49.47955 91.54389 VI Boulder cave 49.48659 91.54849 VII Cliff and exposed 49. 48638 91.55068 bedrock VII Treed fen* 4952254 91.55263 IX Forest at South 49 47526 91.52035 Agimak River X Silhouette trail* 49. 46295 91.53601 XI Clearing at Flayers 49.45061 91.59702 Road XII Flayers Road fen 49.45850 91.60091 XII Forest north of 49.49306 91.56565 Sandbar Lake XIV _ Shoreline of Bog 49.48997 91.58136 Lake* XV_ Ontario rangers 49.47526 91.52035 road XVI North Agimak 49. 49087 91.60870 River DORVAL AND MCMULLIN: SANDBAR LAKE PARK LICHENS AND ALLIED FUNGI 209 Habitat description Campsites near Sandbar Lake with small, dense patches of forest between them, connected by gravel roadways. Red Pine (Pinus resinosa)-dominated forest naturally regenerated after fire, with young Balsam Fir (Abies balsamea) in the understorey, adjacent to the campground. Mixed-wood forest, dominated by Balsam Fir and Balsam Poplar (Populus balsamifera), adjacent to sandy beach along the eastern shoreline of Sandbar Lake. Deciduous-dominated mixed-wood forest with abundant fallen logs and herbaceous groundcover. Jack Pine (Pinus banksiana)-dominated forest with exposed bedrock, young Black Spruce (Picea mariana) understorey, and lichens and mosses forming large thick mats. Caves formed by large boulders leaning against a bedrock cliff, with high humidity and limited light exposure. Exposed bedrock and steep cliffs with mixed-wood forest surrounding the base. Large, humid, lowland area resulting from paludification, with sporadic Tamarack (Larix laricina) and Black Spruce, with sphagnum moss groundcover and a high diversity of wetland vascular plants. Dense, late-successional, mixed-wood forest with little understorey vegetation, near the South Agimak River and associated wetland. Mixed-wood forest along Silhouette trail/roadway, with a vegetative community resulting from past resource extraction. Treeless clearing along gravel road (Flayers Road) surrounded by Jack Pine, with sandy soil, fallen logs, and exposed bedrock. Fen, near Flayers Road, dominated by Tamarack and Black Spruce, with moss covering the ground and some pools of standing water. Mixed-wood forest with highly variable structure and age, located between the northeastern shoreline and | km north of Sandbar Lake. Sedge mat with abundant Tamarack and Black Spruce, few shrubs, and many dead standing trees, surrounding the small lake. Road re-colonized by forest dominated by young Balsam Fir, between two lakes to the north and south of the road. River with exposed and mossy boulders, surrounded by dense mixed-wood forest. *Indicates the sampling sites in each of the park’s five nature reserve zones. Sorensen—Dice coefficient of similarity To compare the lichen community at Sandbar Lake Provincial Park with two other locations in Ontario, we used the Sorensen—Dice coefficient of similarity (Dice 1945; Sorensen 1948). This coefficient is calcu- lated as follows: 2A(2A +B+C) where A is the total number of species at Sandbar Lake Provincial Park and another location (e.g., lo- cation 2), B is the number of species at Sandbar Lake Provincial Park that are absent from location 2, and C is the number of species at location 2 that are ab- sent from Sandbar Lake Provincial Park (Dice 1945; Sorensen 1948). The two locations included for comparison, Awenda Provincial Park (McMullin and Lendemer 2016) and Sandbanks Provincial Park (McMullin and Lewis 2014), are study sites with comparable search efforts nearest to Sandbar Lake Provincial Park. Conservation status We report the conservation status (S ranks) for each species recorded in the park. The Ontario Na- 210 tural History Information Centre (NHIC) assigns these non-legal provincial conservation status ranks to species in Ontario, based on guidelines set out by NatureServe (2018). If adequate information is known about the presence of a species in the prov- ince, then a rank between 1 and 5 is assigned. An S rank of 1 denotes a species that is considered critic- ally imperilled, 2 means imperilled, 3 is vulnerable, 4 is apparently secure, and 5 is secure. Other ranks include NR meaning not ranked, U meaning unrank- able (because of lack of information), and ? meaning the rank is uncertain. Species with a rank of S1 to S3 are provincially tracked. Observations of these spe- cies in the province are considered remarkable and are reported to the NHIC. Species with a rank above S3 or are unranked are not provincially tracked and are considered to be fairly common within the province. Results We located 139 species in 69 genera at Sandbar Lake Provincial Park (see Annotated species list): 54 (39%) species are crustose, 52 (37%) are foliose, and 33 (24%) are fruticose. For 122 species (88%), green algae are the primary photobiont; for 13 (9%), cyano- bacteria are the primary photobiont; and four (3%) are non-lichenized allied fungi. Sorensen—Dice coefficient of similarity Sorensen—Dice coefficients of similarity were de- termined for each of the two other provincial parks and Sandbar Lake Provincial Park. The lichen com- munity at Sandbar Lake Provincial Park is more sim- ilar to that of Sandbanks Provincial Park (coefficient value of 0.65) than to that of Awenda Provincial Park (coefficient value of 0.49). The number of species (n = 139) at Sandbar Lake Provincial Park was also more similar to the number found at Sandbanks (n = 122) than Awenda (n = 203). Sandbar Lake Provincial Park is considerably larger than both other provincial parks in this comparison (Table 2). Conservation status Of the 139 species discovered at Sandbar Lake Provincial Park, 125 have been assigned conservation status ranks. Seven of these species have a conserv- ation status rank between SI and S3 (critically im- perilled, imperilled, vulnerable) and are provincially THE CANADIAN FIELD-NATURALIST Vol. 133 tracked. One species is listed as S1S2: Arthrosporum populorum A. Massl.; two are S2: Bacidia laur- ocerasi (Delise ex Duby) Zahlbr. and Ochrolechia pseudopallescens Brodo; two are S2S3: Calicium parvum Tibell and Chaenothecopsis pusilla (Ach.) A.E.W. Schmidt; and two are S3: Cetrelia chicitae (W.L. Culb.) W.L. Culb. & C.F. Culb. and Melanelixia glabratula (Lamy) Sandler & Arup. The non-tracked species include 19 that are S4, seven that are S4?, 27 that are S4S5, 65 that are S5, nine that are not ranked, and five that are unrankable. Annotated species list This list is organized alphabetically by genus and species, and taxonomic authorities follow the 23rd version of the North American Lichen Checklist (Esslinger 2018), as does most of the nomenclature. Any differences between this list and Esslinger’s re- flects the opinion of the authors. Substrate follows species name and taxonomic authorities. Roman num- erals indicate the collection site (Table 1). Provincial conservation status rank follows the collection site. Non-lichenized fungi typically treated with lichens are preceded by a dagger (‘). Acarospora fuscata (Schrad.) Arnold—Saxicolous. VII. S5 Amandinea punctata (Hoffm.) Coppins & Scheid.— Corticolous on Balsam Fir (Abies balsamea (L.) Miller). I. SS. Arthonia sp—Corticolous on a snag. I. SNR. Arthrosporum populorum A. Mass|—Corticolous on a fallen Trembling Aspen (Populus tremuloides Michaux). XIII. S1S2. Athallia pyracea (Ach.) Arup, Frodén & Sochting— Corticolous ona fallen P. tremuloides. XII. SU. Bacidia laurocerasi (Delise ex Duby) Zahlbr.—Cor- ticolous. I. S2. Baeomyces rufus (Huds.) Rebent.—Terricolous on sandy soil. [X. S4S5. Biatora pycnidiata Printzen & Tonsberg—Cortic- olous on A. balsamea and Black Spruce (Picea mariana (Miller) Britton, Sterns & Poggenburgh). IV, XV. SNR. Biatora vernalis (L.) Fr—Bryicolous. I. S5. Bryoria sp.—Corticolous on P. mariana and dead P. mariana. 1, XII, VII. SNR. TABLE 2. Sorensen—Dice coefficient of similarity between Sandbar Lake Provincial Park community and two other park communities in Ontario. Approximate distance Provincial park from Sandbar Lake Area (ha) Nes SON ce ihe of species coefficient Provincial Park Sandbar Lake 0 8053 139 1 Awenda 1018 km southeast 2915 203 0.49 Sandbanks 1254 km southeast 1551 122 0.65 2019 Bryoria furcellata (Fr.) Brodo & D. Hawksw.—Cor- ticolous on P. mariana and Jack Pine (Pinus bank- siana Lambert). I, III, 1X, XIII, XIV. SS. Bryoria fuscescens (Gyeln.) Brodo & D. Hawksw.— Corticolous on A. balsamea. XII. S5. Bryoria kockiana Velmala, Myllys & Goward—Cor- ticolous on A. balsamea and White Spruce (Picea glauca (Moench) Voss. I. S4. Bryoria trichodes subsp. trichodes (Michx.) Brodo & D. Hawksw.—Corticolous on P. mariana. XII, XIV. SS. Buellia erubescens Arnold—Corticolous on A. bal- samea and Paper Birch (Betula papyrifera Mar- shall). I, X, XV. SS. Calicium parvum Tibell—Corticolous on P. resi- nosa. II. S283. Calicium trabinellum (Ach.) Ach.—Lignicolous on a snag. X. S4S5. Caloplaca arenaria (Pers.) Mill. Arg—Saxicolous. VII. SS. Caloplaca cerina (Ehrh. ex Hedwig) Th. Fr.—Cor- ticolous on a fallen P. tremuloides. XIII. SS. Caloplaca chrysophthalma Degel.—Corticolous on a fallen P. tremuloides. 1, IV. S4? Candelariella lutella (Vainio) Rasénen—Corticolous on Alnus sp. and ona fallen P. tremuloides. 1, XIII. SNR. Candelariella vitellina (Hoffm.) Mull—Saxicolous. VII. SS. Carbonicola anthracophila (Ny1.) Bendiksby & Tim- dal—Lignicolous on burned wood. II. $4? Cetrelia chicitae (W.L. Culb.) W.L. Culb. & C.F. Culb.—Saxicolous. XVI. $3? Chaenotheca brunneola (Ach.) Mill. Arg—Ligni- colous on a snag. II. S4. Chaenotheca chrysocephala Turner ex Ach.) Th. Fr. —Corticolous on P. mariana. XIII. S4. Chaenotheca ferruginea (Turner ex Sm.) Mig.—Cor- ticolous on a charred conifer and P. resinosa. II, X. S4. tChaenothecopsis nana Tibell—Corticolous on P. mariana. XIII. SU. +Chaenothecopsis pusilla (Ach.) A.FW. Schmidt— Corticolous on P. resinosa. II. S2S3. Chrysothrix caesia (Flot.) Korb—Corticolous on B. papyrifera. 1. SS. Cladonia botrytes (K.G. Hagen) Willd. —Lignicolous on dead P. mariana. VIII. SS. Cladonia cenotea (Ach.) Schaerer—Lignicolous on rotting wood. II. SS. Cladonia cornuta (L.) Hoffm.—Terricolous. XI. S4S5. Cladonia cristatella Tuck.—Lignicolous on a rotted stump; saxicolous; terricolous. I, V. S5. Cladonia deformis (L.) Hoffm.—Terricolous on thin soil. V. SS. DoRVAL AND MCMULLIN: SANDBAR LAKE PARK LICHENS AND ALLIED FUNGI 211 Cladonia digitata (L.) Hoffm.—Lignicolous on rot- ting wood and a stump. I, II. S4S5. Cladonia macilenta var. macilenta Hoffm.—Terri- colous. XI. SS. Cladonia merochlorophaea Asah.—Terricolous. I. S4. Cladonia mitis Sandst—Saxicolous. V. S5. Cladonia ochrochlora Flérke—Lignicolous on rot- ting wood; terricolous. II, XI. S5. Cladonia parasitica (Hoftm.) Hoffm.—Lignicolous on a log. II. S4. Cladonia phyllophora Hofftm.—Terricolous. V, XI. SS. Cladonia pyxidata (L.) Hoffm.—Terricolous. I. S5. Cladonia rangiferina (L.) F.H. Wigg.—Saxicolous on bedrock. V. S5. Cladonia stellaris (Opiz) Pouzar & Vézda—Saxico- lous on bedrock. V. S5. Cladonia uncialis (L.) F.H. Wigg.—Terricolous. V. S5. Cladonia verticillata (Hoffm.) Schaer.—Terricolous. XI. S4S5. Dermaatocarpon luridum (With.) J.R. Laundon— Saxicolous. XVI. SS. Dimelaena oreina (Ach.) Norman—Saxicolous. VII. S4. Evernia mesomorpha Ny|.—Corticolous on A. bal- samea, P. mariana, and a snag. I, II, HI. SS. Flavoparmelia caperata (L.) Hale—Corticolous on B. papyrifera, saxicolous. I, VI. SS. Flavopunctelia flaventior (Stirt.) Hale—Corticolous on A. balsamea. I. SS. Fuscidea arboricola Coppins & Tonsberg—Cortico- lous on A. balsamea. XV. SU. Heterodermia speciosa (Wulfen) Trevisan—Bryico- lous. I. S4S5. Hypogymnia incurvoides Rass.—Corticolous on P. mariana. XIV. S4. Hypogymnia physodes (L.) Nyl—Corticolous on A. balsamea and Picea sp. I, I, IV, XII, XV. SS. Hypogymnia tubulosa (Schaer.) Hav.—Corticolous on A. balsamea and Picea sp. I, XII. $4? Imshaugia aleurites (Ach.) S.F. Meyer—Corticolous on a burned snag and dead Eastern White Cedar (Thuja occidentalis L.). I, VIII. S5. Imshaugia placorodia (Ach.) S.F. Meyer—Cortico- lous on a P. mariana log. IX. S485. Julella fallaciosa (Arnold) R.C. Harris—Corticolous on B. papyrifera. |. S4? Lasallia papulosa (Ach.) Llano—Saxicolous. VI. S5. Lecanora albella var. rubescens (Imshaug & Brodo)— Corticolous on B. papyrifera. XII. SNR. Lecanora allophana (Ach.) Nyl.—Corticolous on a snag. I. S5. Lecanora allophana f. sorediata Vain.—Corticolous on B. papyrifera and P. tremuloides. 1, [X. S5. 212 Lecanora circumborealis Brodo & Vitik.—Cortic- olous on Tamarack (Larix laricina (Du Roi) K. Koch). VUL SS. Lecanora polytropa (Ehrh.) Rabenh.—Saxicolous. VII. S5. Lecanora pulicaris (Pers.) Ach——Corticolous on B. papyrifera, lignicolous on a Pinus sp. cone. I, XIII. S5. Lecanora symmicta (Ach.) Ach.—Corticolous on B. papyrifera and Eastern White Pine (Pinus strobus L.). I, X. SS. Lecanora thysanophora R.C. Harris—Corticolous on A. balsamea and a deciduous tree. XIII. S4S5. Lepra trachythallina (Erichsen) Lendemer & R.C. Harris—Corticolous on T: occidentalis. IV. S4. Lepraria finkii(B. de Lesd.) R.C. Harris—Corticolous on 7. occidentalis, saxicolous. VI, XVI. SS. Leptogium cyanescens (Rabenh.) Korb.—Saxicolous on a mossy boulder. I, XVI. S5. Leptorhaphis epidermidis (Ach.) Th. Fr—Cortico- lous on B. papyrifera. 1X. S4. Lobaria pulmonaria (L.) Hoffm.—Corticolous on P. tremuloides. VII. S4. Melanelixia glabratula (Lamy) Sandler & Arup— Corticolous on dead P. strobus. XIV. S3. Melanelixia subaurifera (Nyl.) O. Blanco et al._— Corticolous on B. papyrifera; saxicolous. I, X. S5. Melanohalea exasperatula (Nyl.) O. Blanco et al.— Corticolous on Alnus sp. I. S485. Mycobilimbia berengeriana (A. Massal.) Hafellner & V. Wirth—Terricolous. I. S4S5. Mycoblastus sanguinarius (L.) Norman—Cortico- lous on P. mariana. VIII. S4S5. Mycocalicium subtile (Pers.) Szatala—Lignicolous on a snag. I, II. S4S5. Myelochroa glabina (Ach.) Elix & Hale—Corticolous on B. papyrifera. 1. S4SS. Nephroma helveticum Ach.—Saxicolous. XVI. S4S5. Nephroma parile (Ach.) Ach.—Saxicolous on a mossy rock; terricolous. I, X VI. S4SS5. Nephroma resupinatum (L.) Ach—Corticolous on Mountain Maple (Acer spicatum Lamarck); sax- icolous on a mossy rock; terricolous. I, XIII. $4. Ochrolechia arborea (Kreyer) Almb.—Corticolous on B. papyrifera and P. mariana. I, XIV. S485. Ochrolechia pseudopallescens Brodo—Corticolous on P. mariana and dead P. mariana. VIII, XIV. S2. Parmelia squarrosa Hale—Corticolous on B. papyr- ifera. 1, XIII. SS. Parmelia sulcata Taylor—Corticolous on B. papyr- ifera and ona snag. II, X. S5. Parmeliopsis capitata R.C. Harris ex JW. Hinds & P.L. Hinds—Corticolous on a conifer, L. laricina, P. mariana, and dead P. strobus. I, V, VII, XII. SS. THE CANADIAN FIELD-NATURALIST Vol. 133 Parmeliopsis hyperopta (Ach.) Arnold—Corticolous on dead P. strobus and a snag. I, V. SS. Peltigera aphthosa (L). Willd—Terricolous on mossy soil. II. S5. Peltigera canina (L.) Willd.—Lignicolous on a rot- ted log; terricolous on the base of a rock. I, IV. SS. Peltigera elisabethae Gyeln.—Terricolous on sandy soil. I, III, IV. SS. Peltigera evansiana Gyeln.—Bryicolous on a mossy boulder. XVI. S4. Peltigera extenuata (Nyl. ex Vainio) Lojka—Saxico- lous and terricolous. I, XHI, XVI. $4? Peltigera horizontalis (Huds.) Baumg.—Lignicolous on a rotted log; saxicolous; terricolous on the base of a rock. I, IV, XHI. SS. Peltigera malacea (Ach.) Funck—Terricolous. X. S485. Peltigera neckeri Hepp ex Mull. Arg—Saxicolous. XI. SS. Peltigera polydactylon (Neck.) Hoffm.—Saxicolous. I. SS. Peltigera rufescens (Weiss) Humb.—Terricolous. I. SS. Pertusaria rubefacta Erichsen—Corticolous on A. spicatum. XIII. S4? Phaeophyscia adiastola (Essl.) Ess!—Saxicolous. I, XVI. S4. Phaeophyscia hirtella Ess|—Corticolous on P. tre- muloides. 1. S4. Phaeophyscia pusilloides (Zahlbr.) Essl—Bryico- lous; corticolous on A. spicatum and P. tremul- oides. I, XIII. SS. Physcia adscendens (Fr.) H.Olivier—Corticolous on Alnus sp.; saxicolous. I. $5. Physcia caesia (Hoffm.) Hampe ex Furnr.—Saxico- lous. I. S4S5. Physcia millegrana Degel—Saxicolous. I. S5. Plasmatia tuckermanii (Oakes) W.L. Culb. & C.F. Culb.—Corticolous. XII. S4S5. Polysporina simplex (Taylor) Vézda—Saxicolous. VII. S4SS. Porpidia crustulata (Ach.) Hertel & Knoph—Saxico- lous. XII. S5. Protoparmelia hypotremella Herk, Spier & V. Wirth —Corticolous on dead 7: occidentalis. VII. SNR. Punctelia rudecta (Ach.) Krog—Corticolous on A. balsamea. I. SS. Pyxine sorediata (Ach.) Mont.—Saxicolous. VI. SS. Ramalina dilacerata (Hoftm.) Hoffm.—Corticolous on Picea sp. I, XIII. S4. Ramalina intermedia (Delise ex Nyl.) Nyl—Saxico- lous on a boulder. VII. S5. Rhizocarpon concentricum (Davies) Beltr—Saxico- lous. VI. SNR. Rhizocarpon timdalii thlen & Fryday—Saxicolous. VII. SNR. 2019 Rinodina freyi H. Magn.—Corticolous on a fallen P. tremuloides. XIII. S485. Scoliciosporum chlorococcum (Stenh.) Vézda—Lig- nicolous on a Pinus sp. cone. I. S5. *Sphinctrina anglica Ny|.—Lichenicolous on P. hypo- tremella on T. occidentalis. VIII. S4. tStenocybe pullatula (Ach.) Stein—Corticolous on Alnus sp. I. SU. Stereocaulon dactylophyllum Florke—Saxicolous. I. S4. Stereocaulon grande (H.Magn.) H. Magn.—Saxico- lous. XVI. S4. Stereocaulon tomentosum Fr.—Terricolous on mossy soil and on sandy soil. [X, X, XI, XVI. S4S5. Trapeliopsis granulosa (Hoftm.) Lumbsch—Terric- olous on sandy soil. HI. S5. Tuckermanopsis americana (Sprengel) Hale—Cor- ticolous on B. papyrifera and on a snag. I, XIII. S5. Tuckermanopsis sepincola (Ehrh.) Ach—Cortico- lous on L. laricina. VMI. SS. Umbilicaria deusta (L.) Baumg.—Saxicolous. VII. SS. Umbilicaria mammulata (Ach.) Tuck.—Saxicolous. Ill, VI. S485. Umbilicaria muehlenbergii (Ach.) Tuck.—Saxicolous on a boulder. V, VII. S4S5. Usnea cavernosa Tuck —Corticolous on A. balsamea and on P. mariana. I, III, VIII, XII. S485. Usnea dasopoga (Ach.) Nyl—Corticolous on a con- ifer. XII. S5. Usnea hirta (L.) Weber ex F.H. Wigg.—Corticolous on a snag. II. S5. Usnea lapponica Vain.—Corticolous on a snag. II. S4? Usnea subfloridana Stirt—Corticolous on dead P. mariana. VIII. S4S5. Vulpicida pinastri (Scop.) J.-E. Mattson & M.J. Lai— Corticolous on burned wood and on P. mariana; lignicolous on rotting wood. II, V, VHI, XV, XIV. S5. Xanthomendoza hasseana (Rasanen) Schoting, K4ar- nefelt & S.Y. Kondr—cCorticolous on P. tremul- oides. XIII. S5. Xanthoparmelia plittii (Gyeln.) Hale—Saxicolous. VII. S4S5. Xanthoparmelia viriduloumbrina (Gyeln.) Lendemer —Saxicolous. VII, XVI. SU. Discussion Sandbar Lake Provincial Park hosts a rich divers- ity of lichens, including many species that are rare in the region and province. For example, A. populorum, of which we made a single collection, is only known from nine collections in Ontario, of which only one is in northwestern Ontario (MIN 879779). This spe- DORVAL AND MCMULLIN: SANDBAR LAKE PARK LICHENS AND ALLIED FUNGI 213 cies is almost exclusively collected from the bark of Trembling Aspen, as it has been from Sandbar Lake Provincial Park. Although it is provincially tracked, this species is inconspicuous and may be overlooked in the province. Three species, B. /aurocerasi, C. parvum, and O. pseudopallescens, are also con- sidered rare or very rare in southern Ontario (Wong and Brodo 1992). Within the province, B. /aur- ocerasi has been collected mainly from the area dir- ectly surrounding the Great Lakes. Given the dis- tance of this provincial park from this location (~200 km northwest), this observation is notable. Similarly, C. chicitae is only known from near Lake Superior in Ontario (Brodo ef al. 2001). Geographic patterns found in previous reports of these species are likely affected by past collection efforts being almost exclu- sively in the area surrounding the Great Lakes, so- lidifying the need for further study in inland areas of northwestern Ontario. Overall, the species com- position of the community found at Sandbar Lake Provincial Park reflects the boreal forest in the re- gion; representative species include P. aphthosa, V. pinastri, and 17 species in the genus Cladonia (Brodo et al. 2001). The most common species in Sandbar Lake Provincial Park were B. furcellata and P. canina, which we collected six times each, at two and six collection sites, respectively. Both species are re- ported frequently from the province, as well. Although the northwest region of Ontario is known for a high diversity of lichens, previously, only one study has been geographically focussed (an intensive study within a relatively small, delineated boundary such as within a provincial park of a few hundred ha as opposed to within the entire province or not in a delineated area) in the region, on the Slate Islands (C. Wetmore unpubl. data accessed through CNALH). Focussed studies in areas with delineated boundaries, such as ours, are important for establish- ing baseline data. Lichens are effective indicators of climate change and ecological integrity and, with a sound baseline, can be used to monitor changes in the local environment over time (McMullin et a/. 2017). Coefficients generated through a Sorensen—Dice comparison showed a low level of similarity between the lichen community at Sandbar Lake Provincial Park and two other provincial parks in Ontario. Given the distance and difference in climate and for- est types among the parks, this result was not sur- prising. Awenda and Sandbanks Provincial Parks are over 1000 km southeast of Sandbar Lake Provincial Park and both border large freshwater bodies (Georgian Bay and Lake Ontario). Sandbar Lake is located within the boreal forest region, while Awenda and Sandbanks are in the Great Lakes—St. Lawrence forest region, which has a higher diversity of decidu- 214 ous trees. Both Awenda and Sandbanks Provincial Parks are smaller than our study area. Although the size of Sandbar Lake Provincial Park would likely re- late to higher diversity of lichen species, access to most parts of this park are limited, with few roads and trails outside the campground. Access by trails and roads is also limited in some areas of Sandbanks; however, the small size of the park may make it easier to sample a greater proportion of its area. In contrast, Awenda has trail networks that can be used to access most portions of the park. Nonetheless, these parks were selected because they were the nearest areas with similar search efforts (McMullin and Lewis 2014; McMullin and Lendemer 2016). Sandbar Lake Provincial Park is surrounded by resource extraction operations, especially timber har- vesting. The park, therefore, provides protection for important habitats in the area. Expansions to the park have also facilitated increased ecosystem representa- tiveness, and over time there is potential for mature or old-growth forests to develop—a habitat that is rare in this region (Ontario Parks 2012). Much of the cur- rent area of Sandbar Lake has experienced a variety of disturbances in recent history, including natural processes, such as wildfire, and anthropogenic ones, such as industrial-scale timber harvesting (Ontario Parks 2012). Forest management practices in north- ern Ontario have been shown to have direct effects on lichen community composition (e.g., herbicide contact, loss of microhabitats) and indirect effects (e.g., light exposure, tree species presence, changes in structural diversity, changes to available moisture; McMullin et a/. 2013). As a result, previous disturb- ances in the park will have influenced the lichen biota present now. Our baseline data provide the first foun- dation that can be used to acknowledge and monitor future changes to the lichen community. Our results can also be used to compare with lichen communities on lands outside the park to better understand the ef- fects on lichen biodiversity of resource extraction in the region. Author Contributions Writing — Original Draft Preparation: H.R.D. and R.T.M.; Writing — Review & Editing: R.T.M. and H.R.D.; Conceptualization: R.T.M.; Formal Analy- sis: H.R.D.; Investigation: H.R.D. and R.T.M.; Me- thodology: H.R.D. and R.T.M.; Project Administra- tion: H.R.D. and R.T.M.; Resources: H.R.D. and R.T.M. Acknowledgements We thank the Ministry of Natural Resources and Forestry, Ontario Parks, and the staff of Sandbar Lake Provincial Park for allowing this study to take place and for providing accommodations and natural herit- age information to ensure that fieldwork could be THE CANADIAN FIELD-NATURALIST Vol. 133 conducted with the greatest efficiency. Literature Cited Ahti, T., and J. Crowe. 1995. Additions to the lichens of Thunder Bay district, Ontario. Evansia 12: 21—23. Brodo, I.M., S.D. Sharnoff, and S. Sharnoff. 2001. Lichens of North America. Yale University Press, New Haven, Connecticut, USA. Crowe, J. 1994. The lichens of Thunder Bay district, On- tario, Canada. Evansia 11: 62-75. Dice, L.R. 1945. Measures of the amount of ecologic asso- ciation between species. Ecology 26: 297-302. https:// doi.org/10.2307/1932409 Esslinger, T.L. 2018. A cumulative checklist for lichen- forming, lichenicolous, and allied fungi of the conti- nental United States and Canada, version 22. Dr. Theo- dore L. Esslinger, North Dakota State University, Fargo, North Dakota, USA. Accessed 15 November 2018. https:// www.ndsu.edu/pubweb/~esslinge/chckIst/chcklst7.htm. Lavoie, M., D. Paré, N.J. Fenton, A. Groot, and K. Taylor. 2005. Paludification and management of forested peat- lands in Canada: a literature review. Environmental Reviews 13: 21-50. https://doi.org/10.1139/A05-006 McMullin, R.T., K. Drotos, D. Ireland, and H. Dorval. 2018. Diversity and conservation status of lichens and allied fungi in the Greater Toronto Area: results from four years of the Ontario BioBlitz. Canadian Field- Naturalist 132: 394-406. https://doi.org/10.22621/cfn. v13214.1997 McMullin, R.T., and J.C. Lendemer. 2016. Lichens and al- lied fungi of Awenda Provincial Park, Ontario: diversity and conservation status. American Midland Naturalist 176: 1-19. https://do1.org/10.1674/0003-0031-176.1.1 McMullin, R.T., and C.J. Lewis. 2013. New and inter- esting lichens from Ontario, Canada. Opuscula Philo- lichenum 12: 6-16. Accessed 21 September 2019. http://sweetgum.nybg.org/science/op/biblio_list.php? BucVolume_tab=12. McMullin, R.T., and C.J. Lewis. 2014. The unusual li- chens and allied fungi of Sandbanks Provincial Park, Ontario. Botany 92: 85-92. https://doi.org/10.1139/cjb- 2013-0227 McMullin, R.T., J. Maloles, and S.G. Newmaster. 2015. New and interesting lichens from Ontario, Canada II. Opuscula Philolichenum 14: 93-108. Accessed 21 Sep- tember 2019. http://sweetgum.nybg.org/science/op/biblio_ list.php?BucVolume_tab=14. McMullin, R.T., I. Thompson, and S. Newmaster. 2013. Lichen conservation in heavily managed boreal for- ests. Conservation Biology 27: 1020-1030. https://doi. org/10.1111/cobi.12094 McMullin, R.T., D. Ure, M. Smith, H. Clapp, and Y.F. Wiersma. 2017. Ten years of monitoring air quality and ecological integrity using field-identifiable lichens at Kejimkujik National Park and National Historic Site in Nova Scotia, Canada. Ecological Indicators 81: 214— 221. https://doi.org/10.1016/j.ecolind.2017.05.069 Natureserve. 2018. National and subnational conservation status definitions. NatureServe, Arlington, Virginia, USA. Accessed 16 April 2018. http://explorer.nature serve.org/nsranks.htm. 2019 Newmaster, S.G., R.J. Belland, A. Arsenault, D.H. Vitt, and T.R. Stephens. 2005. The ones we left behind: comparing plot sampling and floristic habitat sam- pling for estimating bryophyte diversity. Diversity and Distributions 11: 57-72. https://doi.org/10.1111/. 1366-9516.2005.00123.x Ontario. 2006. Provincial Parks and Conservation Reserves Act, 2006, S.O. 2006, c.12. Government of Ontario, Toronto, Ontario, Canada. Accessed 24 May 2018. https://www.ontario.ca/laws/statute/06p12. Ontario. 2015. Ontario Department of Lands and Forests: history and function. Ontario Ministry of Government and Consumer Services, Toronto, Ontario, Canada. Ac- cessed 14 April 2018. https://tinyurl.com/ybke3veu. Ontario. 2017. Ontario’s parks and protected areas. Ontario Ministry of Natural Resources and Forestry, Toronto, Ontario, Canada. Accessed 20 April 2018. https://www. ontario.ca/page/ontarios-parks-and-protected-areas. Ontario Parks. 2012. Sandbar Lake park management plan. Ministry of Natural Resources and Forestry, To- ronto, Ontario, Canada. Accessed 21 September 2019. https://www.ontario.ca/page/sandbar-lake-provincial- SUPPLEMENTARY MATERIAL: Appendix S1. Collection details of specimens. DORVAL AND MCMULLIN: SANDBAR LAKE PARK LICHENS AND ALLIED FUNGI 215 park-management-plan. Orange, P.W., and F.J. White. 2001. Microchemical Methods for the Identification of Lichens. British Lichen Society. London, United Kingdom. Sorensen, T. 1948. A method for establishing groups of equal amplitude in plant sociology based on similar- ity of species content and its application to analyses of the vegetation on Danish commons. Biologiske Skrifter Kongelige Danske Videnskabernes Selskab 5: 1-34. Ac- cessed 21 September 2019. http://www.royalacademy. dk/Publications/High/295_ S%C3%B8rensen,%20 Thorvald.pdf. Wetmore, C.M. 1981. Lichens of Voyageurs National Park, Minnesota. Bryologist. 84: 482—491. https://doi.org/10. 2307/3242555 Wong, P., and I. Brodo. 1992. The lichens of southern Ontario, Canada. Syllogeus 69. Canadian Museum of Nature, Ottawa, Ontario, Canada. Received 7 January 2019 Accepted 14 October 2019 The Canadian Field-Naturalist Do turtle warning signs reduce roadkill? Davip C. SEBURN!* and HANNAH McCurpy-ADAMs!? 'Canadian Wildlife Federation, 350 Michael Cowpland Avenue, Ottawa, Ontario K2M 2W1 Canada “Current address: Wildlife Preservation Canada, 5420 Highway 6 North, Guelph, Ontario N1H 6J2 Canada Corresponding author: davids@cwf-fcf.org Seburn, D.C., and H. McCurdy-Adams. 2019. Do turtle warning signs reduce roadkill? Canadian Field-Naturalist 133(3): 216-220. https://doi.org/10.22621/cfn.v13313.2279 Abstract Roadkill is a serious threat for many species of freshwater turtles. One of the most common road mitigation tools is wild- life warning signs to alert drivers. These warning signs have commonly been used for large mammals, although there is little evidence that they are effective in reducing roadkill. We tested the effectiveness of turtle warning signs at four known roadkill hotspots along a provincial highway in eastern Ontario and compared the results with four control sites on a nearby major road in a before-after-control-impact (BACI) study. We found 30 dead turtles in the four hotspots in 2017 before the signs were installed and 27 in 2018 after the signs were installed. The number of turtles killed on the road after the signs were installed did not change significantly (y’, = 1.1, P > 0.2). Although turtle warning signs may alert some drivers, they should not be considered a replacement for more effective road mitigation tools, such as wildlife fencing and crossing structures. Key words: Turtles; reptiles; road mitigation; wildlife signs; BACI design Introduction Roadkill is a major risk for many species of fresh- water turtles (Gibbs and Shriver 2002; Steen and Gibbs 2004; Aresco 2005). It can lead to population declines (Gibbs and Shriver 2002) or male-biased populations from disproportionate roadkill of female turtles (Steen and Gibbs 2004; Dupuis-Désormeaux et al. 2017). Turtle populations are sensitive to any threat that increases the adult mortality rate (Congdon et al. 1993; Cunnington and Brooks 1996) and are ex- tremely slow to rebound from declines (Keevil et al. 2018). As a result, roadkill can have a negative ef- fect on turtle populations near roads (Rytwinski and Fahrig 2012). Wildlife warning signs are one of the most com- monly used tools to attempt to reduce roadkill, although there is little evidence that they are effect- ive (Huijser et a/. 2015). They can take the form of standard road signs as well as enhanced road signs with flashing lights or symbols (Pojar et al. 1975; Huijser et al. 2015). Most studies on the effectiveness of wildlife warning signs have found that they do not significantly reduce roadkill (e.g., Pojar et al. 1975; Coulson 1982; Bullock et al. 2011; but see Found and Boyce 2011). Most wildlife warning sign studies have focussed on large mammals, and we are unaware of any published studies on the effectiveness of standard wildlife signs on reducing turtle roadkill. Given that all eight species of freshwater tur- tles that occur in Canada are listed as species at risk (Government of Canada 2019), it is important to assess whether turtle warning signs lead to a signifi- cant reduction in roadkill. To test the effectiveness of turtle signs (Figure 1) we examined roadkill before and after signs were installed at known hotspots in eastern Ontario. The importance of before-after-con- trol-impact (BACI) research design has been stressed in evaluating the effectiveness of road mitigation strategies (Lesbarreres and Fahrig 2012); thus, we also compared roadkill rates with those on a control road over the same period. Methods As part of a larger project on turtle conservation, road surveys were conducted in eastern Ontario in 2017, and from those surveys four hotspots were iden- tified. In spring 2018, the Ministry of Transportation installed standard turtle signs at these hotspots to help reduce roadkill. The four hotspots were located along provincial highway 15 north of Smiths Falls in Lanark County, along a section of road ~36 km in length (45.0°N, 76.0°W; Figure 2). Turtle warning signs were installed facing oncoming traffic at both ends of each hotspot. The signed road segments at each location averaged 1010 m (range 750—1675 m) in length. Daily traffic at these locations ranged from an A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 216 ©The Ottawa Field-Naturalists’ Club 2019 FicurE 1. Example of turtle sign installed by the Ministry of Transportation along provincial highway 15 in eastern Ontario in May 2018. Photo: David Seburn. annual average daily traffic of 4950 to 9400 vehicles (Ministry of Transportation 2019). The four control road segments were located along Roger Stevens Drive east of Smiths Falls in Lanark County, along a section of road ~28 km in length (Figure 2). Highway 15 and Roger Stevens Drive intersect in Smiths Falls and the two roads are less than 25 km apart at any point. Each control seg- ment was 1000 m in length and was selected based on the presence of at least four roadkilled turtles during 2017. Daily traffic in the four control segments varied Roger Stevens Drive D Smith Falls FiGure 2. Location of two roads used in test of the effect- iveness of turtle signs in eastern Ontario in 2017 and 2018. Roadkill hotspots were located along provincial highway 15 and are numbered 1-4. Turtle signs were installed at each end of all four hotspots in spring 2018. Four segments of road along Roger Stevens Drive, labelled A—D, served as control sections. SEBURN AND MCCurRDy-ADAMS: TURTLE WARNING SIGNS AND ROADKILL 217 by section, and ranged from an annual average daily traffic volume of 2860 to 3900 vehicles (roads depart- ment, Lanark County unpubl. data). Both the control and impact roads were paved, two-lane roads, with a posted speed limit of 80 km/h, although this limit was frequently exceeded by drivers (D.C.S. and H.M.-A. pers. obs.). Road surveys were usually conducted with at least two people in the vehicle, but on some occasions, only one person conducted a road survey. Surveys were conducted during the day, typically from 0900 to 1600. Roads were surveyed by driving at ~SO—60 km/h and scanning the road surface and road shoul- ders for dead turtles. The location of each roadkilled turtle was recorded using a handheld global pos- itioning system unit (eTrex or eTrex 20x, Garmin Ltd., Olathe, Kansas, USA) with a spatial accuracy of at least + 5 m. All dead turtles were removed from the road or road shoulder to prevent double counting ona subsequent survey. Road surveys were conducted ap- proximately weekly from May until early September in 2017 and 2018. Both control and impact roads were typically surveyed on the same day. The turtle warning signs were installed at the end of May 2018. Only dead turtles found in 2018 after the signs were installed were included in the analysis for both control and impact roads. Similarly, for 2017, only turtles from after the end of May were included so that the same period in both years was compared. In addition, all live turtles found on the road were ex- cluded to examine only the effect of the road signs on turtle mortality. Live turtles made up <10% of all tur- tles found on roads. This is as expected, as, if turtles successfully cross a road, they are only present for a few minutes and would only be detected if the cross- ing coincided with the survey. A chi-squared 2x2 contingency table was used to compare differences in the number of turtles in 2017 and 2018 for both roads (Minitab 8.3; Minitab Inc., State College, Pennsylvania, USA). The turtles from all four hotspots were pooled to prevent pseudorepli- cation (Hurlbert 1984) and the two years compared. Similarly the four control road segments were pooled and the two years compared. Statistical significance was defined as P < 0.05. Results Three species of turtles were found during sur- veys: Painted Turtle (Chrysemys picta), Snapping Turtle (Chelydra serpentina), and Blanding’s Turtle (Emydoidea blandingii). We found 30 dead turtles in the four hotspots in 2017 before the signs were in- stalled, and 27 in 2018 after the signs were installed. In the four control sections, we found 19 dead turtles in 2017 and 26 in 2018 after the signs were installed 218 along the other road. There was no statistically sig- nificant difference in the number of turtles found be- fore or after the signs were installed (Table 1; y?, = 1.1, P>0.2). Discussion Our road surveys likely did not detect all of the turtles killed on the roads, as they were conducted ap- proximately weekly and turtle carcasses along roads may not persist that long (Santos ef a/. 2011). In addi- tion, compared with walking surveys, driving sur- veys may fail to detect some carcasses (Santos ef al. 2016). There is no reason to assume that carcass per- sistence or detectability would have differed signifi- cantly between the two years, and survey methods and survey frequency were the same in both years. There were similar numbers of roadkilled turtles in the control road sections in both years, suggesting that roadkill numbers in the impact road sections would also have been similar in both years without the presence of any mitigation. Thus, any significant changes in roadkill numbers in the impact road sec- tions between 2017 and 2018 should be attributable to the road signs. The lack of any significant decrease in roadkill indicates that the signs were not effective. A larger sample size would have increased our chances of detecting a statistically significant difference in the amount of roadkill, if one existed. Nonetheless, a de- crease of only 10% in roadkill in 2018 from 2017 is not indicative of effective mitigation, as wildlife bar- riers and crossing structures can reduce roadkill by more than 90% (Dodd et al. 2004). Any road mitiga- tion strategy that results in only a 10% reduction in roadkill should be considered a failure. Wildlife warning signs are one of the most com- monly installed road mitigation tools (Huijser et al. 2015), likely because of their low cost. However, de- spite their wide use, there is little evidence that such warning signs are effective at reducing roadkill. Few drivers are even aware of such warning signs. In one study, only 5—10% of drivers who were stopped 200 m after passing a warning sign were able to recall the sign (Drory and Shinar 1982). For warning signs to be effective, they should result in drivers reducing their speed. Animated deer (Odocoileus spp.) warning signs have led to a reduc- TABLE 1. Results of 2x2 chi-squared contingency table comparing the observed number of dead turtles on the con- trol and impact roads, both before and after turtle signs were installed. Roadkill (expected value) Site Before After Impact (with signs) 30 (27.4) 27 (29.6) Control (no signs) 19 (21.6) 26 (23.4) THE CANADIAN FIELD-NATURALIST Vol. 133 tion in speed, but only by <5 km/h (Pojar et al. 1975). Similarly, camel (Camelus spp.) warning signs have resulted in a decline in vehicle speed, but only by 3-7 km/h (Al-Ghamdi and AlGadhi 2004). Moose (Alces americanus) warning, signs reduced driving speeds by only 1.5 km/h in a driving simulator (Jagerbrand et al. 2018). Greater speed reductions (~10 km/h) have occurred when deer carcasses were placed next to warning signs to emphasize the reality of the threat (Pojar et al. 1975). The effectiveness of animal warn- ing signs on driving speed may also decline over time as drivers become habituated to the signs (Pojar et al. 1975; Khalilikhah and Heaslip 2017). Hence, it seems that even large-mammal warning signs may only have a small effect on vehicle speeds, even though collisions can result in the injury or death of the driver (e.g., Conover et al. 1995; Niemi et al. 2017). Ultimately, the main issue is whether animal warning signs result in a reduction in collisions and roadkill. Deer crossing signs did not reduce the num- ber of deer killed in Colorado (Pojar et al. 1975), but deer collisions were reduced, at least for the first year, after warning signs were installed at known hotspots in the city of Edmonton, Alberta (Found and Boyce 2011). Temporary, flashing warning signs installed at known deer migration locations resulted in a sig- nificant reduction in vehicle collisions, but this effect lessened during the second year of the study (Sullivan et al. 2004). Warning signs were also not effective at reducing roadkill of kangaroos in Australia (Coulson 1982; Shima et al. 2018), mammals and birds along a major road in South Africa (Bullock et a/. 2011), or snakes in Illinois (Shepard et al. 2008). Enhanced warning signs may be effective under some limited circumstances. Diamond-backed Terra- pins (Malaclemys terrapin) suffer high rates of road mortality during nesting forays, which are associ- ated with diurnal high tides (Crawford et al. 2014). Flashing warning signs installed but only activated for a 2-h period each day corresponding to the diurnal high tide during the nesting season, significantly re- duced Diamond-backed Terrapin roadkill (Crawford et al. 2018). It is also possible that wildlife warning signs may be more effective along roads with a lower speed limit as speed limit is often positively correl- ated with roadkill (Farmer and Brooks 2012). Although wildlife warning signs may not sig- nificantly reduce roadkill, they can still be valuable within a comprehensive mitigation strategy for pub- lic education and sending a message that roadkill of wildlife is a serious issue. Wildlife warning signs should not replace more effective road mitigation tools such as wildlife fencing and crossing structures (e.g., Dodd et al. 2004; Aresco 2005; Baxter-Gilbert et al. 2015; Crawford et al. 2017). 2019 Acknowledgements We are grateful to all of the Canadian Wildlife Federation summer staff who assisted with road sur- veys in 2017 and 2018: Mackenzie Burns, Hannah Delion, Brandon Holden, Holly Jasmine Long, and Georgia McLay. Thanks to the Ministry of Transportation for their concern about turtle road- kill and for installing the turtle signs. Financial sup- port for this work came from the Rogers Foundation, the Canada Summer Jobs Program, and the CleanTech Internship program of Environment and Climate Change Canada. We are grateful to the reviewers for their comments, which helped us improve this paper. Literature Cited Al-Ghamdi, A.S., and S.A. AlGadhi. 2004. Warning signs as countermeasures to camel—vehicle collisions in Saudi Arabia. Accident Analysis and Prevention 36: 749-760. https://doi.org/10.1016/j.aap.2003.05.006 Aresco, M.J. 2005. Mitigation measures to reduce highway mortality of turtles and other herpetofauna at a north Florida lake. Journal of Wildlife Management 69: 549— 560. https://doi.org/10.2193/0022-541x(2005)069[0549: mmtrhm]2.0.co;2 Baxter-Gilbert, J.H., J.L. Riley, D. Lesbarréres, and J.D. Litzgus. 2015. Mitigating reptile road mortality: fence failures compromise ecopassage effectiveness. PLoS ONE 10: e0120537. https://doi.org/10.1371/journal.pone. 0120537 Bullock, K.L., G. Malan, and M.D. Pretorius. 2011. Mammal and bird road mortalities on the Upington to Twee Rivieren main road in the southern Kalahari, South Africa. African Zoology 46: 60-71. https://doi.org /10.1080/15627020.2011.11407479 Congdon, J.D., A.E. Dunham, and R.C. van Loben Sels. 1993. Delayed sexual maturity and demographics of Blanding’s turtles (Emydoidea blandingii). implications for conservation and management of long-lived organ- isms. Conservation Biology 7: 826—833. https://doi.org/ 10.1046/}.1523-1739.1993.740826.x Conover M.R., W.C. Pitt, K.K. Kessler, T.J. DuBow, and W.A. Sandborn. 1995. Review of human injuries, ill- nesses, and economic losses caused by wildlife in the United States. Wildlife Society Bulletin 23: 407—414. Coulson, G.M. 1982. Road-kills of macropods on a section of highway in central Victoria. Wildlife Research 9: 21— 26. https://doi.org/10.1071/wr9820021 Crawford, B.A., J.C. Maerz, N.P. Nibbelink, K.A. Buhlmann, T.M. Norton, and S.E. Albeke. 2014. Hot spots and hot moments of diamondback terrapin road- crossing activity. Journal of Applied Ecology 51: 367— 375. https://doi.org/10.1111/1365-2664.12195 Crawford, B.A., C.T. Moore, T.M. Norton, and J.C. Maerz. 2017. Mitigating road mortality of diamond- backed terrapins (Malaclemys terrapin) with hybrid bar- riers at crossing hot spots. Herpetological Conservation and Biology 12: 202-211. Crawford, B.A., C.T. Moore, T.M. Norton, and J.C. Maerz. 2018. Integrated analysis for population esti- SEBURN AND MCCurDy-ADAMS: TURTLE WARNING SIGNS AND ROADKILL 219 mation, management impact evaluation, and decision- making for a declining species. Biological Conservation 222: 33-43. https://do1.org/10.1016/j.biocon.2018.03.023 Cunnington, D.C., and R.J. Brooks. 1996. Bet-hedging theory and eigenelasticity: a comparison of the life hist- ories of loggerhead sea turtles (Caretta caretta) and snap- ping turtles (Chelydra serpentina). Canadian Journal of Zoology 74: 291-296. https://doi.org/10.1139/z96-036 Dodd, Jr., C.K., W.J. Barichivich, and L.L. Smith. 2004. Effectiveness of a barrier wall and culverts in reducing wildlife mortality on a heavily traveled highway in Florida. Biological Conservation 118: 619-631. https:// doi.org/10.1016/j.biocon.2003.10.011 Drory, A., and D. Shinar. 1982. The effects of roadway environment and fatigue on sign perception. Journal of Safety Research 13: 25-32. https://doi.org/10.1016/ 0022-4375(82)90015-9 Dupuis-Désormeaux, M., V. D’ Elia, C. Cook, J. Pearson, V. Adhikari, and S.E. MacDonald. 2017. Remarkable male bias in a population of midland painted tur- tles (Chrysemys picta marginata) in Ontario, Canada. Herpetological Conservation and Biology 12: 225-232. Farmer, R.G., and R.J. Brooks. 2012. Integrated risk fac- tors for vertebrate roadkill in southern Ontario. Journal of Wildlife Management 76: 1215-1224. https://doi.org/ 10.1002/jwmg.358 Found, R., and M.S. Boyce. 2011. Warning signs mitigate deer—vehicle collisions in an urban area. Wildlife Society Bulletin 35: 291-295. https://doi.org/10.1002/wsb.12 Gibbs, J.P., and W.G. Shriver. 2002. Estimating the effects of road mortality on turtle populations. Conserva- tion Biology 16: 1647-1652. https://doi.org/10.1046/j.15 23-1739.2002.01215.x Government of Canada. 2019. Species at risk public regis- try. Government of Canada, Ottawa, Ontario, Canada. Accessed 4 February 2019. https://www.canada.ca/en/ environment-climate-change/services/species-risk- public-registry.html. Huijser, M.P., C. Mosler-Berger, M. Olsson, and M. Strein. 2015. Wildlife warning signs and animal de- tection systems aimed at reducing wildlife-vehicle col- lisions. Pages 198-212 in Handbook of Road Ecology. Edited by R. van der Ree, D.J. Smith, and C. Grilo. John Wiley & Sons, Chichester, United Kingdom. https://doi. org/10.1002/9781118568170.ch24 Hurlbert, S.H. 1984. Pseudoreplication and design of eco- logical field experiments. Ecological Monographs 54: 187-211. https://doi.org/10.2307/1942661 Jaigerbrand, A.K., H. Antonson, and C. Ahlstrém. 2018. Speed reduction effects over distance of animal—vehicle collision countermeasures—a driving simulator study. European Transport Research Review 10: 40. https:// doi.org/10.1186/s12544-018-0314-8 Keevil, M.G., R.J. Brooks, and J.D. Litzgus. 2018. Post- catastrophe patterns of abundance and survival reveal no evidence of population recovery in a long-lived animal. Ecosphere 9: e02396. https://doi.org/10.1002/ecs2.2396 Khalilikhah, M., and K. Heaslip. 2017. Improvement of the performance of animal crossing warning signs. Journal of Safety Research 62: 1-12. https://doi.org/10. 1016/j.jsr.2017.04.003 220 Lesbarréres, D., and L. Fahrig. 2012. Measures to reduce population fragmentation by roads: what has worked and how do we know? Trends in Ecology & Evolution 27: 374-380. https://doi.org/10.1016/j.tree.2012.01.015 Ministry of Transportation. 2019. Ontario provincial highways traffic volumes on demand. Ministry of Transportation, Toronto, Ontario, Canada. Accessed 20 January 2019. http://www.ragsa.mto.gov.on.ca/tech pubs/TrafficVolumes.nsf/tvweb?OpenForm& Seq=l. Niemi, M., C.M. Rolandsen, W. Neumann, T. Kukko, R. Tiilikainen, J. Pusenius, E.J. Solberg, and G. Erics- son. 2017. Temporal patterns of moose—vehicle colli- sions with and without personal injuries. Accident Analysis & Prevention 98: 167-173. https://doi.org/10. 1016/j.aap.2016.09.024 Pojar, T.M., R.A. Proscence, D.F. Reed, and T.N. Woodard. 1975. Effectiveness of lighted, animated deer crossing sign. Journal of Wildlife Management 39: 87— 91. https://doi.org/10.2307/3800469 Rytwinski, T., and L. Fahrig. 2012. Do species life history traits explain population responses to roads? A meta- analysis. Biological Conservation 147: 87—98. https:// doi.org/10.1016/j.biocon.2011.11.023 Santos, R.A.L., S.M. Santos, M. Santos-Reis, A.P. de Figueiredo, A. Bager, L.M. Aguiar, and F. Ascensao. 2016. Carcass persistence and detectability: reducing the uncertainty surrounding wildlife—vehicle collision surveys. PLoS ONE 11: e0165608. https://doi.org/10.13 71/journal.pone.0165608 THE CANADIAN FIELD-NATURALIST Vol. 133 Santos, S.M., F. Carvalho, and A. Mira. 2011. How long do the dead survive on the road? Carcass persistence probability and implications for road-kill monitoring surveys. PLoS ONE 6: e25383. https://doi.org/10.1371/ journal .pone.0025383 Shepard, D.B., M.J. Dreslik, B.C. Jellen, and C.A. Phillips. 2008. Reptile road mortality around an oasis in the Illinois corn desert with emphasis on the en- dangered Eastern Massasauga. Copeia 2008: 350-359. https://doi.org/10.1643/CE-06-276 Shima, A.L., D.S. Gillieson, G.M. Crowley, R.G. Dwyer, and L. Berger. 2018. Factors affecting the mortality of Lumholtz’s tree-kangaroo (Dendrolagus lumholtzi) by vehicle strike. Wildlife Research 45: 559-569. https:// doi.org/10.1071/WR17143 Steen, D.A., and J.P. Gibbs. 2004. Effects of roads on the structure of freshwater turtle populations. Conservation Biology 18: 1143-1148. https://doi.org/10.1111/.1523-17 39.2004.00240.x Sullivan, T.L., A.F. Williams, T.A. Messmer, L.A. Hel- linga, and S.Y. Kyrychenko. 2004. Effectiveness of temporary warning signs in reducing deer—vehicle col- lisions during mule deer migrations. Wildlife So- ciety Bulletin 32: 907-915. https://do1.org/10.2193/00 91-7648(2004)032[0907:eotwsi]2.0.co;2 Received 13 May 2019 Accepted 20 December 2019 The Canadian Field-Naturalist Surveys for terrestrial gastropods in the Kootenay region of British Columbia, with new records and range extensions KRISTIINA OvASKA!*, LENNART SOPUCK!, and JENNIFER HERON? 'Biolinx Environmental Research Ltd., 1759 Colburne Place, North Saanich, British Columbia V8L 5A2 Canada *Ministry of Environment and Climate Change Strategy, Conservation Science Section, Suite 200, 10428 153rd Street, Surrey, British Columbia V3R 1E1 Canada “Corresponding author: ke.ovaska@gmail.com Ovaska, K., L. Sopuck, and J. Heron. 2019. Surveys for terrestrial gastropods in the Kootenay region of British Columbia, with new records and range extensions. Canadian Field-Naturalist 133(3): 221-234. https://do1.org/10.22621/cfn. v13313.2287 Abstract The northern Columbia River basin, extending from the Kootenay region in British Columbia southward to the Idaho pan- handle and northwestern Montana, contains a unique terrestrial gastropod fauna, but in Canada few surveys have specifi- cally targetted this group. Here we report on terrestrial gastropods detected during surveys of 314 sites distributed in five biogeoclimatic zones across the Kootenay region. The surveys occurred on 65 days over seven years from 2007 to 2015, usually in September—October. We detected 45 taxa identified to species, belonging to 33 genera; micro-snails of the genus Vertigo (apart from Vertigo ovata) were combined into a single category, and snails belonging to Succineidae were not identified further. Regionally endemic species previously reported from the region included Western Banded Tigersnail (Anguispira kochi occidentalis), Coeur @Alene Oregonian (Cryptomastix mullani), Rocky Mountainsnail (Oreohelix stri- gosa), Subalpine Mountainsnail (Oreohelix subrudis), and Pale Jumping-slug (Hemphillia camelus), which was widespread across the region. Magnum Mantleslug (Magnipelta mycophaga), the distribution of which extends beyond the Kootenay region, was detected at several widely spaced sites. Two species new to Canada were detected, Pygmy Slug (Kootenaia burkei) and Sheathed Slug (Zacoleus idahoensis), both of which were subsequently assessed to be of conservation concern both provincially and nationally. Other notable observations included the detection of Fir Pinwheel (Radiodiscus abietum), a regional endemic, which has been previously reported only once, and three species common in coastal forests but not pre- viously reported from the region: Pacific Banana Slug (Ariolimax columbianus), Robust Lancetooth (Haplotrema vancou- verense), and Northwest Hesperian (Vespericola columbianus). Further surveys, especially at higher elevations, may reveal other additional or unusual species. Key words: Terrestrial gastropods; new distribution records; Kootenays; inventory Introduction Mesic forests of the northern Columbia River basin support many unique plants and animals and species with vicarious distributions, separated from their Pacific coastal counterparts by 300 km or more of arid landscapes (Brunsfeld et al. 2001). This unique area extends from southeastern British Columbia (BC) and northeastern Washington southward through the Idaho Panhandle into northwestern Montana. In BC, it encompasses the Kootenay region, which supports a diverse gastropod fauna, including species that are found nowhere else in Canada (Forsyth 1999, 2004). Few studies have specifically targetted this group, and until recently our knowledge of it was based on brief historical accounts, records in Pilsbry’s (1939, 1940, 1946, 1948) monograph, and largely serendipit- ous observations (reviewed by Forsyth 1999). Recent 221 targetted surveys include the Royal British Columbia Museum’s Living Landscape expedition (Forsyth 1999) and surveys by Nekola ef al. (2011) 1n the cen- tral Selkirk Mountains and their vicinity in sup- port of a proposed Selkirk Mountains Caribou Park. Increased survey efforts in this biologically rich area continue to provide new records and document spe- cies new to the province. Here we report on surveys targetting terrestrial gastropods in southeastern BC during seven annual surveys from 2007 to 2015 (no surveys were con- ducted in 2011-2013), including documentation of two species of slugs new to Canada. The surveys were in support of conservation assessments by the province of BC and by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) and focussed on species deemed to be rare or at risk. ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 222 Focal species initially included the snails Western Banded Tigersnail (Anguispira kochi occidentalis), Coeur d’Alene Oregonian (Cryptomastix mullani), and mountainsnail (Oreohelix) species, and the slugs Magnum Mantleslug (Magnipelta mycophaga) and Pale Jumping-slug (Hemphillia camelus). Two spe- cies, Pygmy Slug (Kootenaia burkei) and Sheathed Slug (Zacoleus idahoensis), were added after their discovery as part of this study in 2007 and 2009, re- spectively. The primary objective was to clarify dis- tributions of the focal species. A secondary objective was to investigate the presence of possible undocu- mented species of the northern Columbia basin fauna, the distributions of which may extend northward across the international border into Canada. Study Area This study was conducted in the Kootenay region of southeastern BC, bounded by the Rocky Mountains to the east, the Canada—United States (USA) border if \ } a f | 4 } . Fa BRITISH COLUMBIA’. é i f*Nelsoni e~ ~- f pees) ofS * CANADA / \ THE CANADIAN FIELD-NATURALIST Vol. 133 to the south, Shuswap/Okanagan Highlands to the west, and the 51.6° latitude to the north (Figure 1). The area consists of a series of rugged northwest— southeast oriented mountain ranges (Rocky, Purcell, Selkirk, and Monashee), separated by large valleys, rivers, and lakes. The varied terrain and climate, which can change across short distances, result in a diversity of ecosystems, which have strong influences on terrestrial gastropod distributions. The study area overlaps five of 14 biogeoclimatic zones in BC (BECP n.d.). A zone is classified accord- ing to the unique assemblage and distribution of cli- max and late-seral plant communities, energy flow, nutrient-cycling and soils, topography, and regional climate of a geographic area. Biogeoclimatic zones of the study area include high elevation Interior Mountain-heather Alpine (IMA) and Engelmann Spruce (Picea engelmannii Engelmann)—Subalpine Fir (Abies lasiocarpa (Hooker) Nuttall) (ESSF) zones; mid-elevation Interior Cedar-Hemlock (ICH) and e Survey Sites Study area boundary | . ‘, | Invermere, Figure 1. Location of the study area and sites surveyed in 2007-2015 in southeastern British Columbia. 2019 Montane Spruce (MS) zones; low-elevation Interior Douglas Fir (Pseudotsuga menziesii (Mirbel) Franco) (IDF) zone; and Ponderosa Pine (Pinus ponderosa Douglas ex Lawson & C. Lawson) zone (MacKillop et al. 2018; BECP n.d.). Of the forested zones, the ESSF and ICH zones tend to have wet climates, whereas the MS and IDF zones tend to be dry. Moist Engelmann Spruce—Subalpine Fir forests dominate the higher elevations, White Spruce (Picea glauca (Moench) Voss), Western Hemlock (7suga hetero- phylla (Rafinesque) Sargent), and Western Redcedar (Thuja plicata Donn ex D. Don) forests dominate the wetter low- to mid-elevations; Lodge-pole Pine (Pinus contorta Douglas ex Loudon), Western Larch (Larix occidentalis Nuttall), and Douglas Fir forests occur on the drier mountain slopes; Ponderosa Pine and grass- lands occur in the dry, low-elevation valley bottoms. In recent years, logging, wildfires, hydroelectric reservoirs, and Mountain Pine Beetle (Dendroctonus ponderosae) epidemics have disturbed large areas of forest in the study area. The effects of cattle ranch- ing are localized, occurring mainly in grassland and open forest habitats at low elevations. Human de- velopments are relatively sparse, with settlements and farming occurring mainly in low-elevation river valleys, such as along the Columbia and Kootenay rivers. Several federal and provincial parks contain large areas of relatively undisturbed forest, but these are mostly restricted to higher elevations. Methods Survey sites and effort We surveyed 314 sites, which were at least 500 m apart and distributed across the study area (Figure 1, Appendix S1); geopositions were recorded with a handheld global positioning system unit (GPSMAP 60Cx; Garmin, Olathe, Kansas, USA). Survey efforts focussed on BC Crown forestry lands, which were accessed through logging roads that crisscrossed the area. The sites also included provincial parks (n = 5), national parks (n = 4), Ktunaxa First Nation reserve lands (n = 17), municipal lands (n = 6), and private lands used for forestry, ranching, or recreation (n = 22). Most sites were searched only once, but repeated surveys were conducted at eight sites. The total time spent intensively searching for gastropods was 347 person-hours, with the median survey time one hour per site during 65 days over a seven-year period. Survey dates, conditions, and methods The surveys took place on the following dates: 10— 16 July and 22—27 September 2007; 3-8 September and 4—7 October 2008; 6—9 October 2009; 20 July and 6-16 September 2010; 20-25 September 2013: 15-29 September 2014; and 20-25 September 2015. It rained during 16.5% of the surveys, and the median OVASKA ET AL.: KOOTENAY GASTROPOD DISTRIBUTION 223 ambient temperature (measured at ground level at the start of the survey) was 11°C in September—October and 24°C in July. To locate gastropods, usually two observers (K.O. and L.S.) walked along meandering transects through habitat deemed suitable for gastropods and examined decaying logs, piles of bark, stumps, rocks, other moist refuges, and accumulations of leaf litter, fo- cussing on points where concentrations of such fea- tures were present. Most surveys took place during daylight hours, but a few (7 = 5) took place on wet nights after dark to detect gastropods active on the surface. At night, we used high-watt flashlights to scan the ground surface and tree trunks while walk- ing along trails or traversing suitable habitat and/or driving slowly along side-roads using fog lights to il- luminate the road surface. Identification and vouchers We identified all gastropods found, in some cases only to genus (such as micro-snails of Vertigo), based on external characteristics. Identification was usually done in the field, but we collected samples of small snails and confirmed identification in the laboratory of the Royal British Columbia Museum (RBCM) using a dissecting microscope. Three specimens of Hemphillia were sent to Lyle Chichester to confirm identity through examination of distal reproduct- ive anatomy; several species of Hemphillia occur south of the border in the USA and have not been re- ported from Canada. Identification was based on de- scriptions in Pilsbry (1940, 1948), Forsyth (2004), and Burke (2013). Nomenclature for species fol- lowed Forsyth (2004) and, for families, Bouchet et al. (2017). Voucher specimens were deposited in col- lections at the RBCM (Appendix S2); photographic vouchers were retained in personal collections by the authors. Results The surveys resulted in the detection of 45 taxa identified to species, belonging to 33 genera (Table 1). Micro-snails of the genus Vertigo (apart from Ovate Vertigo, Vertigo ovata) were combined into a single category, which included nominal taxa of Vertigo columbiana, Vertigo cristata, Vertigo gouldii, Ver- tigo modesta, and possibly other taxa (vouchers at RBCM). Collections of Vertigo from British Colum- bia await re-examination in light of the recent revi- sion of the genus (Nekola e¢ a/. 2018). Snails of the family Succineidae were not identified below this level because of complications associated with iden- tification based on shell morphology. The most commonly encountered taxa were Brown Hive (Euconulus fulvus, 57% of sites surveyed), Forest Disc (Discus whitneyi, 45%), Vertigo species group Vol. 133 197 OLTZ O€ BC 6£ TE Il 90 Ill It ical N 5 91 67 < oe : tl TE << Z rl 90 Q — aa a 4 6b7 €8 a < Z | @ Oreohelix spp. | British Coluitibia, V4) tor a ' Pe Alberta r ne wee \ s< 4° | © Cryptomastix mullani | ‘ I mea : »< 4° | @ Vespericola columbianus ah, FIGURE 2. Continued. 232 THE CANADIAN FIELD-NATURALIST Vol. 133 a bth a - - % en ee” Seeee ye ‘en | FiGure 3. Selected species detected during surveys in the Kootenay region, 2007-2015. a. Sheathed Slug, Zacoleus idaho- ensis (September 2013, site 158, 25 mm, in situ); b. Pygmy Slug, Kootenaia burkei (September 2014, site 230, 12 mm, in situ); c. Pale Jumping-slug, Hemphillia camelus (September 2014, site 83, juvenile 35 mm, in situ); d. Magnum Mantleslug, Magnipelta mycophaga (July 2007, site 267, 65 mm, in situ), e. Coeur d’Alene Oregonian, Cryptomastix mullani (September 2007, site 142, 15 mm); f. Rocky Mountainsnail, Oreohelix strigosa (September 2007, site 142, 20 mm); g. Western Banded Tigersnail, Anguispira kochi occidentalis (3 September 2008, site 78, 25 mm, in situ); h. Idaho Forestsnail, A/logona ptychophora (September 2010, site 14, 22 mm). Size is extended length for slugs, shell width for snails. Photos: Kristiina Ovaska. 2019 Ficure 4. a. Fir Pinwheel, Radiodiscus abietum (23 September 2013, site 85) from the Kootenay region. b. Detail of shell OVASKA ET AL.: KOOTENAY GASTROPOD DISTRIBUTION 233 We. with diagnostic fine spiral striae. Photos: Kristiina Ovaska and Heidi Gartner. of these species represents natural distribution or re- sulted from recent introductions by humans from the coast remains unknown. Non-native species encountered during the sur- veys include the snails Iroquois Vallonia (Vallonia ex- centrica) at one site, adjacent to hot springs at a recrea- tional site, and Grovesnail (Cepaea nemoralis) at three lower elevation sites, two of them in provincial parks. Introduced slugs of three families (Agriolimacidae, Arionidae, Limacidae) were widespread within the study area and occurred at 51 sites (16.2%), distrib- uted across all five biogeoclimatic zones sampled and often associated with human-modified habitats. This study contributes to a growing body of in- formation on distributions of terrestrial gastropods of southeastern BC. Most of the sampling sites were in the ICH biogeoclimatic zone, which was expected to provide suitable habitat for most of the focal species. Higher elevation sites in the ESSR and MS zones, in particular, merit further surveys, but pose logistic challenges because of difficulties of access. Yet, these habitats may contain unique species and faunas that might be particularly vulnerable to climate change 1m- pacts, as reported in other areas (Miller et a/. 2009). Author Contributions Writing — Original draft: K.O. and L.S.; Writing — Review & editing: K.O., L.S., and J.H.; Conceptu- alization: K.O., L.S., and J.H.; Surveys & specimen identification: K.O. and L.S.; Logistics & landowner contacts: J.H.; Funding acquisition: J.H. Acknowledgements We are grateful to all individuals and organizations who partnered with us and helped make this project possible, including landholders and managers who al- lowed us on their lands. We particularly acknowledge Ktunaxa First Nations representatives on Aqam and Akisqnuk Reserves, Ted Antifeau, Lindsay Anderson, Kari Stewart-Smith, Ian Adams, Darren Komonosk1, Zehnder Farms, Wayne Stetski, and Nancy Newhouse, for their assistance in selecting and accessing survey sites. Laura Parkinson, Kyle Shottanana, and Christian Engelstoft provided able field assistance during parts of the study. Lyle Chichester performed dissections of Hemphillia specimens. Heidi Gartner diligently and expediently catalogued voucher specimens at Royal British Columbia Museum. Robert Forsyth and Dwayne Lepitzki provided helpful comments, which greatly improved the manuscript. Funding was pro- vided by the BC Ministry of Environment and Climate Change Strategy and BC Ministry of Forests, Lands, Natural Resource Operations and Rural Development. Additional funding came from Environment Canada through the Habitat Stewardship Program for surveys in 2010 and through the Committee on the Status of Endangered Wildlife in Canada for field verification associated with the preparation of status reports in 2013 and 2014. The Royal British Columbia Museum and BC Parks Enhancement Fund contributed to travel expenses associated with the surveys. Literature Cited BECP (Biogeoclimatic Ecosystem Classification Pro- gram). n.d. Zone and provincial classification reports. Research Branch, Ministry of Forestry, Victoria, British Columbia, Canada. Accessed April 2019. https://www. for.gov.bc.ca/HRE/becweb/resources/classification reports/provincial/index. html. Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Niitzel, P. Parkhaev, M. Schrédl, and E.E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran fami- lies. Malacologia 61: 1-526. https://doi.org/10.4002/040. 061.0201 Brunsfeld, S.J., J. Sullivan, D.E. Soltis, and P.S. Soltis. 2001. Comparative phylogeography of northwestern North America: a synthesis. Pages 319-339 in Integrating Ecology and Evolution in a Spatial Context. Edited by J. Silvertown and J. Antonovics. Blackwell Science, Ox- ford, United Kingdom. 234 Burke, T. 2013. Snails and Slugs of the Pacific Northwest. Oregon State University Press, Corvallis, Oregon, USA. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2016a. COSEWIC assessment and status report on the Pygmy Slug Kootenaia burkei in Canada. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2016b. COSEWIC assessment and status report on the Sheathed Slug Zacoleus idahoensis in Canada. COSEWIC, Ottawa, Ontario, Canada. Forsyth, R.G. 1999. Terrestrial malacology in the Colum- bia Basin, British Columbia. Living Landscape proj- ect, Royal British Columbia Museum, Victoria, British Columbia. Accessed March 2019. https://royalbcmuseum. bc.ca/exhibits/living-landscapes/cbasin/molluscs/intro duction2.html. Forsyth, R.G. 2004. Land Snails of British Columbia. Royal British Columbia Museum, Victoria, British Columbia, Canada. Horsakova, V., J.C. Nekola, and M. Horsak. 2019. When is a “cryptic” species not a cryptic species: a consider- ation from the holarctic micro-landsnail genus Euconu- lus (Gastropoda: Stylommatophora). Molecular Phylo- genetics and Evolution 132: 307-320. https://doi.org/10. 1016/j.ympev.2018.12.004 Leonard, W.P., L. Chichester, J. Baugh, and T. Wilke. 2003. Kootenaia burkei, a new genus and species of slug from northern Idaho, United States (Gastropoda: Pulmonata: Arionidae). Zootaxa 355: 1—16. https://doi. org/10.11646/zootaxa.355.1.1 MackKillop, D., A. Ehman, K. Iverson, and E. McKenzie. 2018. A Field Guide to Ecosystem Classification and Identification for Southeast British Columbia: the East Kootenay. Land Management Handbook 71. Govern- ment of British Columbia, Victoria, British Columbia, Canada. Accessed 14 January 2020. https://www.for. gov.bc.ca/hfd/pubs/docs/Imh/LMH71.pdf. SUPPLEMENTARY MATERIAL: THE CANADIAN FIELD-NATURALIST Vol. 133 Miller, J., C. Bassler, C. Strétz, B. Klécking, and R. Brandl. 2009. Molluscs and climate warming in a low mountain range national park. Malacologia 51: 89-109. https://doi.org/10.4002/040.051.0106 Nekola, J.C., S. Chiba, B.F. Coles, C.A. Drost, T. von Proschwitz, and M. Horsak. 2018. A phylogenetic over- view of the genus Vertigo O. F. Miller, 1773 (Gastropoda: Pulmonata: Pupillidae: Vertigininae). Malacologia 61: 21-161. https://doi.org/10.4002/040.062.0104 Nekola, J., B. Coles, and M. Horsak. 2011. Land snail bio- diversity assessment for the Selkirk Mountains Park re- gion in southeastern British Columbia. Unpublished re- port. Valhalla Wilderness Society, New Denver, British Columbia, Canada. Pilsbry, H.A. 1939. Land Mollusca of North America (north of Mexico). Monograph 3, 1(1): i-xvii, 1-573, i-ix. Academy of Natural Sciences of Philadelphia, Philadelphia, USA. Pilsbry, H.A. 1940. Land Mollusca of North America (north of Mexico). Monograph 3, 1(2): 575-994, i-1x. Academy of Natural Sciences of Philadelphia, Philadelphia, USA. Pilsbry, H.A. 1946. Land Mollusca of North America (north of Mexico). Monograph 3, 2(1): i-iv, 1-520, 1-1x, fron- tispiece. Academy of Natural Sciences of Philadelphia, Philadelphia, USA. Pilsbry, H.A. 1948. Land Mollusca of North America (north of Mexico). Monograph 3, 2(2): i—xlvui, 521-1113. Academy of Natural Sciences of Philadelphia, Phila- delphia, USA. Reise, H., J.M.C. Hutchinson, S. Schunack, and B. Schlitt. 2011. Deroceras panormitanum and congeners from Malta and Sicily, with a redescription of the wide- spread pest slug as Deroceras invadens n. sp. Folia Malacologica 19: 201-233. https://doi.org/10.2478/v101 25-011-0028-1 Received 15 May 2019 Accepted 27 December 2019 Appendix S1. List of sites surveyed for terrestrial gastropods in the Kootenay region, British Columbia, 2007-2015. Appendix S82. List of voucher specimens deposited to collections at Royal British Columbia Museum, Victoria, British Columbia. The Canadian Field-Naturalist Conspecific cues encourage Barn Swallow (Hirundo rustica erythrogaster) prospecting, but not nesting, at new nesting structures ANDREW J. CAMPOMIZzzI""*, ZOE M. LEBRUN-SOUTHCOTT', and KRISTYN RICHARDSON? 'Bird Ecology and Conservation Ontario, 114 Vaughan Road, Suite 307, Toronto, Ontario M6C 2M1 Canada ‘Bird Studies Canada, P.O. Box 160, 115 Front Street, Port Rowan, Ontario NOE 1MO Canada Corresponding author: andy@beco-birds.org Campomizzi, A.J., Z.M. Lebrun-Southcott, and K. Richardson. 2019. Conspecific cues encourage Barn Swallow (Hirundo rustica erythrogaster) prospecting, but not nesting, at new nesting structures. Canadian Field-Naturalist 133(3): 235-245. https://doi.org/10.22621/cfn.v13313.2233 Abstract Shed-like structures are being built to provide Barn Swallow (Hirundo rustica erythrogaster) nesting habitat in response to population declines. However, Barn Swallow use of these structures is unavailable in the literature. We conducted three manipulative experiments to test if adding conspecific cues (i.e., vocalizations and decoys) to newly-built structures affected prospecting visits by Barn Swallows (1) during pre-breeding, (2) during post-breeding, and (3) during or after broadcasts of vocalizations compared to before broadcasts. Additionally, we monitored nesting following pre- and post-breeding cues. We built one nesting structure with and one without conspecific cues at each of 10 study sites in southern Ontario, Canada where nesting habitat was recently lost. We detected about twice as many Barn Swallows immediately after conspecific broadcasts compared to before. We did not find substantial differences in abundance and interactions with new nesting structures for other comparisons involving conspecific cues. Following pre-breeding cues at 10 sites, six nests were built in three of 10 structures treated with conspecific cues, compared to five nests in four of 10 structures without cues. In the sub- sequent breeding season following post-breeding cues at eight sites, four nests were built in two of eight structures treated with conspecific cues, compared to four nests in three of eight structures without cues. Conspecific vocalizations appeared to increase prospecting behaviour, but not the number of nests, at new nesting structures. The paucity of nests on new structures suggests that building shed-like structures may not be an effective method of mitigating loss of nesting habitat. Key words: Aerial insectivore; conspecific attraction; habitat restoration; nesting habitat; Ontario; prospecting; public in- formation; social cues Introduction Social cues provide inadvertent information from an animal engaged in its activities and convey infor- mation about a species’ habitat that can be observed by other animals (Danchin ef a/. 2004). There is em- pirical evidence that territorial and colonial-nesting migratory birds can be attracted to nesting areas by experiments that introduce conspecific cues (e.g., Ahlering and Faaborg 2006; Hahn and Silverman 2006; Farrell et al. 2012). Thus, conspecific cues have potential application in conservation of various species to attract nesting birds to restored or protected habitat. Previous research has shown that migratory song- birds can be attracted with conspecific cues to loca- tions that do not provide typical conditions of a spe- cies’ breeding habitat (Nocera et al. 2006). Such circumstances could produce an ecological trap, in which individuals identify a location as breeding habitat because of artificial conspecific cues, but the location negatively affects breeding (Schlaepfer et al. 2002). Alternatively, if conspecific cues increase the size of a breeding colony, there may be increases in re- productive success through various mechanisms such as predator dilution, group vigilance, or extra-pair pa- ternity (Parrish and Edelstein-Keshet 1999; Danchin et al. 2000). Prospecting behaviour to visit potential nesting areas can occur before, during, or after the breeding season for adults and late in the breeding season for hatch-year birds, after they are independ- ent from parents (Reed et al. 1999; Doligez et al. 2004; Ward 2005). Conspecific cues could potentially aid conserv- ation of Barn Swallow (Hirundo rustica eryth- rogaster), an aerial insectivore. Populations of birds that forage on flying insects while in flight have de- clined markedly over the last several decades in North America (Nebel et a/. 2010; Sauer et al. 2013, 2017; Smith et al. 2015), leading to conservation concern and recovery efforts. These aerial insecti- vores include species from four taxonomic families: 235 ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 236 nighthawks and nightjars (Caprimulgidae), swifts (Apodidae), tyrant flycatchers (Tyrannidae), and swallows (Hirundinidae). Barn Swallow is the most abundant and widespread swallow species world- wide (Brown and Brown 1999) and considered least concern by the International Union for Conservation of Nature (BirdLife International 2016). Although still common in many rural areas, the Barn Swallow population declined by 80% in Canada and 66% in Ontario between 1970 and 2012 (Heagy ef al. 2014), leading to its listing as threatened by the govern- ments of Canada (Government of Canada 2017) and Ontario (MECP 2012). The reasons for its popula- tion decline are not well understood, but potential causes include: (1) loss of nesting habitat; (2) loss or degradation of foraging habitat impacting prey in- sects; (3) climate change and mortality from extreme cold weather events on breeding grounds; (4) pollu- tion and pesticides; (5) reduced fecundity because of predation, ectoparasites, and persecution by humans; and (6) loss of, and human disturbance at, roosts (COSEWIC 2011; Heagy et al. 2014). In Ontario, Barn Swallows breed predominantly south of the Canadian Shield in the Mixedwood Plains ecozone (Lepage 2007). They breed in vari- ous non-forested areas and are typically associated with human-built structures that provide nesting op- portunities, such as barns, bridges, and sheds (Brown and Brown 1999). Recently, structures specifically designed as Barn Swallow nesting habitat have been built. In Ontario, most nesting structures are built as mitigation for habitat loss as required by the Ontario Ministry of Environment, Conservation and Parks (e.g., due to building or bridge demolition or reno- vation; MECP 2013); others are built to provide new nesting habitat. There are reports providing informa- tion about nesting in these structures (e.g., Heagy et al. 2014; K.R. unpubl. data), but we were unable to find information in the literature. Overall, the fre- quency of use of these structures for nesting by Barn Swallows is unclear because few results are avail- able. Although loss of nesting habitat is only one po- tential factor contributing to Barn Swallow popula- tion declines, it is important for conservation efforts that address habitat loss to make the best use of funds and opportunities by providing nesting habitat that is most likely to be used productively by the species. Barn Swallows often nest colonially (Brown and Brown 1999), suggesting they may use conspecific cues (e.g., the presence of adults at a nesting struc- ture) when selecting nest sites. There is some evi- dence of success in using conspecific cues to attract Purple Martin (Progne subis, another swallow spe- cies) to nest in previously unoccupied martin houses (Kostka 2000). We hypothesized that introducing THE CANADIAN FIELD-NATURALIST Vol. 133 conspecific cues (i.e., decoys and vocalizations), to make it appear that a structure was already being used by nesting Barn Swallows, would increase the likelihood of nesting at a new structure. There is evidence that the presence of old nests influences the use of a nest site by Barn Swallows. Safran (2004) found that removing old nests before birds arrived on the breeding grounds reduced the proportion of immigrant female Barn Swallows at sites in New York. Additionally, birds that reused old nests had higher seasonal fecundity than those that built new nests (Safran 2004). Settlement patterns of females breeding at a site for the first time were as- sociated with the number of old nests, rather than the prevalence of colourful males or opportunities for extra-pair copulations (Safran 2007). Ringhofer and Hasegawa (2014) found that the number of old un- damaged nests was associated with the spring arrival date of male Barn Swallows at nest sites. Thus, both conspecific cues and the presence of old nests likely influence the use of nest sites by Barn Swallows. Barn Swallows likely gather information about numerous potential nest sites before selecting one for breeding, as occurs with other migratory songbirds (Brown and Brown 1999; Reed et al. 1999). Visits to nesting structures by Barn Swallows likely indi- cate that individuals are gathering information about the structure for potential future use for nesting. Attracting Barn Swallows to new nesting structures could positively or negatively influence reproduct- ive success. Breeding success of Barn Swallows can decrease with increasing number of proximate nests (Shields and Crook 1987); however, it is unknown how the use of conspecific cues might influence the reproductive success of the species. Our goal was to test the influence of conspecific cues on the use of newly built nesting structures by Barn Swallows to inform conservation efforts that include the creation of nesting structures. In ex- periment one, we monitored pre-breeding prospect- ing visits by Barn Swallows to assess if birds visited structures with conspecific cues (i.e., vocalizations and decoys) more frequently than structures without cues. In experiment two, we monitored post-breeding prospecting visits to assess if birds visited structures with conspecific cues more frequently than struc- tures without cues. In experiment three, we further investigated the immediate response to conspecific vocalizations by assessing if Barn Swallows visited structures more frequently during or after vocal- ization broadcasts compared to before broadcasts. Additionally, we monitored nesting following pre- and post-breeding cues to assess if conspecific cues influenced the number and success of Barn Swallow nests at nesting structures. 2019 Study Area We located study sites where an old structure with nesting Barn Swallows was removed or made unavail- able to the birds. This situation enabled us to simulate the circumstances under which many new nesting structures are being built in Ontario (i.e., mitiga- tion for loss of nesting habitat under the Endangered Species Act [MECP 2013]). We coordinated the con- struction of two new nesting structures at each site, during the fall or spring, prior to the breeding sea- son. The result was a paired design, with two new nesting structures on each study site, one treatment with conspecific cues and one control without con- specific cues, thus allowing us to assess the influence of the cues on Barn Swallow use of the new nesting structures. We flipped a coin to randomly select one of the two structures on each of the 10 study sites to have conspecific cues (i.e., decoys and a broad- cast box). We established 10 study sites in southern Ontario between Erin (43.766°N, 80.058°W) in the north and Port Rowan (42.626°N, 80.452°W) in the south. We opportunistically identified study sites through existing contacts and by directly contacting land- owners. Barn Swallow nesting habitat was lost at study sites prior to our study because structures were demolished or access to structures was blocked due to building renovation, nesting deterrents, or a need to keep doors closed (Table 1). The number of Barn Swallow nests in structures where habitat was lost varied, from one to ~50, across study sites (Table 1). We were unable to estimate the number of nests that were used in the year before habitat was lost, which would have provided better information about the number of nesting pairs compared to the number of nests. To the best of our knowledge, Barn Swallows CAMPOMIZZI ET AL.. ATTRACTING BARN SWALLOWS TO STRUCTURES 231 nested at >8 of the 10 study sites in the year prior to our experiment. We used the following criteria to guide where to place new nesting structures. We placed structures in non-forested areas with foraging habitat for Barn Swallows (1.e., grassland, cropland; Brown and Brown 1999). Additionally, we attempted to build new struc- tures <1 km from the previous nesting location to meet mitigation guidelines (MECP 2013) and so that Barn Swallows returning to the site could easily en- counter the new structures. We attempted to place the two new structures equal distances from the location of the demolished, renovated, or closed structure and about 400 m apart from each other to minimize the effects of the conspecific cues on the control structure (1.e., to ensure that broadcasted vocalizations were inaudible at control structures). Additionally, we at- tempted to place structures >100 m from forest edges to maximize availability of proximate foraging habi- tat. Because of constraints on study sites, we placed nesting structures 81-1220 m (mean = 427 m) from the location where Barn Swallows nested previously, 265-589 m (mean = 378 m) apart from each other, and 16—474 m (mean = 167 m) from the nearest forest edge based on land cover data from the Southern Ontario Land Resource Information System (MNRF 2000). Methods Structures We designed nesting structures using the best available information about what Barn Swallows would most likely use (Brown and Brown 1999), However, information about structures built for Barn Swallow nesting is limited and not in the literature. The best available information suggested building TABLE 1. Reason for habitat loss, number of Barn Swallow (Hirundo rustica erythrogaster) nests in lost habitat in year before monitoring (number of previously-active nests unknown), number of new nests in structures with and without pre- breeding conspecific cues built to replace lost habitat, and the year each study site was monitored. Site Old structure No. nests in new structure Year Habitat lost No. nests Conspecific cues No cues monitored AN Barn demolished ~12 2 2: 2014 CH Barn access denied Unknown* 0 0 2015 DA Barn access denied 1 to 27 0 0 2015 DR Barn access denied ~6 pairs 0 0 2015 GU Barn access denied 1§ 0 0 2015 LA Barn demolished 6 0 1 2014 LE Barn demolished 20 to 50 2 1 2015 RA Eaves access denied 1 2 0 2015 WA Barn access denied 4to5 0 1 2015 Wi 3 buildings demolished ~15 0 0 2015 *Landowner observed several nesting pairs previously using structure, but structure was inaccessible to confirm presence of nests. *Nests active in 2012. tNumber of nests unknown, landowner estimated six nesting pairs. SUnknown if nest active in 2014. 238 structures with similar characteristics to bridges and barns that are used for nesting, including rough ver- tical surfaces on which birds can build nests, shelter from wind and rain, visual barriers between nests, and a structure large enough to support several nest- ing pairs (MECP 2013; L. Sarris pers. comm. 13 February 2014; K.R. unpubl. data). We designed a wooden structure with a metal roof, 4.9 m long, 1.3 m wide at the nesting compartments, and 3.7 m tall at the peak of the roof (Figure 1). The structure included 16 nesting compartments, two rows of eight compart- ments along the 4.9-m length of the structure. In each row of eight compartments, we alternated available nest supports by providing a wooden nest cup (i.e., a wooden replica of a nest) in one compartment and bridging in the shape of the letter X, as found in some old barns, in the next compartment. Each compart- ment was bordered by 5 x 25 cm lumber along the center and along the outside of the structure and 5 x 15 cm lumber between compartments on the inside of the structure to provide a visual barrier between nests. Compartments had a flat ceiling above and no obstructions below. To provide shelter from weather, we added 40 cm of lumber along the outside of the structure below the nesting compartments. FicureE 1. One of the nesting structures built to test the impact of conspecific cues on prospecting and nesting by Barn Swallows (Hirundo rustica erythrogaster) in southern Ontario, Canada. Photo: A.J. Campomizzi. THE CANADIAN FIELD-NATURALIST Vol. 133 Each structure was equipped with nest cups, perches, and predator deterrents to encourage use by Barn Swallows and decrease risk of nest preda- tion. We placed nest cups on all structures because they are required for mitigation projects in Ontario (MECP 2013). Although the presence of old nests may increase the chance of nesting, Barn Swallow re- sponse to nest cups is unknown. We placed 16 nest cups on each structure, eight on the inside and eight on the outside of the structure. We placed nest cups far enough from ceilings (6.4 cm) and roofs (6.4 cm under roof peaks and 10.8 cm under sloped roofs) to allow the birds to build a mud rim along the top of the cup. The distance between the top of the nest cup and the ceiling or roof above was based on instructions provided by a nest cup supplier (American Artifacts, Taneytown, USA) and Barn Swallow nest placement (1.e., typically 2.5 to 6.0 cm from a ceiling; Brown and Brown 1999). We attached four perches to each structure; two on the inside and two on the outside. We included perches on the structures because there is evidence that adults lead juveniles from the nest to a perch, fledglings frequently perch after leaving the nest, and fledglings are initially fed by parents while perched (Brown and Brown 1999). To deter mam- malian nest predators from accessing and preying on nests, we covered each support post with galvanized sheet metal (Figure 1). Experiment one: pre-breeding prospecting We monitored structures at 10 study sites (two sites in 2014 and eight in 2015). We placed seven carved Barn Swallow decoys (Olde World Carvings, Spartanburg, South Carolinia, USA; Starr Decoys, Weybridge, Vermont, USA) at each treatment struc- ture on perches and nests to make it appear as though the structure was being used, but not fully occupied, by nesting Barn Swallows. We attached the broadcast box to a small shelf about 1.5 m from the ground on a post that supported each treatment structure. We largely followed Farrell and Campomizzi (2011) for the design of the broad- cast box, resulting in a plastic box containing a com- pact disc player, amplified speaker, battery, and timer that broadcasted Barn Swallow vocalizations period- ically throughout the day. We made a 30-min audio track of Barn Swallow songs, non-alarm calls, and periods of silence to simulate an active Barn Swallow nesting colony. To assemble the 30-min track, we ob- tained eight audio recordings made in Ontario and ad- jacent US states from the Cornell Lab of Ornithology (2014). To provide vocalizations throughout the day, we programmed the timer to turn the broadcast box on for 30 min at 0600, 0700, 0800, 1000, 1200, 1500, 1700, and 1900. We used the literature about Barn Swallow vocalizations to guide our selection of songs 2019 and calls to include, when to broadcast the vocaliza- tions during the day, and the number of vocalizations interspersed with silence (Samuel 1971; Brown 1985; Brown and Brown 1999). We installed conspecific cues for the return of Barn Swallows to the study area for breeding in the spring. We continued broadcasting vocalizations for the duration of the nesting season, until late August in 2014 and early September in 2015. Across 2014 and 2015, we monitored 10 treatment and 10 control structures for pre-breeding prospect- ing visits by Barn Swallows to assess if birds visited structures with conspecific cues more frequently than structures without cues and to record their behav- iour. We conducted prospecting surveys at a desig- nated survey location 50 m from each structure twice per week, between sunrise and sunset. Pre-breeding prospecting surveys occurred from 24 April to 14 June. This period corresponds to Barn Swallow ar- rival in the study area and the beginning of nesting. Seven to 21 days pass between pair formation and egg laying (Brown and Brown 1999) and the earliest egg date for Ontario is 10 May (Peck and James 1987). During each 10-min survey, we recorded each Barn Swallow detected within 50 horizontal m of the struc- ture. During each 2-min interval of the 10-min sur- vey, we recorded each individual detected, its distance from the structure, if it behaviourally interacted with the structure (perched on, flew under, or flew into or out of the structure). We also recorded if we detected an individual carrying nest material during the 10-min survey and if birds interacted with, perched next to, or attempted to copulate with decoys. We recorded a conservative estimate of the number of individuals to avoid counting one individual multiple times during a survey. We conducted surveys during weather condu- cive to Barn Swallow activity and detection (i.e., not during rain or strong wind). We noted if nest predators were on or attempting to get on the structure. After each survey, we walked to the structure to see if birds were inside and to monitor nests, as described below. Experiment two: post-breeding prospecting Following the pre-breeding prospecting experi- ment at each study site, we monitored 10 treatment and 10 control structures for post-breeding prospect- ing visits by Barn Swallows to assess if birds vis- ited structures with conspecific cues more frequently than structures without cues, and to record their be- haviour. Using the same broadcast schedule of Barn Swallow vocalizations and bird survey methods de- scribed above, we conducted post-breeding surveys from 20 July to 5 September, a range that includes when pairs not attempting second broods are finish- ing caring for dependent fledglings to when we no longer saw birds in breeding areas. The latest egg date for Ontario is 21 August (Peck and James 1987). CAMPOMIZZI ET AL.. ATTRACTING BARN SWALLOWS TO STRUCTURES 239 Experiment three: prospecting before, during, and after In 2016, we placed conspecific vocal cues at three structures that were randomly selected as treat- ments in 2015 but were not used for nesting by Barn Swallows in 2015. In 2016, each of the three study sites had a pair of structures, one with and one with- out conspecific vocal cues. We did not use decoys for experiment three. We changed the frequency and duration of vocal- izations played on each day at each treatment struc- ture compared to 2015 to enable assessment of Barn Swallow visits before, during, and after broadcasts of vocalizations. Vocalizations played for 20 min at the start of each hour between 0600 and 2100. Broadcasts began on 19 April and ceased on 6 June. In 2016, we surveyed the three nesting structures for pre-breeding prospecting visits by Barn Swallows. We designed surveys to document Barn Swallows searching for nest sites (particularly behavioural interactions with structures) and if conspecific cues influenced the frequency of detections. We visited treatment structures twice per week, once in the mor- ning and once in the afternoon or evening, for a one hour survey. We scheduled the majority of surveys during the morning and evening because, in 2014 and 2015, we observed more Barn Swallow activity dur- ing these times compared to other times. The survey hour consisted of 20 min before the broadcast, 20 min of broadcast, and 20 min after the broadcast. We ob- served treatment structures from a designated sur- vey location 50 m away, recording all individual Barn Swallows that came within 50 horizontal m. Survey periods were broken into 5-min intervals to record possible variation in bird abundance and behaviour throughout the survey. During each 5-min interval, we recorded detections of each individual. For each Barn Swallow detected, we recorded its horizontal distance from, and interactions with, the nesting structure. We conducted surveys during weather conducive to Barn Swallow activity and detection. After each survey, we approached the treatment structure to look for signs of nesting and check active nests. We did not conduct prospecting surveys of the control structures on the three study sites because we were testing Barn Swallow response to broad- casts at treatment structures only (prospecting sur- veys at control structures were conducted for the pre- breeding and post-breeding prospecting experiments, see above). We checked for nesting activity at con- trol structures after surveys were completed at treat- ment structures. Nest monitoring We monitored nests to assess differences in the number of nests and nest success of Barn Swallows 240 between structures with and without conspecific cues. We monitored 10 study sites across 2014 and 2015 following pre-breeding conspecific cues. Addi- tionally, we monitored nesting at eight study sites in 2016 following post-breeding conspecific cues ap- plied in 2015. Nest monitoring occurred from 12 May to 22 August. We followed nest monitoring protocols for Barn Swallows provided on Bird Studies Canada’s Project NestWatch website (http://www.birdscanada. org/volunteer/pnw/index.jsp?targetpg=barsmonitor), with minor modifications. We looked for evidence of nest building while conducting bird surveys early in the breeding season. At the five study sites without bird surveys in 2016, the frequency of nest monitor- ing visits varied based on whether there were active nests at the site. We checked active nests approxi- mately once per week until nesting activity ceased. Sites without active nests were checked periodically throughout the season. We began monitoring nests with a mirror to ob- serve nest contents on the visit after a nest appeared fully built, to minimize the chance of nest abandon- ment. We checked nest contents with a mirror once every five to seven days. During each nest check, we recorded the number of eggs, number and age of young, brood parasitism, adult activity, and condition of the nest. We did not check nest contents with a mir- ror if nestlings were >10 days old, to avoid potentially causing young to fledge prematurely; instead, we ob- served the nest from a distance with binoculars. We continued to check nesting structures for active nests throughout the breeding season. Analyses We did not use statistical analyses for nest data because sample size of nests was too small. Instead, we described nesting activity. For bird survey data, we first explored data through graphs and descrip- tive statistics. We excluded survey data collected while Barn Swallow nests were active at a structure to ensure that detections were of prospecting birds, not adults attending to nests. We used means and CI to assess the direction, magnitude, and precision of effects (Johnson 1999; Wasserstein and Lazar 2016), and interpreted their biological importance (Guthery et al. 2001; Nakagawa and Cuthill 2007; Nuzzo 2014). We calculated means and CI for the difference in Barn Swallow detections and interactions with struc- tures from spatially and temporally paired surveys described below. We conducted analyses using R (version 3.4.1, R Core Team 2017). Experiment one: pre-breeding prospecting. We separately compared the abundance of Barn Swallows detected and interacting with structures during pre- breeding surveys. We compared the difference in abundance of Barn Swallows detected between (1) THE CANADIAN FIELD-NATURALIST Vol. 133 broadcast and non-broadcast times at treatments, (2) broadcast and non-broadcast times at controls, and (3) all surveys at treatments compared to controls. We made the same three comparisons in the differ- ence in the abundance of Barn Swallows interacting with structures. For comparisons during broadcast and non-broadcast times at treatments and controls, we paired surveys conducted during the same week for each structure. For example, to calculate the dif- ference in abundance between broadcast and non- broadcast times at each treatment structure for each week of surveys, we subtracted the number of Barn Swallows detected during the non-broadcast time from the number of individuals detected while con- specific vocalizations were broadcasted. For com- parisons between treatments and controls, we paired surveys conducted on the same day for each pair of treatment and control structures at each study site. These analyses resulted in three estimates of the dif- ference in abundance of Barn Swallows detected at structures (Figure 2a) and another three estimates of the difference in abundance of individuals inter- acting with structures (Figure 2b) during pre-breed- ing surveys. Estimated means greater than zero indi- cate more Barn Swallows detected or interacting with structures during broadcast compared to non-broad- cast or treatment compared to control. Experiment two: post-breeding prospecting: For post-breeding surveys, we made the same compari- sons as pre-breeding surveys with one exception. We used surveys at treatments during broadcast and con- trols during non-broadcast only because mean abun- dance at treatments was more than twice as high during broadcast compared to non-broadcast times, suggesting a potential numerical response by the birds. These an- alyses resulted in an additional three estimates of the difference in abundance of Barn Swallows detected at structures (Figure 2a) and three estimates of the dif- ference in abundance of individuals interacting with structures (Figure 2b) during post-breeding surveys. Experiment three: prospecting before, during, and after. We compared the difference in abundance of Barn Swallows detected at treatment structures dur- ing broadcast versus before broadcast and after broadcast versus before. We paired data for surveys conducted on the same day for each structure. We were unable to make comparisons of the abundance of Barn Swallows interacting with structures because we did not detect interactions during these surveys. These analyses resulted in two estimates of the differ- ence in abundance of Barn Swallows detected at struc- tures (Figure 3). Estimated means greater than zero indicate more Barn Swallows detected during broad- cast compared to before broadcast or after broadcast compared to before. 2019 +8) 2.0 1.5 1.0 0.5 0.0 Difference in Barn Swallow abundance -1.0 Tr Con Tr - Con io a Pre-breeding mean e Post-breeding mean — 95% Cl Difference in Barn Swallow interactions Tr Con Tr - Con Comparison FIGURE 2. Mean and 95% CI of the difference in abun- dance of Barn Swallows (Hirundo rustica erythrogaster) a. detected and b. interacting with nesting structures (i.e., perched on, flew under, or flew into or out of the structure) with and without conspecific cues in southern Ontario, Canada in 2014 and 2015. Comparisons during pre-breed- ing are for structures treated with conspecific cues dur- ing broadcast minus non-broadcast surveys (Tr), control structures during broadcast minus non-broadcast surveys (Con), and all treatment minus control surveys (Tr - Con). Comparisons during post-breeding are the same for Tr and Con; the third comparison is of treatment during broad- cast minus control during non-broadcast surveys only (Tr - Con). Results Across all surveys in 2014 and 2015, we de- tected Barn Swallows on 33% (n = 263) of surveys at structures with conspecific cues and 38% (n = 263) of surveys at structures without cues. Across CAMPOMIZZI ET AL.. ATTRACTING BARN SWALLOWS TO STRUCTURES 241 & Mean — 95% Cl Data Difference in Barn Swallow abundance Broadcast - before After - before Comparison FiGurE 3. Mean, 95% CI, and observed data of the differ- ence in abundance of Barn Swallows (Hirundo rustica ery- throgaster) detected on one hour surveys conducted for 20 min before, during, and after conspecific vocalizations at three nesting structures in southern Ontario, Canada in 2016. surveys with Barn Swallow detections, we detected 279 Barn Swallows on 88 surveys at structures with conspecific cues and 299 Barn Swallows on 99 sur- veys at structures without cues. These results include data from surveys of structures with active nests. As noted above, we reduced the dataset for the compari- sons below. Experiment one: pre-breeding prospecting During pre-breeding, the mean difference in Barn Swallow abundance during broadcast compared to non-broadcast times at treatments was 0.04 (n = 46) and not substantially different from zero (Figure 2a). Similarly, the mean difference in Barn Swallow abun- dance during broadcast compared to non-broadcast times at controls (—0.26, n = 50) and at treatments compared to controls (—0.21, 7 = 98) was not substan- tially different from zero (Figure 2a). Experiment two: post-breeding prospecting During post-breeding, all three mean differences in Barn Swallow abundance were larger than during pre-breeding and greater than zero. The mean differ- ence in Barn Swallow abundance during broadcast compared to non-broadcast times was 0.78 (n = 42) at treatments, 0.27 (n = 48) at controls, and 0.69 (n = 52) at treatments compared to controls. These differ- ences suggest an effect of ~0.5 individuals per sur- vey, but 95% CI included zero, although marginally for treatments compared to controls (lower 95% CI: —0.06; Figure 2a). 242 The largest differences in abundance of Barn Swallows interacting with structures was during post- breeding. The mean difference in Barn Swallows in- teracting with structures during broadcast compared to non-broadcast was 0.28 (n = 42) at treatments and 0.21 (n = 52) for treatments compared to controls (Figure 2b). Both CI marginally included zero (lower 95% CI: —0.09 for treatments and —0.01 for treat- ments compared to controls), indicating some lack of confidence in an effect of ~0.2 individuals per survey interacting with structures (Figure 2b). Experiment three: prospecting before, during, and after We detected 40 individual Barn Swallows on prospecting surveys in 2016 and 45% (n = 42) of sur- veys for this experiment had Barn Swallow detec- tions. We detected 12, 20, and 26 individuals before, during, and after conspecific broadcast, respectively. Mean difference in abundance of Barn Swallows de- tected at treatment structures was higher both during broadcast compared to before broadcast (0.19, n = 42) and after broadcast compared to before (0.33, n = 42; Figure 3). The CI for after broadcast compared to be- fore was greater than zero (95% CI: 0.01, 0.65; Figure 3). We did not observe Barn Swallows behaviourally interacting with structures (i.e., perching on or flying inside of a structure) in 2016. Nesting Experiment one: pre-breeding prospecting: Across 2014 and 2015, there were six nests on three struc- tures with conspecific cues and five nests on four structures without cues (Table 1). All nests observed with eggs eventually fledged young. Two additional nests were built on structures without cues; however, we never observed eggs in these nests. All three nest- ing pairs at structures with conspecific cues appeared to fledge a second clutch, compared to one of four pairs nesting at structures without cues. The earliest nest initiation date (i.e., first egg date) was 20 May at structures with conspecific cues and 21 May at struc- tures without cues. All nests were built in wooden nest cups in the interior of the structures. For all 11 nests monitored following pre-breeding cues, Barn Swallows added a mud rim to the top of the wooden nest cup, mak- ing the top of the nest look similar to a typical Barn Swallow nest. Experiment 2: post-breeding prospecting: In 2016, four nests were built in two of eight structures treated with post-breeding conspecific cues in 2015, com- pared to four nests in three of eight structures with- out post-breeding cues in 2015. All eight nests were in nest cups in the interior of the structure and fledged young. One additional nest was initiated on the ex- THE CANADIAN FIELD-NATURALIST Vol. 133 terior of a structure but was abandoned after some addition of mud to a nest cup; we did not observe eggs in this nest. Two nests from 2015 were reused in 2016. Three of the eight nests appeared to be second clutches. Experiment 3: prospecting before, during, and after: Barn Swallows did not nest at the three sites used for the experiment comparing prospecting be- fore, during, and after broadcasts of vocalizations in 2016. Discussion To our knowledge, this is the first evaluation of Barn Swallow use of new nesting structures specif- ically built for the species. Our study provides some evidence to link prospecting behaviour by Barn Swallows at new nesting structures to broadcasts of conspecific vocalizations. During pre-breeding, Barn Swallow abundance was higher immediately after conspecific broadcasts compared to before broad- casts. However, we did not find substantial differ- ences in Barn Swallow abundance and interactions with new nesting structures for other comparisons involving conspecific cues. Most importantly, Barn Swallows did not nest more frequently on structures treated with pre- or post-breeding conspecific cues; they nested on and fledged young from structures with and without cues, albeit in low numbers. Our results provide some evidence that Barn Swallows can be attracted to at least investigate new nesting structures by introducing conspecific cues. On several occasions, we observed Barn Swallows approaching nesting structures immediately after the broadcast started, anecdotally suggesting a response to the start of the vocalizations. On a few occasions, we observed Barn Swallows interacting with decoys by perching on, singing at, and attempting to copu- late with decoys. Previous research in the literature has not documented Barn Swallow prospecting be- haviour in response to simulated conspecific cues at nesting sites. A study on Chimney Swift (Chaetura pelagica, another aerial insectivore species) showed that introducing conspecific vocalizations and de- coys increased the length of time that the birds spent near new nesting towers (Finity and Nocera 2012). Additionally, conspecific cues introduced during post-breeding have been shown to influence habi- tat selection in the next breeding season for migra- tory songbirds (Nocera et al. 2006, Betts et al. 2008). In our study, however, Barn Swallows did not build more nests in 2016 at structures treated with con- specific cues during post-breeding in 2015, compared to structures without cues. Future research to explore the impacts of broadcasted conspecific vocalizations on prospecting behaviour and nesting may help in- 2019 form future efforts to create nesting habitat for vari- ous species. Some of the structures included in this project provided nesting habitat for Barn Swallows and all nests observed with eggs fledged young. We did not observe a difference in the number of nests built on structures with and without conspecific cues that were provided during pre-breeding or post-breeding. Although nesting structures provided opportunities for birds to nest on X-shaped bridging in addition to nest cups, all nests were built in nest cups. Nest cups may be an important feature of new nesting struc- tures because they provide a nesting substrate and essentially a partially-built nest. The nest cups may attract Barn Swallows to new structures if they func- tion similarly to old nests (sensu Safran 2004, 2007; Ringhofer and Hasegawa 2014) and enable birds to begin nesting earlier in the season because the birds do not need to build an entire nest. Re-using old nests can increase reproductive success (Safran 2007; but see Barclay 1988). Therefore, nest cups could be im- portant for conservation because they may enable Barn Swallow pairs to raise a second brood, thus increasing fecundity. Combining conspecific cues and the presence of old nests (perhaps by providing wooden nest cups) may maximize the number of Barn Swallows that prospect at new nesting structures but may not lead to more nests at new structures. It is possible that the structures with and without cues were not far enough apart to completely separ- ate the effect of the conspecific cues. Although the distance Barn Swallows travel to prospect for nest sites is unknown, adults will forage up to 500 m from nesting colonies (Moller 1987), suggesting in- dividuals encountering one structure on a study site could encounter the other structure. Future research to assess if conspecific cues at one structure can af- fect prospecting at multiple structures, or if prospect- ing is greater at structures with conspecific cues com- pared to structures without cues (at greater distances than we tested), may be helpful for understanding nest site selection and guiding conservation efforts. Additionally, most of our study sites had few nests in the nesting habitat that was lost. With few Barn Swallows returning to nest at these sites, there may have been few Barn Swallows within hearing dis- tance of the vocalizations. The number of philopatric Barn Swallows may impact the magnitude of the ef- fect of conspecific cues on prospecting birds. We are uncertain how many nesting pairs could nest simultaneously on the structures used for our experiment. However, building a few of these new structures is unlikely to replace the lost nesting habitat provided by bridges or old barns with large nesting colonies (e.g., 50 breeding pairs). Building CAMPOMIZZI ET AL.. ATTRACTING BARN SWALLOWS TO STRUCTURES 243 one nesting structure cost ~$2500 to $3500 (CAD). Regulators and land managers should consider if this expense is worth the benefit or if funds could be used in more effective ways to support Barn Swallow nest- ing habitat. A potential alternative is to provide in- centives for landowners to repair and maintain aging barns that can provide nesting habitat for larger col- onies of Barn Swallows and for more years than new structures (Heagy ef al. 2014). It may also be benefi- cial for future research to investigate the relationship between colony size and characteristics of nesting structures and the surrounding landscape. Building new nesting structures may be an option for creat- ing new Barn Swallow nesting habitat in locations with appropriate foraging habitat (i.e., grassland, cropland; Brown and Brown 1999), where no struc- ture currently exists and there is an interest in con- tributing to Barn Swallow conservation. When struc- tures are built for Barn Swallow nesting habitat, we recommend including wooden nest cups in the in- terior of the structure, which was the location of all nests in our experiment. However, more research is needed to assess if loss of nesting habitat is limiting the Barn Swallow population to determine if creating or maintaining nesting habitat is likely to have a posi- tive impact on the population or if resources should be directed to addressing other threats to the species. Our results confirm that new structures can pro- vide nesting habitat for Barn Swallows but provid- ing conspecific cues may not enhance this conserva- tion strategy. The paucity of nests built on structures raises questions about the efficacy and efficiency of building new nesting structures to mitigate the loss of nesting habitat. Author Contributions Conceptualization: Z.M.L., A.J.C., and K.R.; Data Curation: A.J.C. and Z.M.L.; Formal Analysis: A.J.C.; Funding Acquisition: K.R., Z.M.L., and A.J.C.; Investigation: Z.M.L., A.J.C., and K.R.; Methodology: A.J.C., Z.M.L., and K.R.; Project Administration: Z.M.L., K.R., and A.J.C.; Resources: Z.M.L. and K.R.; Supervision: Z.M.L. and K.R.; Validation: A.J.C. and Z.M.L.; Visualization: A.J.C. and Z.M.L.; Writing — Original Draft: A.J.C. and Z.M.L.; Writing — Review & Editing: A.J.C., Z.M.L., and K.R. Acknowledgements Funding for this project was provided by the Government of Ontario, TD Friends of the Environ- ment Foundation, The City of Waterloo, Colleges and Institutes Canada Clean Tech Internship pro- gram, Echo Foundation, the Government of Canada (Canada Summer Jobs Program), Ontario Soil and Crop Improvement Association through the Species 244 at Risk Farm Incentive Program (SARFIP), and indi- vidual donors. We are grateful for assistance with field monitoring from Betty Chanyi, Timothy Fernandes, Rebecca Howe, Jaelyn Kloepfer, Katherine Robbins, Gail Tako, Bob Wood, Graham Wood, Karen Wood, and Carolyn Zanchetta. Thanks to Becky Stewart, Bird Studies Canada, for the opportunity to undertake this project. We also thank the Nature Conservancy of Canada, rare Charitable Research Reserve, The Arboretum at the University of Guelph, The City of Waterloo, and several private landowners for allowing construction and monitoring of nesting structures on their properties. Richard Van Vleck (American Artifacts) and Larry Sarris (Ontario Ministry of Transportation) provided substantial helpful informa- tion about structures for Barn Swallows. Thanks to Jeff Sauder (for assistance designing structures and leading construction) and the other builders. Mike Cadman, Tara Imlay, and two anonymous review- ers provided helpful comments on previous versions of this manuscript. The views expressed herein are those of the authors, not funders or other entities. Literature Cited Ahlering, M.A., and J. Faaborg. 2006. Avian habi- tat management meets conspecific attraction: if you build it, will they come? Auk 123: 301-312. https://doi. org/10.1642/0004-8038(2006)123[301:ahmmca]2.0.co;2 Barclay, R.M.R. 1988. Variation in the costs, benefits, and frequency of nest reuse by Barn Swallows (Hirundo rustica). Auk 105: 53—60. https://doi.org/10.1093/auk/ 105.1.53 Betts, M.G., A.S. Hadley, N. Rodenhouse, and J.J. Nocera. 2008. Social information trumps vegetation structure in breeding-site selection by a migrant song- bird. Proceedings of the Royal Society of London B: Biological Sciences 275: 2257-2263. https://doi.org/10. 1098/rspb.2008.0217 Bird Life International. 2016. Hirundo rustica. The IUCN Red List of Threatened Species 2016: e.T22712252A8 7461332. http://doi.org/10.2305/iucn.uk.2016-3.rlts.t227 12252a87461332.en Brown, C.R. 1985. Vocalizations of Barn and Cliff Swal- lows. Southwestern Naturalist 30: 325-333. https://do1. org/10.2307/3671263 Brown, C.R., and M.B. Brown. 1999. Barn Swallow (Hi- rundo rustica), The Birds of North America Online. Edited by A. Poole. Cornell Lab of Ornithology, Ithaca, New York, USA. Accessed 1 February 2014. https:// birdsna.org/Species-Account/bna/species/barswa/ introduction. Cornell Lab of Ornithology. 2014. Macaulay Library. Accessed 14 April 2014. https:/(www.macaulaylibrary. org/. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2011. COSEWIC assessment and status report on the Barn Swallow Hirundo rustica in Canada. COSEWIC, Ottawa, Ontario, Canada. Danchin, E., L.A. Giraldeau, T.J. Valone, and R.H. Wag- THE CANADIAN FIELD-NATURALIST Vol. 133 ner. 2004. Public information: from nosy neighbors to cultural evolution. Science 305: 487-491. https://doi. org/10.1126/science.1098254 Danchin, E., R.H. Wagner, J.K. Parrish, and L. Edelstein-Keshet. 2000. Benefits of membership. Sci- ence 287: 804—807. https://doi.org/10.1126/science.287. 5454.803e Doligez, B., T. Part, and E. Danchin. 2004. Prospecting in the collared flycatcher: gathering public information for future breeding habitat selection? Animal Behaviour 67: 457-466. https://do1.org/10.1016/j.anbehav.2003.03.010 Farrell, S.L., and A.J. Campomizzi. 2011. A component system for broadcasting sound for research and man- agement. Journal of Wildlife Management 75: 463—466. https://doi.org/10.1002/jwmg.45 Farrell, S.L., M.L. Morrison, A.J. Campomizzi, and R.N. Wilkins. 2012. Conspecific cues and breeding habitat selection in an endangered woodland warbler. Journal of Animal Ecology 81: 1056-1064. https://doi. org/10.1111/j.1365-2656.2012.01995.x Finity, L., and J.J. Nocera. 2012. Vocal and visual con- specific cues influence the behavior of Chimney Swifts at provisioned habitat. Condor 114: 323-328. https://doi. org/10.1525/cond.2012.110145 Government of Canada. 2017. Species profile: Barn Swal- low. Accessed 20 March 2019. https://wildlife- species. canada.ca/species-risk-registry/species/species Details_e.cfm?sid=1147. Guthery, F.S., J.J. Lusk, and M.J. Peterson. 2001. The fall of the null hypothesis: liabilities and opportunities. Journal of Wildlife Management 65: 379-384. https:// doi.org/10.2307/3803089 Hahn, B.A., and E.D. Silverman. 2006. Social cues fa- cilitate habitat selection: American redstarts establish breeding territories in response to song. Biology Letters 2: 337-340. https://doi.org/10.1098/rsbl.2006.0472 Heagy, A., D. Badzinski, D. Bradley, M. Falconer, J. McCracken, R.A. Reid, and K. Richardson. 2014. Recovery Strategy for the Barn Swallow (Hirundo rustica) in Ontario. Ontario Recovery Strategy Se- ries. Accessed 20 March 2019. http://files.ontario.ca/ environment-and-energy/species-at-risk/mnr_sar_rs_ brn_swl_en.pdf. Johnson, D.H. 1999. The insignificance of statistical sig- nificance testing. Journal of Wildlife Management 63: 763-772. https://doi.org/10.2307/3802789 Kostka, K. 2000. Social attraction: a new technique for establishing a Purple Martin colony site. Purple Martin Conservation Association. Accessed 1 December 2015. http://www. purple-martin.org/PMArticles/Social AttractionPMPA htm. Lepage, D. 2007. Barn Swallow. Pages 398-399 in Atlas of the breeding birds of Ontario, 2001-2005. Edited by M.D. Cadman, D.A. Sutherland, G.G. Beck, D. Lepage, and A.R. Couturier. Bird Studies Canada, Environment Canada, Ontario Field Ornithologists, Ontario Ministry of Natural Resources, Ontario Nature, Toronto, Ontario, Canada. 2019 MECP (Ministry of Environment, Conservation and Parks. 2012. Species at risk in Ontario. Accessed 21 March 2019. https://(www.ontario.ca/page/species-risk- ontario. MECP (Ministry of Environment, Conservation and Parks). 2013. Alter a structure (habitat for Barn Swal- low). Accessed 21 March 2019. https://www.ontario.ca/ page/alter-structure-habitat-barn-swallow. MNRF (Ministry of Natural Resources and Forestry). 2000. Southern Ontario Land Resource Information System (SOLRIS). Accessed 21 March 2019. https:// data.ontario.ca/dataset/southern-ontario-land-resource- information-system-solris-20. Moller, A.P. 1987. Advantages and disadvantages of co- loniality in the swallow, Hirundo rustica. Animal Be- haviour 35: 819-832. https://doi.org/10.1016/S0003- 3472(87)80118-5 Nakagawa, S., and I.C. Cuthill. 2007. Effect size, confi- dence interval and statistical significance: a practical guide for biologists. Biological Reviews 82: 591-605. https://doi.org/10.1111/j.1469-185X.2007.00027.x Nebel, S., A. Mills, J.D. McCracken, and P.D. Taylor. 2010. Declines of aerial insectivores in North America follow a geographic gradient. Avian Conservation and Ecology 5(2): 1. http://doi.org/10.5751/ACE-00391-050201 Nocera, J.J., G.J. Forbes, and L.A. Giraldeau. 2006. Inadvertent social information in breeding site selec- tion of natal dispersing birds. Proceedings of the Royal Society of London B: Biological Sciences 273: 349-355. https://doi.org/10.1098/rspb.2005.3318 Nuzzo, R. 2014. Statistical errors: P values, the “gold stan- dard” of statistical validity, are not as reliable as many scientists assume. Nature 506: 150-152. https://doi.org/ 10.1038/506150a Parrish, J.K., and L. Edelstein-Keshet. 1999. Complex- ity, pattern, and evolutionary trade-offs in animal ag- gregation. Science 284: 99-101. https://doi.org/10.1126/ science.284.5411.99 Peck, G.K., and R.D. James. 1987. Breeding Birds of On- tario: Nidiology and Distribution Volume 2: Passerines. The Royal Ontario Museum, Toronto, Ontario, Canada. https://doi.org/10.5962/bh1 title.60694 R Core Team. 2017. R: a language and environment for sta- tistical computing. R Foundation for Statistical Com- puting, Vienna, Austria. Reed, J.M., T. Boulinier, E. Danchin, and L.W. Oring. 1999. Informed dispersal: prospecting by birds for breed- ing sites. Current Ornithology 15: 189-259. https://doi.org/ 10.1007/978-1-4757-4901-4_5 CAMPOMIZZI ET AL.. ATTRACTING BARN SWALLOWS TO STRUCTURES 245 Ringhofer, M., and T. Hasegawa. 2014. Social cues are preferred over resource cues for breeding-site selection in Barn Swallows. Journal of Ornithology 155: 531-538. https://doi.org/10.1007/s10336-013-1035-3 Safran, R.J. 2004. Adaptive site selection rules and vari- ation in group size of barn swallows: individual deci- sions predict population patterns. American Naturalist 164: 121-131. https://doi.org/10.1086/422198 Safran, R.J. 2007. Settlement patterns of female barn swal- lows Hirundo rustica across different group sizes: ac- cess to colorful males or favored nests? Behavioral Ecology and Sociobiology 61: 1359-1368. https://doi.org/ 10.1007/s00265-007-0366-6 Samuel, D.E. 1971. Vocal repertoires of sympatric Barn and Cliff Swallows. Auk 88: 839-855. https://doi.org/10. 2307/4083842 Sauer, J.R., W.A. Link, J.E. Fallon, K.L. Pardieck, and D.J. Ziolkowski, Jr. 2013. The North American Breeding Bird Survey 1966-2011: summary analysis and species accounts. North American Fauna 79: 1-32. https:// doi.org/10.3996/nafa.79.0001 Sauer, J.R., K.L. Pardieck, D.J. Ziolkowski, Jr., A.C. Smith, M.R. Hudson, V. Rodriguez, H. Berlanga, D.K. Niven, and W.A. Link. 2017. The first 50 years of the North American Breeding Bird Survey. Condor 119: 576-593. https://doi.org/10.1650/condor-17-83.1 Schlaepfer, M.A., M.C. Runge, and P.W. Sherman. 2002. Ecological and evolutionary traps. Trends in Ecology & Evolution 17: 474-480. https://doi.org/10.1016/S0169- 5347(02)02580-6 Shields, W.M., and J.R. Crook. 1987. Barn Swallow col- oniality: a net cost for group breeding in the Adiron- dacks? Ecology 68: 1373-1386. https://doi.org/10.2307/ 1939221 Smith, A.C., M.R. Hudson, C.M. Downes, and C.M. Francis. 2015. Change points in the population trends of aerial-insectivorous birds in North America: synchron- ized in time across species and regions. PLoS ONE 10: e0130768. https://doi.org/10.1371/journal.pone.0130768 Ward, M.P. 2005. Habitat selection by dispersing yellow- headed blackbirds: evidence of prospecting and the use of public information. Oecologia 145: 650—657. https:// doi.org/10.1007/s00442-005-0179-0 Wasserstein, R.L., and N.A. Lazar. 2016. The ASA’s statement on p-values: context, process, and purpose. American Statistician 70: 129-133. https://do1.org/10.1 080/00031305.2016.1154108 Received 27 March 2019 Accepted 18 December 2019 The Canadian Field-Naturalist Seasonal movements of White-tailed Deer (Odocoileus virginianus) in the Rocky Mountains of British Columbia TREVOR A. KINLEY Parks Canada Agency, P.O. Box 220, Radium Hot Springs, British Columbia VOA 1MO Canada; email: trevor.kinley@ canada.ca Kinley, T.A. 2019. Seasonal movements of White-tailed Deer (Odocoileus virginianus) in the Rocky Mountains of British Columbia. Canadian Field-Naturalist 133(3): 246-252. https://doi.org/10.22621/cfn.v13313.2201 Abstract Nineteen adult female White-tailed Deer (Odocoileus virginianus), fitted with very high frequency or global positioning system collars in the Rocky Mountains of southeast British Columbia, exhibited straight-line seasonal movements ranging from <4 km to 109 km. Movement was almost entirely along the floor of both low- and high-elevation valleys, although there was some use of mid-elevation mountainsides during early winter. Spatial locations of deer spanned 891—2234 m above sea level. Seasonal movements of these deer from a single winter range extended to two provinces, three national parks, one provincial park, non-park provincial Crown land, and private land. Deer populations with similar movement patterns may be most effectively managed by considering their extensive movements and coordinating approaches across jurisdictions. Key words: Kootenay National Park; Odocoileus virginianus, seasonal movement; summer range; White-tailed Deer; winter range Introduction White-tailed Deer (Odocoileus virginianus) in- habit a range of ecosystems across North and South America. Some individuals and populations exhibit migration (round-trip movements between distinct seasonal ranges, sensu Berger 2004) of tens of kilo- metres, with greater movements typical in north- ern or mountainous locations (Baumeister 1992; Demarais et al. 2000; Robinson et al. 2002; Nelson et al. 2004; Brinkman et al. 2005; Stewart et al. 2011). In some study areas, individuals may be sedentary, facultative migrators, or obligate migrators (Sabine et al. 2002; Brinkman et al. 2005; Fieberg et al. 2008; Grovenburg ef a/. 2011). Deer on low-quality win- ter range may be more likely to migrate as a result of density-dependent competition (Henderson ef al. 2018). During spring and summer, an advancing line of greening vegetation offers ungulates in the Rocky Mountains the opportunity to follow high-quality habitat up slope (Merkle et al. 2016; Middleton et al. 2018). Based on long-term roadside surveys and inci- dental observations, White-tailed Deer in Kootenay National Park (K NP), British Columbia (BC), Canada, are common from spring through fall, but absent or nearly so during winter (S. Wrazej unpubl. data). Considering those observations in the context of the strong elevation gradient in the area, seasonal eleva- tion differences reported for nearby deer populations (Robinson et al. 2002; Hoekman et al. 2006), and an expectation of generally low-quality winter ranges for deer in snowy, mountainous areas in the northern part of their range, I speculated that deer summering in the park overwintered at lower elevations south of the park. I collared adult female White-tailed Deer south of the park and within the park, and investi- gated their movement patterns. Deer were monitored for variable periods and could not all be confirmed to make return movements (migrations); thus, I use the more general term “seasonal movement”. I report on seasonal movements and elevation-use patterns of these collared deer, including in relation to jurisdic- tional boundaries potentially affecting management regimes. Study Area The Beaverfoot and upper Kootenay Rivers are part of the Columbia River watershed of southeast BC (Figure 1). Their headwaters rise in the same valley in the Rocky Mountains. From there, the Beaverfoot River flows generally north by northwest into Yoho National Park where it joins the Kicking Horse River, which eventually exits the Rocky Mountains and flows into the Columbia River in the Rocky Mountain 246 ©Her Majesty the Queen in Right of Canada, as represented by the Minister of Environment and Climate Change Canada and Responsible for Parks Canada, 2020. 2019 KINLEY: SEASONAL MOVEMENTS OF WHITE-TAILED DEER 247 Sake Louise : — ed British Columbia system (black dots) or very high frequency (white dots) collars in the upper Kootenay River valley of British Columbia, 2011-2016. Winter range is oval in lower right. 248 Trench (hereafter “Trench’”). The Kootenay River flows south by southeast, passing through and beyond KNP before exiting the Rockies into the Trench. The elevation of the valley bottom is ~1250 m at the head- waters and ~1050 m at the downstream ends of the parks. Mountains adjacent to the rivers reach a max- imum elevation of 2400-3000 m. Leading tree species vary with elevation and lo- cation, but along the valley bottoms are primarily Lodgepole Pine (Pinus contorta Douglas ex Loudon), Engelmann Spruce (Picea engelmannii Engelmann), Trembling Aspen (Populus tremuloides Michaux), Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco), Western Larch (Larix occidentalis Nuttall), and, locally, Western Red Cedar (Thuja plicata Donn ex D. Don). At higher elevations, leading tree spe- cies are Engelmann Spruce and Subalpine Fir (Abies lasiocarpa (Hooker) Nuttall). Vegetation is domin- ated by mixed-age, mixed-species stands of those conifers interspersed with burns, wetlands, cutblocks from past logging (outside of parks), and non-forested areas on the highest peaks. This variety includes the range of grass, shrub, and open forest cover types normally selected by deer nearby and also the greater canopy cover selected under deep snow conditions (Hoekman et al. 2006). There is no agriculture within the study area. Methods Deer were captured in Clover traps (VerCauteren et al. 1999) baited with either hay and liquid and dried commercial deer attractants or hay, salt, apple, and dried molasses. One deer was immobilized by free- range darting (Dan-Inject APS, Berkop, Denmark) using a medetomidine—ketamine mixture (Caulkett et al. 2000). Deer captures were undertaken primarily in February 2014 on a winter range 15—20 km south of the southern boundary of KNP (Figure 1). This was at the confluence of the Kootenay and Palliser Rivers, at an elevation of 950-1100 m. Capture also occurred during November 2011, April 2012, and November— December 2015 within or beside anthropogenic forest openings in KNP, at ~1160 m elevation. Females <11 months old and all males were re- leased. All other females were fitted with collars, either global positioning system (GPS; G2110D, Advanced Telemetry Systems, Isanti, Minnesota, USA) or very high frequency (VHF; LMRT-2, Lotek Wireless, Newmarket, Ontario, Canada). One female originally fitted with a VHF collar was later recap- tured and fitted with a GPS collar. GPS collars at- tempted fixes hourly, were programmed to detach about 10 months after collaring, and were down- loaded on retrieval. VHF-collared deer were re- located on an approximately two-week schedule THE CANADIAN FIELD-NATURALIST Vol. 133 through ground monitoring. Aerial monitoring was undertaken twice in late winter of the first year, when only one deer was collared and snow depth prevented ground access, and once for all deer in another year as they left the winter range when not all deer could be located from the ground. Maximum straight-line movements were deter- mined for each deer monitored from at least January or February to July or August or the reverse for one or more years, 1.e., for those with potential to dem- onstrate seasonal movements. Universal Transverse Mercator coordinates were used to calculate Euclid- ean distances between the most distant points. Elevations reported here are only as recorded on- board GPS collars because, in the mountainous study area, relatively small horizontal errors would trans- late into considerable elevation errors if extracted via a geographic information system, especially for VHF collar data. All maximum and minimum ele- vation records were confirmed to be within clusters of sequential locations and, hence, unlikely to reflect significant GPS error. To represent movement vec- tors at an appropriate scale, sequential records of deer locations were manually approximated graphically. Where overlapping individuals were not distinguish- able, these representations were further linearized for visual clarity. Results Time from date of collaring to death, collar drop, or cessation of monitoring ranged from 299 to 1417 days for the 10 VHF-collared deer (x = 667, SD 295) and from 166 to 320 days for the nine GPS-collared deer (x = 286, SD 56) for which seasonal movements were calculated. Maximum straight-line movements (Figure 2) ranged from 6.1 to 82.4 km for VHF- collared deer (x = 33.0 km, SD 27.3) and from 3.4 to 109.2 km for deer with GPS collars (x = 48.7 km, SD 40.0). Among GPS-collared deer, maximum eleva- tions ranged from 1199 m to 2234 m and minimum elevations from 891 m to 997 m (pooled sample: 10% < 1010 m, 10% > 1382 m). Variation in elevation use was evident during summer, with two deer occurring at maximum elevations (>1900 m) at a time when all others were below 1500 m, and during early winter when two deer used elevations above 1600 m while others were below 1300 m (Figure 3). Seventeen deer were not recorded outside the con- tiguous Kootenay—Beaverfoot—K icking Horse valley (Figures 1 and 2), but two moved into a major tribu- tary valley or crossed the Continental Divide into Alberta. All 19 occurred for at least part of the year on provincial Crown land in BC, of which at least nine also made use of Kootenay, Yoho, or Banff na- tional parks, one of Spray Valley Provincial Park, 2019 KINLEY: SEASONAL MOVEMENTS OF WHITE-TAILED DEER 249 ~ = < ; 4 J os , “e - Sake Louise i Ti Figure 2. Maximum extent of movements of 19 female White-tailed Deer (Odocoileus virginianus) fitted with global positioning system (black lines) or very high frequency (white lines) collars in the upper Kootenay River valley of British Columbia, 2011-2016. Movements are presented as linear vectors for visual clarity. Maximum elevation (m) THE CANADIAN FIELD-NATURALIST Vol. 133 Month FiGurE 3. Maximum elevation per month of nine White-tailed Deer (Odocoileus virginianus) captured in the upper Kootenay River valley of British Columbia and fitted with global positioning system collars, 2012-2016. Alberta, and five of private land in BC. Of the nine using one or more national parks for part of the year, at least eight were on Crown or private land in BC during part or all of the current regular, youth, or bow-only “antlerless” hunting seasons from October through December (MFLNRORD 2018). One col- lared deer summered south of the winter range; all others were generally north. Discussion Given the lower frequency of monitoring of VHF collars and potential effects of limited access on manual monitoring, data from VHF collars likely underrepresent deer movements and use of high elevations relative to GPS collars. The apparently shorter maximum movements of VHF-collared deer despite longer duration of monitoring may reflect that. However, for both collar types, a wide range and broadly similar distribution of maximum movements was recorded. Even with the possibility that some deer movements reported in other studies may have represented dispersals, mean and maximum seasonal movements reported here are greater than values re- ported from nearby studies in the mountains of BC and northwest United States (Morgan 1993; Secord 1994; Robinson et al. 2002; Hoekman et al. 2006) and in 10 earlier studies from the same region summar- ized by Baumeister (1992: 56) and similar to those observed by Baumeister (1992). Compared with the findings of most of those authors and Henderson et al. (2018), maximum movements of the deer in my study were an order of magnitude greater. A range of elevation-use strategies was appar- ent. Use of minimum elevations was similar among individuals, but maximum elevations varied con- siderably. Most deer activity was along the floor of the main valley in which they wintered. When deer left that valley, movements typically followed the floors of the tributary valleys in which they travelled. However, the higher elevations of those tributaries, along with some limited forays from the main val- ley to adjacent mountain slopes, were associated with several other patterns of elevation use. Some deer oc- curred at high elevations for at least part of the sum- mer, and some moved to relatively high elevations during early winter. Relatively long-distance movement by deer in this study area may have reflected the abundance and broad distribution of moderate- to high-elevation summer habitats and the limited elevation gradient at valley- floor positions, such that any deer gaining the advan- tage of following the wave of greening vegetation up- slope (Mysterud 2013; Merkle et al. 2016, Middleton et al. 2018) without leaving valley floors would be obliged to move considerable distances in this land- scape. The existence of a wintering population in an elevated valley proximal to the Trench is notable. The Trench is as close as 15 km to summer and winter ac- tivity (Figure 1), is accessible via several passes or downstream movement, is at lower elevations with less snow and warmer winter temperatures, includes agricultural fields and extensive riparian areas, and 2019 is notable for an abundant large-mammal fauna and high-quality winter range (Benson 1970) includ- ing for deer. Delaying movement to winter ranges is advantageous to some ungulates (Mysterud 2013), yet White-tailed Deer are poorly adapted to snow (Stelfox and Taber 1969; Telfer and Kelsall 1984). For deer in my study, further travel through snow to reach the Trench may be prohibitive late in the season de- spite apparently higher-quality habitat used by other deer in the Trench. Alternatively, the shorter return distance to summer habitats and the ability to more precisely gauge the initiation of green-up may offer advantages to remaining within the mountains dur- ing winter. Additional collaring on summer ranges within this region of the Rocky Mountains would indicate whether some White-tailed Deer summering there do seasonally join other deer in the Trench. Deer wintering in the upper Kootenay River val- ley occurred in Kootenay and Yoho national parks in BC, Banff National Park and Spray Valley Provincial Park in Alberta, and both private land and non-park Crown land in BC. For a species as resilient as White- tailed Deer (Halls 1978), a lack of protective man- agement across jurisdictions is unlikely to have the severe population effects experienced by many mi- gratory ungulates (Bolger et al. 2008). However, management goals for resource extraction, fire, deer hunting, predator hunting and trapping, ecological in- tegrity, and recreation have the potential to constrain or enhance deer populations and movements. Cross- jurisdictional differences may influence the ability of any agency to achieve its wildlife or ecosystem ob- jectives. For example, managing predators or enhan- cing habitat to benefit deer would have less effect if done only on a portion of the population’s annual range, and maintaining a naturally functioning sys- tem may be affected by activities on other land bases, such as hunting during the “antlerless” deer season on provincial and private land. It would be prudent for resource managers to coordinate their efforts with nearby jurisdictions or at least consider the ef- fect of extensive seasonal movements when manag- ing White-tailed Deer in the Rocky Mountains. Acknowledgements Many Parks Canada Agency staff, students, and volunteers helped capture, monitor, retrieve collars, or manage data. I thank Jenny Burrows, Blair Fyten, Matt Kennedy, Justin Kinnersley, Isabel McFetridge, Sonia Nicholl, Tom Niddrie, Margaret Pak, Darren Quinn, Adam Sherriff, Geoff Skinner, Brian Spread- bury, Natalie Stafl, Kate Williams, Shelagh Wrazej, and Anna Yuill. I particularly appreciate the cheer- ful company and dedicated cold-weather trapping of Laura Kroesen and Patrick Langan, the provision of KINLEY: SEASONAL MOVEMENTS OF WHITE-TAILED DEER 251 roadside survey data by Shelagh Wrazej, and the geo- matics talent of Dan Teleki, all of Parks Canada in Radium Hot Springs, British Columbia. Alan Dibb and two anonymous reviewers provided helpful comments on earlier drafts. Permits were issued by the British Columbia Ministry of Natural Resource Operations and by Parks Canada’s animal care com- mittee. Capture and handling protocols were ap- proved under research permit KOONP-2011-7812 (and subsequent renewals) and animal care permit 8979 for work conducted in Kootenay National Park, and under Wildlife Act permit CB12-76585 and CB13- 92061 for work on provincial land. Project funds were provided by Parks Canada Agency through its Conservation and Restoration Program. Literature Cited Baumeister, T.R. 1992. Ecology of white-tailed deer on summer-fall range in northcentral Idaho. M.Sc. thesis, Montana State University, Bozeman, Montana, USA. Benson, W.A. 1970. The land capability analysis of the East Kootenay area (annotated map). Queen’s Printer, Ottawa, Ontario, Canada. Berger, J. 2004. The last mile: how to sustain long-distance migration in animals. Conservation Biology 18: 320— 331. https://doi.org/10.1111/).1523-1739.2004.00548.x Bolger, D.T., W.D. Newmark, T.A. Morrison, and D.F. Doak. 2008. The need for integrative approaches to understand and conserve migratory ungulates. Ecology Letters 11: 63-77. https://doi.org/10.1111/).1461-0248.20 07.01109.x Brinkman, T.J., C.S. DePerno, J.A. Jenks, B.S. Har- oldson, and R.G. Osborn. 2005. Movement of fe- male white-tailed deer: effects of climate and intensive row-crop agriculture. Journal of Wildlife Management 69: 1099-1111. https://doi.org/10.2193/0022-541 X(2005) 069[1099: mofwde]2.0.co;2 Caulkett, N.A., P.H. Cribb, and J.C. Haigh. 2000. Com- parative cardiopulmonary effects of carfentanil—xyla- zine and medetomidine—ketamine used for immobil- ization of mule deer and mule deer/white-tailed deer hybrids. Canadian Journal of Veterinary Research 64: 64-68. Demarais, S., K.V. Miller, and H.A. Jacobson. 2000. White-tailed deer. Pages 601-628 in Ecology and Management of Large Mammals in North America. Edited by S. Demarais and P.R. Krausman. Prentice- Hall, Upper Saddle River, New Jersey, USA. Fieberg, J., D.W. Kuehn, and G.D. DelGiudice. 2008. Understanding variation in autumn migration of north- ern white-tailed deer by long-term study. Journal of Mammalogy 89: 1529-1539. https://doi.org/10.1644/ 07-mamm-a-277.1 Grovenburg, T.W., C.N. Jacques, R.W. Klaver, C.S. DePerno, T.J. Brinkman, C.C. Swanson, and J.A. Jenks. 2011. Influence of landscape characteristics on migration strategies of white-tailed deer. Journal of Mammalogy 92: 534-543. https://doi.org/10.1644/ 09-mamm-a-407.1 252 Halls, L.K. 1978. White-tailed deer. Pages 43-65 in Big Game of North America: Ecology and Management. Edited by J.L. Schmidt and D.L. Gilbert. Stackpole Books, Harrisburg, Pennsylvania, USA. Henderson, Jr., C.R., M.S. Mitchell, W.L. Meyers, P.M. Lukacs, and G.P. Nelson. 2018. Attributes of seasonal home range influence choice of migratory strategy in white-tailed deer. Journal of Mammalogy 99: 89-96. https://doi.org/10.1093/jmammal/gyx148 Hoekman, S.T., G.L. Dusek, and A.K. Wood. 2006. Spatial relationships and resource selection. Pages 109-186 in Ecology of White-tailed Deer in the Salish Mountains, Northwest Montana. Montana Fish, Wildlife & Parks, Wildlife Division, Helena, Montana, USA. Merkle, J.A., K.L. Monteith, E.O. Aikens, M.M. Hayes, K.R. Hersey, A.D. Middleton, B.A. Oates, H. Saw- yer, B.M. Scurlock, and M.J. Kauffman. 2016. Large herbivores surf waves of green-up during spring. Pro- ceedings of the Royal Society B 283: 20160456. https:// doi.org/10.1098/rspb.2016.0456 MFLNRORD (Ministry of Forests, Lands, Natural Resource Operations and Rural Development). 2018. 2018-2020 hunting and trapping regulations synopsis. Black Press Media, Victoria, British Columbia, Canada. Middleton, A.D., J.A. Merkle, D.E. McWhirter, J.G. Cook, R.C. Cook, P.J. White, and M.J. Kauffman. 2018. Green-wave surfing increases fat gain in a migra- tory ungulate. Oikos 127: 1060-1068. https://doi.org/10. 1111/oik.05227 Morgan, J.T. 1993. Summer habitat use of white-tailed deer on the Tally Lake Ranger District, Flathead Na- tional Forest. Ph.D. thesis, Montana State University, Bozeman, Montana, USA. Mysterud, A.T. 2013. Ungulate migration, plant phenol- ogy, and large carnivores: the times they are a-changin’. Ecology 94: 1257-1261. https://doi.org/10. 1890/12-0505.1 Nelson, M.E., L.D. Mech, and P.F. Frame. 2004. Tracking of white-tailed deer migration by global positioning sys- THE CANADIAN FIELD-NATURALIST Vol. 133 tem. Journal of Mammalogy 85: 505-510. https://doi. org/10.1644/bos-120 Robinson, H.S., J.C. Gwilliam, and I. Parfitt. 2002. Movements, survival and mortality of white-tailed deer in the Pend d’Oreille River valley. Columbia Basin Fish and Wildlife Compensation Program, Nelson, British Columbia, Canada. Sabine, D.L., S.F. Morrison, H.A. Whitlaw, W.B. Bal- lard, G.J. Forbes, and J. Bowman. 2002. Migration behavior of white-tailed deer under varying winter climate regimes in New Brunswick. Journal of Wild- life Management 66: 718-728. https://doi.org/10.2307/ 3803137 Secord, M.L. 1994. Winter habitat use, migration, and spring and summer use of clearcuts by white-tailed deer in the Priest Lake watershed of northern Idaho. M.Sc. thesis, University of Montana, Missoula, Montana, USA. Stelfox, J.G., and R.D. Taber. 1969. Big game in the north- ern Rocky Mountain coniferous forest. Pages 197-222 in Coniferous Forests of the Northern Rocky Mountains: Proceedings of the 1968 Symposium. Edited by R.D. Taber. University of Montana Foundation, Missoula, Montana, USA. Stewart, K.M., R.T. Bowyer, and P.J. Weisberg. 2011. Spatial use of landscapes. Pages 181-218 in Biology and Management of White-tailed Deer. Edited by D.G. Hewitt. CRC Press, Boca Raton, Florida, USA. https:// dot.org/10.1201/9781482295986 Telfer, E.S., and J.P. Kelsall. 1984. Adaptation of some large North American mammals for survival in snow. Ecology 65: 1828-1834. https://doi.org/10.2307/1937779 VerCauteren, K., J. Beringer, and S.E. Hyngstrom. 1999. Use of netted cage traps for capturing white-tailed deer. Pages 155-164 in Mammal Trapping. Edited by G. Proulx. Alpha Wildlife Research and Management Ltd., Sherwood Park, Alberta, Canada. Received 7 January 2019 Accepted 18 December 2019 The Canadian Field-Naturalist Sharp-tailed Grouse (7ympanuchus phasianellus) population dynamics and restoration of fire-dependent northern mixed-grass prairie RosBert K. Murpuy'?" and KAREN A. SmitH’? 'United States Fish and Wildlife Service, Des Lacs National Wildlife Refuge Complex, Kenmare, North Dakota 58746 USA *Current address: Eagle Environmental, Inc., 12 Longview Road, Sandia Park, New Mexico 87047 USA 3Current address: 8210 88th Street NW, Kenmare, North Dakota 58746 USA “Corresponding author: murph@eagleenvironmental.net Murphy, R.K., and K.A. Smith. 2019. Sharp-tailed Grouse (Zympanuchus phasianellus) population dynamics and restora- tion of fire-dependent northern mixed-grass prairie 133(3): 253-262. https://doi.org/10.22621/cfn.v133i3.2115 Abstract Case studies of Sharp-tailed Grouse (7ympanuchus phasianellus) population dynamics before and during re-introduction of fire to northern mixed-grass prairies that lacked fire for multiple decades are unavailable. At a 108-km? northern mixed- grass prairie refuge in North Dakota, fire was suppressed from the early 1900s through late 1970s. Nine management units (total area 16.8 km’, 15.7% of the refuge) received initial prescribed fire treatments during 1979-1984. The mean annual density of male Sharp-tailed Grouse attending leks on these units during 1981-1985 (9.0 males/km7) was twice that on the same units during 1961-1965 (4.2 males/km7’), amid the fire exclusion era; nonoverlap of 90% CIs encompassing the means suggested a significant treatment effect. However, densities of males on units managed without prescribed fire during 1961— 1965 and 1981-1985 did not change between the two periods. By 1987, fire had been re—introduced to >50% of the refuge overall. Mean annual abundance (i.e., total numbers) of lekking males on the entire refuge did not differ between 1961-1965 and 1981-1985 but was significantly greater during 1989-1993 than during 1961-1965 and 1981-1985. Changes in density and abundance of lekking males coincided with fire-induced reductions in woody cover and increases in herbaceous cover. Our study illustrates the marked capacity of Sharp-tailed Grouse to respond to reductions of tree and shrub cover resulting from prescribed fire in northern mixed-grass prairie and the species’ attraction to habitat disturbance in general. Key words: Sharp-tailed Grouse; 7ympanuchus phasianellus, northern mixed-grass prairie; prairie management; prescribed fire; North Dakota Introduction During the 1900s, tree and shrub cover increased markedly on present-day national wildlife refuges in the northern mixed-grass prairie (NMGP) region of North America’s Great Plains, mainly due to fire sup- pression (Grant and Murphy 2005). Fire-intolerant Trembling Aspen (Populus tremuloides Michaux), Silverberry (Elaeagnus commutata Bernhardi ex Rydberg), and Western Snowberry (Symphoricarpos occidentalis Hooker) were common tree and shrub species that proliferated. Some species of grassland- dependent passeriform birds that bred on the refuges became rare or absent in areas invaded by trees and shrubs (Madden et al. 1999; Grant et al. 2004; Murphy and Smith 2007). Sharp-tailed Grouse (7yvmpanuchus Phasianellus), a conspicuous member of the NMGP breeding bird community, may abandon landscapes that become invaded by trees and shrubs in the ab- sence of fire, e.g., in Aspen Parkland (Caldwell 1976; Moyles 1981; Berger and Baydack 1992). However, there are no published reports of changes in num- bers of the species during years encompassing pre- scribed fires to restore grassland landscapes invaded by woody vegetation in the NMGP region. While compiling a case study on the manage- ment of 108-km? Lostwood National Wildlife Refuge (LNWR), centred in the NMGP region, we found that data on annual counts of Sharp-tailed Grouse and concurrent records of fire re-introduction to the area after roughly 60 years of fire exclusion could con- tribute to the knowledge of fire’s role in Sharp-tailed Grouse management. Moreover, the case study could be supported by published data on changes in domin- ance of woody versus herbaceous vegetation associ- ated with prescribed fire on the refuge (Madden et al. 1999; Murphy and Smith 2007; Smith and Murphy 2007) given a major refuge management objective: to restore the historic (before settlement by Euro- 253 ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0) 254 American peoples) balance of woody versus herb- aceous vegetation to favour native grassland bird spe- cies and other native wildlife (U.S. Fish and Wildlife Service 1998). Our first study objective, carried out at a local spatial scale, was to compare the density (_e., number/km7?) of lekking male Sharp-tailed Grouse on prairie management units during 1961-1965, amid the fire exclusion era, to the density of lekking males on the same units under initial prescribed fire treat- ments during 1981-1985. Our second study objective, pursued at a landscape scale, was to compare overall abundance (i.e., total number) of lekking male Sharp- tailed Grouse on all of LNWR among three 5-yr per- iods: (1) 1961-1965, amid the fire exclusion era; (2) 1981-1985, the initial fire re-introduction period; and (3) 1989-1993, after >50% of the refuge had been treated by prescribed fire at least once. As part of this objective, we also examined relationships between abundance or density of lekking male Sharp-tailed Grouse and fire history, 1.e., number of fires con- ducted, across management areas of the entire refuge. Methods LNWR, in Burke and Mountrail counties, north- western North Dakota (48.617°N, 102.450°W), is roll- ing to hilly native prairie (55% of refuge area) and tame grasslands (21%) interspersed with about 4000, 0.1- to 224-ha wetlands (20%; Murphy 1993). Before settlement by Euro-Americans in the early 1900s, the upland landscape was mixed-grass prairie dom- inated by needlegrasses (Nassella viridula [Trinius] Barkworth, Hesperostipa comata Oldham and Brinker, Heterostipa spartea |Trinius] Barkworth), Western Wheatgrass (Pascopyrum smithii [Rydberg] A. Léve), Blue Grama (Bouteloua gracilis [Kunth] Lagasca ex Griffiths), and a variety of native forb species (Barker and Whitman 1988). Shrubs probably covered ~5% of the uplands and trees were rare (U.S. Soil Conservation Service 1975). Patches of tree-size Trembling Aspen began to appear by the 1930s (as de- tected in aerial photos) after 10-20 years of active fire suppression (Murphy 1993). The density and mean size of aspen patches on LNWR increased from 1.5 patches/km? and 0.13 ha in the mid—1930s, when the refuge was established, to 4.8 patches/km? and 0.36 ha in the early 1980s, respectively. Shrub cover dom- inated by Western Snowberry increased from ~25% in the mid—1930s to >50% by the early 1980s. Counts of lekking male Sharp-tailed Grouse can yield reliable population indices if done within nar- row constraints with bias accounted for (Drummer et al. 2011). All Sharp-tailed Grouse leks on LNWR were located systematically in early spring annually during 1961-1965 and 1981-1993, following standard guidelines (Kirsch 1956). Each lek had at least two THE CANADIAN FIELD-NATURALIST Vol. 133 males by definition. To maximize accuracy, males were counted two to four times on each lek between 0.5 h before sunrise to 2 h after sunrise during mid- April through early-May, encompassing the peak per- iod of lek attendance by breeding hens in the region (Connelly et al. 1998). Males on some leks could be counted by observation with binoculars from nearby hilltops. Most leks, however, were higher than their surroundings such that some or most males could not be viewed directly from any single location. In such a case, the observer crept to within ~3—-10 m of the lek’s edge and listened to determine whether females were present; acoustic displays by male Sharp-tailed Grouse on leks are distinctly more intense when fe- males are there (Connelly et al. 1998). If females were present, they would flush immediately when the ob- server stood slowly. In contrast, males typically hesi- tated to flush for several seconds after all females flushed, then flushed and flew together in a cohesive flock. The flocked males typically could be counted by the observer at least twice before landing or flying from view. The same procedure would be followed at the respective lek on at least one other morning until counts of total males on the lek were consistent among mornings. To address our first objective of comparing the density of lekking male Sharp-tailed Grouse on man- agement units during 1961-1965 to the density of lekking males on the same units during initial pre- scribed fire treatments ~20 years later, we used con- current changes in densities of lekking males on units not prescribe-burned as a baseline for comparison. Relying on refuge records, we categorized the 1940— 1985 management history of units as either (1) grazed by cattle <19 years; (2) grazed by cattle >29 years; (3) prolonged rest; (4) treated by prescribed fire dur- ing 1979-1984; or (5) burn perimeter (Table 1, Figure 1). Prescribed fires at LNWR were conducted by using a surround technique, described in Murphy and Smith (2007). Most fires consumed >80% of above- ground vegetation (Table 2). We categorized the area within 0.8 km of burn units as burn perimeter (a mix of grazed <19 years, grazed >29 years, and prolonged rest categories; Figure 1) because prescribed fires could indirectly influence densities of lekking male Sharp-tailed Grouse on adjacent management units, and 0.8 km approximates the mean distance between the species’ leks and nest sites (reviewed in Connelly et al. 1998). We could not formally test for differences in density of lekking male Sharp-tailed Grouse among management history categories because manage- ment treatment types were not assigned randomly to the various units, and prolonged rest and burn per- imeter categories were represented by only one and 2019 MURPHY AND SMITH: PRESCRIBED FIRE AND SHARP-TAILED GROUSE 255 TABLE 1. Management history of prairie management units at Lostwood National Wildlife Refuge in northwestern North Dakota during 1940-1985. Number of Total area Unit category mais (km?) Management history description Grazed <19 years’ 3 13.8 Grazed by cattle at light stocking rates (0.4—0.6 Animal Unit Months/ha) during July—October every 1—4 years during 1940-1979 and rested during 1980-1985 Grazed >29 years’ S 16.0 Grazed by cattle at light stocking rates (0.4—0.6 Animal Unit Months/ha) during July—October every 1—2 years during 1940-1979 and rested during 1980-1985 Prolonged rest 1 7.1 Not grazed or prescribe-burned Burned 1979-1984 9 16.8 Rested and periodically grazed 1940-1978 then prescribe-burned one, two, or three times in late spring or summer during 1979-1984, with 2—3 years between prescribed fires on units burned two or three times* Burn perimeter 2 19.2 Not prescribe-burned; a mix of grazed <19, grazed >29, and prolonged rest categories *Range 11-18 years. ‘Range 30-36 years. ‘Table 2 presents detailed 1979-1984 fire treatment history for each unit. coo Lakes (excluded) Miscellaneous areas (excluded) Grazed <19 (11-18) years Grazed >29 (30-36) years Prolonged Rest Burned 1979-1984 (unit numbers indicated) SOC f Va = x Burn Perimeter Figure 1. Location and management history of prairie management units on Lostwood National Wildlife Refuge in north- western North Dakota as of 1981-1985, when density (individuals/km7’) of male Sharp-tailed Grouse (Zympanuchus pha- sianellus) displaying on leks in spring was documented annually on units of five management categories. These were compared to densities of lekking males documented on the respective units during 1961-1965. The refuge area south of Highway 50 was excluded from the comparison of densities as it was open to hunting of Sharp-tailed Grouse. Inset map (upper right) indicates the study area (black dot) in relation to North America’s northern mixed-grass prairie region (grey shaded). two units, respectively (Table 1). Our conclusions were thus limited. We considered, however, that ten- tative evidence of a treatment effect might be implied for a given management history category if 90% CIs around the respective 1961-1965 and 1981-1985 mean densities did not overlap. Moreover, our ap- proach to assessing male Sharp-tailed Grouse density in relation to prescribed fire included counts of lek- king males on all prescribed fire units each spring, such that in a given spring, residual vegetation was 256 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Size, year of treatment, and thoroughness of burns on prairie management units treated by prescribed fire on Lostwood National Wildlife Refuge in northwestern North Dakota during June, July, or August, 1979-1984. Unit number Area (km7’) 1979 1980 1 1.0 ct 2 3.7 b 3 5.0 4 0.5 a 5 0.9 c 6 1.0 7 0.4 8 0.5 9 3.8 Year 1981 1982 1983 1984 Cc c Cc c c b Cc a b ¢ b Cc Cc Cc Cc *A pproximate percentage of above-ground live and residual vegetation removed by prescribed fire: (a) 35-50%; (b) 51-80%; (c) >80%. limited on units that had been burned the previous summer. This provides for a conservative picture of the species’ response to prescribed fire because in the first spring after summer fires, vegetation on LNWR is relatively short and sparse (Madden et al. 1999), less than optimal for females seeking nest sites. Because male Sharp-tailed Grouse compete for space on leks near areas frequented by females seek- ing ideal nesting cover (Gratson 1988), lek attendance by males is likely to be reduced during the first spring after summer fires. We acknowledge that small num- bers of male Sharp-tailed Grouse may not attend leks in some years or do so infrequently (Gratson et al. 1991) but believe this possibility would negligibly in- fluence our conclusions. During 1986-1993, following the initial pre- scribed fires, incrementally greater proportions of LNWR were treated by prescribed fire such that units lacking prescribed fire histories including burn per- imeter became less and less available. This change hindered longer-term comparison of male Sharp- tailed Grouse density among management history categories. We could, however, assess landscape-level changes in abundance (i.e., total numbers) of lekking male Sharp-tailed Grouse across all of LNWR in re- lation to prescribed fire. For our second objective, we sought to examine abundance of lekking males in the first five-year period after 1985 during which >50% of the refuge was treated with prescribed fire. We also sought a five-year period during which mean annual precipitation, from one year before the start to one year before the end of the period, was similar to that in the 1961-1965 baseline (40.2 cm) and 1981-1985 initial prescribed fire periods (41.7 cm). For a given year, we considered precipitation level in the previ- ous year as a key potential confounding factor in our comparison because it can markedly influence sur- vival of Sharp-tailed Grouse in general (Cartwright 1944). The years 1989-1993 met these two criteria (55-64% of refuge burned; 40.0 cm mean annual precipitation during 1988-1992), except that an un- naturally severe wildfire burned a 22.7-km? unit of the refuge in August 1988. We considered the wild- fire an anomaly because it occurred when there was an abnormally high buildup of residual vegetation combined with ambient temperature, relative humid- ity, fuel moisture, and windspeed conditions far ex- ceeding bounds for conducting prescribed fires (U.S. Fish and Wildlife Service unpubl. data). The wildfire burned into the humus layer and in many places to mineral soil, removing all residual and live, above- ground herbaceous and woody vegetation except for ~40% of tree-size Trembling Aspen and scant patches of Western Snowberry. Therefore, we replaced the following spring’s (1989) count of lekking males with like data from spring 1987, the only year following prescribed fire treatment (July 1986) of the area, al- though total precipitation in the previous year (1986) was slightly greater (45.7 cm) than the 1988-1992 mean. Last, we considered the August 1988 wildfire to be, in effect, a 1989 prescribed fire with regard to its influence on residual vegetation and numbers of lekking male grouse the following spring. We therefore consider the 1989-1993 period as a landscape-level prescribed fire period (hereafter the Landscape Fire period). In a similar vein, we here- after refer to the 1961-1965 period, that neared the end of ~60 years of fire suppression, as the Fire Exclusion period, and the 1981-1985 years associated with initial prescribed fire treatments as the Initial Fire period. During the mid-1980s, LNWR was partitioned into 20 “management blocks” (MBs) averaging 4.7 km? in area (SE = 1.0, range = 0.6—22.7). To assess whether total abundance of Iekking male Sharp- tailed Grouse on LNWR changed between any two 2019 of the three time periods, we first combined small (<5 km/7), adjoining MBs into five MB groups composed of two to four MBs each. Each of five other “groups” were represented by one large MB. Thus, a total of 10 MB groups were available to provide adequate sample sizes for a matched-pairs analyses of tem- poral change in abundance of Iekking male grouse. We next summed the total number of Iekking males in each MB group, for each year in each of the three time periods, then calculated the mean annual abun- dance of lekking males for each MB group by period. To test for a difference in total abundance between two given periods, we paired the periods’ annual means for each MB group and used the non-paramet- ric Wilcoxon matched-pairs signed-ranks test (Daniel 1990) to assess whether overall means of the paired observations differed. Specifically, we used a two- tailed version of the test with a set at 0.1 to deter- mine whether total abundance of lekking male Sharp- tailed Grouse differed between (1) the Fire Exclusion and Initial Fire periods, (2) Initial Fire and Landscape Fire periods, and (3) Fire Exclusion and Landscape Fire periods. A one-tailed test would seem appro- priate based on knowledge that Sharp-tailed Grouse abundance can increase when prescribed fire is incor- porated into the habitat disturbance regime (Kirsch and Kruse 1972; Kirsch et al. 1973). However, one- tailed Wilcoxon tests performed with small sizes, as in our case (10 matched pairs), can result in incorrect P-values (Mundry and Fischer 1998), so we used the more conservative two-tailed approach. Before con- ducting the tests, we plotted distributions of the dif- ferences between pairs and found the distributions to be reasonably symmetrical, an assumption of the Wilcoxon signed-ranks test (Daniel 1990). Finally, we categorized each MB by the num- ber of prescribed fires (1.e., fire history) applied to it during 1979-1992, from none up to four (Figure 2). However, most area covered by two MBs in a three or four burns category also was treated with inten- sive, prolonged grazing by cattle during two and three (respectively) late spring-early summer seasons of the Landscape Fire period. So, we placed them in a unique fire history category named “3 or 4 Burns plus Grazing” (3+G in Figure 2). The 22.7-km/? unit that experienced a severe wildfire in August 1988 was placed in its own MB category, named “1 Burn plus 1 Wildfire” (1+W), because the wildfire event distinguished it from other MBs; this MB also was unique due to its large size, nearly four times larger than any other. We used non-overlap of 90% CIs en- compassing means as tentative evidence of differ- ences in abundance of Iekking male Sharp-tailed Grouse among fire history categories in different per- iods. We also calculated period-specific densities of MURPHY AND SMITH: PRESCRIBED FIRE AND SHARP-TAILED GROUSE 257 lekking male Sharp-tailed Grouse for each fire his- tory category. Results Based on non-overlapping 90% CIs (Figure 3), the mean annual density of lekking male Sharp-tailed Grouse on prairie management units that were treated by prescribed fire during the Initial Fire period was more than double what it was on the same units ~20 years earlier, during the Fire Exclusion period. Meanwhile, densities of lekking males did not dif- fer between the periods on rested, grazed, and burn perimeter units, based on substantial overlap among 90% Cls (Figure 3). Across all of LNWR, the mean annual abundance of lekking male Sharp-tailed Grouse did not differ be- Prescribed fire history categories of Management Blocks (number of blocks in parentheses): 0 O Burns (6) 1 1 Burn (3) 1+W 1 Burn plus 1 Wildfire (1) 2 2 Burns (4) 3 3 or 4 Burns (4) 3+G 3or4 Burns plus Grazing (2) FiGure 2. Number of prescribed fires applied during 1979— 1992 on management blocks at Lostwood National Wildlife Refuge in northwestern North Dakota. Wildfires had been effectively suppressed on the area before 1979. Management block boundaries were designated during the mid-1980s. 258 © 1961-1965 @ 1981-1985 Mean annual density of lekking males (n/km7) Management history category FiGurE 3. Mean annual densities (individuals/km7) of lek- king male Sharp-tailed Grouse (7ympanuchus phasianel- lus) at Lostwood National Wildlife Refuge in northwestern North Dakota during 1961-1965 and 1981-1985 on multiple prairie management units of each of five management his- tory categories (see Table 1); density was assessed on the same units of each category during both time periods. After ~60 years of fire suppression, prescribed fire was re-intro- duced to the refuge during 1979-1984, specifically on units in the Burn 1979-1984 category. Error bars are 90% CIs. tween the Fire Exclusion and Initial Fire periods (W* = 23.5,n= 10, P=0.70; Figure 4). However, mean an- nual abundance of Ilekking male Sharp-tailed Grouse across all of LNWR was significantly greater dur- ing the Landscape Fire period than during the Fire Exclusion (W* = 7, n = 10, P = 0.04) and Initial Fire periods (W* =2, n= 10, P= 0.006; Figure 4). A 32% increase in mean annual abundance from the Initial Fire to Landscape Fire periods coincided with a sub- stantial rise in the percentage of LNWR treated by prescribed fire at least once, from 6.0-15.7% during 1979-1984 to 53.4—63.1% during 1987-1992 (Figure 4). Up to 1185 lekking males were recorded in one year (1992), representing an overall density of 12.6 males/km? (based on 94.2 km/? total refuge area ex- cluding major lakes). Most of the increase in total numbers arose from MBs burned once or twice be- tween 1979 and 1992 (Figure 5a). During this per- iod, density of lekking males seemed consistently high across all categories of number of fires experi- enced (range of means = 9.7 to 10.7 males/km7; range THE CANADIAN FIELD-NATURALIST Vol. 133 of CIs, + 1.7 to+2.7), except for the “3 or 4 Burns plus Grazing” category, where no lekking males were de- tected, and the “O Burns” category (Figure 5b). Discussion Our case study of the re-introduction of fire to a large tract of NMPG deprived of fire for >60 years and associated changes in density and abundance of lekking male Sharp-tailed Grouse is unique for the NMGP region. It may well illustrate the spe- cies’ marked capacity to respond to the reduction of trees and shrubs by prescribed fires in a prairie eco- system, and to major perturbations in general. During the 1981-1985 Initial Fire period, the mean annual density of lekking males was about two-fold greater on prairie management units receiving initial pre- scribed fire treatments than on the same units during the Fire Exclusion period two decades earlier, when the units had been managed by prolonged rest or rest and grazing. This increase occurred even though the Initial Fire dataset included many counts of males on units treated by prescribed fire in previous sum- mers. Scattered unburned patches of vegetation may have attracted nesting females to such areas; un- burned “skips” comprised a mean of 14.5% of three units burned during 1982-1984 (Kruse and Piehl 1984). Moreover, the lack of change in male densities between the Fire Exclusion and Initial Fire periods on all but the burn units suggests that the increased density of lekking male Sharp-tailed Grouse on burn units can be attributed to the general growth in bird numbers on those units rather than just shifts in loca- tions of males from other units. In contrast with the markedly increased density of male Sharp-tailed Grouse between the Fire Exclusion and Initial Fire periods on management units in the burn category, we found no evidence of concur- rent change in male density on units categorized as either prolonged rest, <19 years grazed, or >29 years grazed. Under the latter management regimes, how- ever, densities likely would have declined during time intervals exceeding two decades, as trees and shrubs continued to displace grass- and forb-domin- ated prairie. Indeed, in the absence of fire for roughly six decades, much of the NMGP refuge had become aspen parkland, with some 2.59-km? sections having >15 aspen tree patches (Murphy 1993). In southern Manitoba, Berger and Baydack (1992) documented a severe decline in the number of Sharp-tailed Grouse leks as prairie gradually transformed into Trembling Aspen-dominated forest during only 21 years of fire suppression; on average, the birds abandoned a given lek if forest cover within 1 km exceeded 56%. Tree-size Trembling Aspen cover at LNWR aver- aged far less, only about 0.6% in 1969 and 0.7% in 2019 MURPHY AND SMITH: PRESCRIBED FIRE AND SHARP-TAILED GROUSE 299 1200 70 [E35] nMales — —% Burned 5 “3 1000 60 2 E 5 bo 580.8 mi = 50 6 we — $ 800 5 ee wo 0 40 § 2 600 S 5 30 3 c — = °o £ 400 8 be 20 g = om E : Cc & 200 10 € rs 0 0) Fire Exclusion: 1961-1965 Initial Prescribed Fire: 1981-1985 Landscape-level Prescribed Fire: 1989-1993 Figure 4. Change in annual abundance, i.e., total number, of male Sharp-tailed Grouse (Zympanuchus phasianellus) attending leks in spring at Lostwood National Wildlife Refuge, northwestern North Dakota during three historic time per- iods in relation to percentage of the refuge treated by prescribed fire at least once beginning in 1979, after ~60 years of fire suppression. Prescribed fires were conducted during late spring through summer. Due to an artificially severe wildfire on a 22.7-km* management block in August 1988, the 1989 count of lekking males was replaced by like data from 1987, follow- ing the management block’s first prescribed fire in 1986 (see text). Above each five-year group of columns is the respective group mean + | SE. 1985 (Figure 4 in Grant and Murphy [2005]). Thus, the near-complete removal of aspen trees at LNWR via burning or combinations of grazing and burning (Smith and Murphy 2007) probably contributed less to increases in male Sharp-tailed Grouse abundance than did the conversion of much shrub cover (mainly Western Snowberry and Silverberry) to grass-forb cover types via prescribed fire (Madden et al. 1999). Such a conversion has been critical in restoring other key members of the refuge’s grassland bird commun- ity. For example, the endemic Sprague’s Pipit (Anthus spragueii) and Baird’s Sparrow (Ammodramus bairdii) were absent and rare, respectively, on two 90-ha tracts rested and lightly grazed for >40 years but reappeared and increased after four prescribed fires were conducted during a ~16-year period. These changes coincided with a shift in grass-dominated cover from 45% to 84% and a 33% reduction in over- all vegetation height and density (Murphy and Smith 2007). The increase in Sharp-tailed Grouse density on management units undergoing initial prescribed fire treatments in our case study was consistent with a 32% increase in total abundance of males at the landscape scale during the Landscape Fire period, when >50% of the refuge had been burned at least once. Historically, the fire return interval for the eastern, more mesic part of the NMGP that encom- passes LNWR averaged roughly six years (Bragg 1995; Madden et al. 1999). A mosaic of manage- ment units under short (2—4 years) and moderate (5—7 years) fire return intervals seems optimal for most na- tive grassland bird species at LNWR (Madden et al. 1999). The heterogenous structure and general com- position of vegetation in units managed under these fire return intervals (Madden et al. 1999, 2000) may be ideal for Sharp-tailed Grouse as well. Indeed, the mean annual density of Ilekking males during the Initial Fire period was high on units treated by pre- scribed fire even though most units were burned two to three times with only 1-2 years between treat- ments. Although Sharp-tailed Grouse have a gen- eral affinity for frequent disturbance and early suc- cessional stages (Kirsch et al. 1973; Connelly et al. 1998), the consistently high mean annual densities of lekking males across MBs of all fire history categor- ies (excluding MBs that also were intensively grazed; Figure 5b) during the Landscape Fire period, sug- gest that the bird’s abundance in NMGP does not ne- cessarily increase with increasing “fire experience” (sensu Madden et al. 1999), at least during 10- to 15- year periods. Conclusions about effects of a given fire return interval on the structure and general composition of NMGP—and thus on attractiveness of the habitat for Sharp-tailed Grouse—should be considered tenta- tive, particularly if other types of defoliation treat- ments are applied between prescribed fires. The type, frequency, duration, and intensity of any such treat- ments likely influence effects of a given fire return 260 01961-1965 Mean annual total number of lekking male Sharp-tailed Grouse THE CANADIAN FIELD-NATURALIST Vol. 133 B® 1989-1993 O Burns 1 Burn 1Burnplus 2 Burns 3o0r4 3or4 1 Wildfire Burns Burns plus Grazing wo 5a" = 2 —G 12 =e 10 >= ze ° o © Se ° S2 4 Cc © s © 93 c oO o wD 0 2% O Burns 1 Burn 1Burnplus 2 Burns 3or4 3or4 1 Wildfire Burns Burns plus Grazing Category of 1979-1992 fire history Figure 5. Mean annual abundance (i.e., total number of individuals, a) and density (i.e., number of individuals/km7, b) of male Sharp-tailed Grouse (Zympanuchus phasianellus) attending leks in spring at Lostwood National Wildlife Refuge, northwestern North Dakota: (1) during 1961-1965, amid ~60 years of fire suppression; and (2) during 1989-1993, by which time >50% of the refuge area had been prescribe-burned at least once, beginning in 1979. The refuge is divided into six cat- egories of management blocks based on numbers of fires experienced during 1979-1992. Thus, values during 1961-1965 reflect abundance and density before fire was re-introduced to the refuge. Each category of fire history is represented by two to six management blocks except the “1 Burn plus 1 Wildfire” (1+W) category, which is represented by a single but very large (22.7 km?) management block. Due to an artificially severe wildfire on this latter area in August 1988, the 1989 count of lekking males was replaced by like data from 1987, following the management block’s first prescribed fire in 1986 (see text). In addition to having multiple prescribed fires, management blocks in the “3 or 4 Burns plus Grazing” (3+G) cat- egory received intensive, prolonged grazing by cattle in spring and early summer for 2—3 years during 1988-1992, leaving little residual nesting cover for Sharp-tailed Grouse in subsequent springs. Error bars are 90% CIs. Asterisks indicate no lekking males detected. interval on prairie vegetation in general, e.g., by re- ducing fuel loads (Engle and Bidwell 2001). Also, in- fluences of various fire return intervals on vegeta- tion structure and composition may be confounded by the presence of Smooth Brome (Bromus inermis Leysser) and Kentucky Bluegrass (Poa pratensis L.), two introduced, cool-season grass species that have become pervasive in much of the NMGP (Romo ef al. 1990; Murphy and Grant 2005). These grasses ap- pear to be increasing regardless of prairie manage- ment treatment history (Ellis-Felege et a/. 2013; but see Kobiela et a/. 2017), a change that may reduce the availability and attractiveness of cover for Sharp- tailed Grouse. Our comparisons of Sharp-tailed Grouse density and abundance among the Fire Exclusion, Initial Fire, and Landscape Fire periods included some basic components of a before-after-control-impact (BACI) study design, but our overall case study was observa- tional in nature and lacked robust replication. Ideally, a Statistically valid experimental design with repli- cation across a more extensive landscape would be used to distinguish effects of habitat management on Sharp-tailed Grouse from confounding, non-manage- ment, factors, e.g., precipitation extremes. Aldridge et al. (2004) attempted this in an aspen parkland land- scape. Regardless, long-term monitoring of Sharp- tailed Grouse abundance at LNWR enables passive 2019 adaptive management of the species’ NMGP habitat (Aldridge et al. 2004). Acknowledgements We dedicate this paper to the memory of three northern prairie and prairie grouse management ex- perts: Michael Gratson, Leo Kirsch, and Arnie Kruse. During 1961-1965, counts of lekking male Sharp- tailed Grouse at Lostwood National Wildlife Refuge were conducted by Merrill Hammond, Ned Peabody, and Don White. Bob Danley and Ken Maruskie as- sisted us with lek counts during the 1990s. Discussions with Ryan Nielson (Eagle Environmental Inc.) en- hanced our approach to data analyses. We thank Todd Grant, two anonymous reviewers, and Associate Edi- tor Jenn Foote for many comments that markedly im- proved our manuscript. Findings and conclusions in this article are those of the authors and do not neces- sarily represent views of the United States Fish and Wildlife Service. Literature Cited Aldridge, C.L., M.S. Boyce, and R.K. Baydack. 2004. Adaptive management of prairie grouse: how do we get there? Wildlife Society Bulletin 32: 92-103. https://do1. org/10.2193/0091-7648(2004)32[92:amopgh]2.0.co;2 Barker, W.B., and W.C. Whitman. 1988. Vegetation of the Northern Great Plains. Rangelands 10: 266-272. Accessed 13 May 2019. https://journals.uair.arizona.edu/ index.php/rangelands/article/viewFile/10975/10248. Berger, R.P., and R.K. Baydack. 1992. Effects of aspen succession on Sharp-tailed Grouse, Zymphanuchus pha- sianellus, in the interlake region of Manitoba. Canadian Field-Naturalist 106: 185-191. Accessed 13 May 2019. https://biodiversitylibrary.org/page/34347405. Bragg, T.B. 1995. Climate, soils and fire: the physical envi- ronment of North American grasslands. Pages 49-81 in The Changing Prairie. Edited by K.H. Keeler and A. Joern. Oxford University Press, New York, New York, USA. Caldwell, P.J. 1976. Energetic and population considera- tion of Sharp-tailed Grouse in the Aspen Parkland of Canada. Ph.D. thesis, Kansas State University, Manhat- tan, Kansas, USA. Cartwright, B.W. 1944. The “crash” decline in Sharp- tailed Grouse and Hungarian Partridge in western Canada and the role of the predator. Transactions of the North American Wildlife Conference 9: 324-329. Connelly, J.W., M.W. Gratson, and K.P. Reese. 1998. Sharp-tailed Grouse (Zympanuchus phasianellus), ver- sion 2.0. Jn The Birds of North America. Edited by A.F. Poole and F.B. Gill. Cornell Lab of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.354 Daniel, W.W. 1990. Applied Non-parametric Statistics. PWS-Kent, Boston, Massachusetts, USA. Drummer, T.D., R.G. Corace, and S.J. Sjogren. 2011. Sharp-tailed Grouse lek attendance and fidelity in upper Michigan. Journal of Wildlife Management 75: 311-318. https://doi.org/10.1002/jwmg.42 Ellis-Felege, S.N., C.S. Dixon, and S.D. Wilson. 2013. MURPHY AND SMITH: PRESCRIBED FIRE AND SHARP-TAILED GROUSE 261 Impacts and management of invasive cool-season grasses in the Northern Great Plains: challenges and opportun- ities for wildlife. Wildlife Society Bulletin 37: 510-516. https://doi.org/10.1002/wsb.321 Engle, D.M., and T.G. Bidwell. 2001. The response of cen- tral North American prairies to seasonal fire. Journal of Range Management 54: 2-10. https://do1.org/10.2458/azu_ jrm_y54il_engle Grant, T.A., E. Madden, and G.B. Berkey. 2004. Tree and shrub invasion in northern mixed-grass prairie: im- plications for breeding grassland birds. Wildlife Society Bulletin 32: 807-818. https://doi.org/10.2193/0091-7648 (2004)032[0807:tasiin|]2.0.co;2 Grant, T.A., and R.K. Murphy. 2005. Changes in wood- land cover on prairie refuges in North Dakota, USA. Natural Areas Journal 25: 359-368. Gratson, M.W. 1988. Spatial patterns, movements, and cover selection by Sharp-tailed Grouse. Pages 158-192 in Adaptive Strategies and Population Ecology of Northern Grouse. Edited by A.T. Bergerud and M.W Gratson. University of Minnesota Press, Minneapolis, Minnesota, USA. Gratson, M.W., G.K. Gratson, and A.T. Bergerud. 1991. Male dominance and copulation disruption do not ex- plain variance in male mating success on Sharp-tailed Grouse (Zympanuchus phasianellus) \eks. Behaviour 118: 187-213. https://doi.org/10.1163/156853991x00283 Kirsch, L.M. 1956. Spring grouse census and habitat in- ventory methods. U.S. Fish and Wildlife Service, Wild- life Management Series Number 13. Kirsch, L.M., A.T. Klett, and H.W. Miller. 1973. Land use and prairie grouse population relationships in North Dakota. Journal of Wildlife Management 37: 449-453. https://doi.org/10.2307/3800307 Kirsch, L.M., and A.D. Kruse. 1972. Prairie fires and wildlife. Tall Timbers Fire Ecology Conference 12: 289— 303. Kobiela, B., J. Quast, C. Dixon, and E.S. DeKeyser. 2017. Targeting introduced species to improve plant com- munity composition on USFWS-managed prairie rem- nants. Natural Areas Journal 37: 150-161. https://do1.org/ 10.3375/043.037.0204 Kruse, A.D., and J.L. Piehl. 1984. The impact of pre- scribed burning on ground-nesting birds. Proceedings of the North American Prairie Conference 9: 153- 156. Accessed 13 May 2019. http://digicoll.library.wisc. edu/cgi-bin/EcoNatRes/EcoNatRes-idx?ty pe=article& did=EcoNatRes.NAPC09.AKruse&id=EcoNatRes. NAPCO9&isize=M. Madden, E.M., A.J. Hansen, and R.K. Murphy. 1999. Influence of prescribed fire history on habitat and abun- dance of passerine birds in northern mixed-grass prai- rie. Canadian Field-Naturalist 113: 627-640. Accessed 13 May 2019. https://biodiversitylibrary.org/page/3423 5497. Madden, E.M., R.K. Murphy, A.J. Hansen, and L. Mur- ray. 2000. Models for guiding management of prairie bird habitat in northwestern North Dakota. American Mid- land Naturalist 144: 377-393. https://doi.org/10.1674/00 03-0031(2000)144[0377:mfgmop]2.0.co;2 Moyles, D.L.J. 1981. Seasonal and daily use of plant com- 262 munities by Sharp-tailed Grouse (Pedioecetes phas- ianellus) in the parklands of Alberta. Canadian Field- Naturalist 95: 287-291. Accessed 13 May 2019. https:// www. biodiversitylibrary.org/page/28061990. Mundry, R., and J. Fischer. 1998. Use of statistical pro- grams for nonparametric tests of small samples often leads to incorrect values: examples from animal be- havior. Animal Behavior 6: 256—259. https://doi.org/10. 1006/anbe.1998 0756 Murphy, R.K. 1993. History, nesting biology, and pred- ation ecology of breeding raptors in the Missouri Coteau of northwestern North Dakota. Ph.D. thesis, Montana State University, Bozeman, Montana, USA. Accessed 13 May 2019. https://scholarworks.montana. edu/xmlui/bitstream/handle/1/7252/31762102084736. pdf;sequence=1. Murphy, R.K., and T.A. Grant. 2005. Land manage- ment history and floristics in mixed-grass prairie, North Dakota, USA. Natural Areas Journal 25: 351-358. Murphy, R.K., and K.A. Smith. 2007. Long-term trends in abundance of breeding songbirds during restora- tion of northern mixed-grass prairie. Transactions of THE CANADIAN FIELD-NATURALIST Vol. 133 the North American Prairie Conference 20: 359-370. Accessed 13 May 2019. https://digitalcommons.unl.edu/ usfwspubs/79/. Romo, J.T., P.L. Grilz, and E.A. Driver. 1990. Invasion of the Canadian prairies by an exotic perennial. Blue Jay 48: 131-135. Smith, K.A., and R.K. Murphy. 2007. Burning and graz- ing regime decimates Quaking Aspen in a northern mixed-grass prairie (North Dakota). Ecological Resto- ration 25: 63-64. U.S. Fish and Wildlife Service. 1998. Lostwood National Wildlife Refuge Comprehensive Conservation Plan. U.S. Fish and Wildlife Service, Mountain-Prairie Region, Denver, Colorado, USA. Accessed 13 May 2019. https:// www.fws.gov/mountain-prairie/refuges/completed PlanPDFs_F-L/Ist_1998_ccpfinal_all.pdf. U.S. Soil Conservation Service. 1975. Field technical guide. U.S. Department of Agriculture, Soil Conservation Ser- vice, Bismarck, North Dakota, USA. Received 27 August 2018 Accepted 8 December 2019 The Canadian Field-Naturalist Humpback Whale (Megaptera novaeangliae) observations in Laskeek Bay, western Hecate Strait, in spring and early summer, 1990-2018 ANTHOony J. GASTON!*, NEIL G. PILGRIM!, and VIVIAN PATTISON? 'Laskeek Bay Conservation Society, P.O. Box 867, Queen Charlotte City, British Columbia VOT 1S0 Canada ~Department of Geography, University of Victoria, 3800 Finnerty Road, Victoria, British Columbia V8P 5C2 Canada Corresponding author: tonygastonconsult@gmail.com Gaston, A.J., N.G. Pilgrim, and V. Pattison. 2019. Humpback Whale (Megaptera novaeangliae) observations in Laskeek Bay, western Hecate Strait, in spring and early summer, 1990-2018. Canadian Field-Naturalist 133(3): 263-269. https://doi.org/10.22621/cfn.v13313.2231 Abstract We describe observations of Humpback Whales (Megaptera novaeangliae) made along the west side of central Hecate Strait, British Columbia, during the spring and summer of 1990-2018. From none in March, the frequency of sightings increased from early April to a peak in May, then fell in June with few in July. The frequency of sightings during the peak period (1 May—20 June) increased over the course of the study at a mean rate of 6% a year, similar to increases recorded elsewhere in British Columbian waters. The frequency of sightings was highest in years when the Oceanic Nifio Index for January—March was low and peaked earlier in years when the Oceanic Nifio Index was high. Both of these relationships suggest a connection between Humpback Whale sightings in western Hecate Strait and the larger oceanographic context, with sightings more frequent in years of lower water temperatures. Key words: Humpback Whale; Megaptera novaeangliae, Hecate Strait; seasonal occurrence; population trends; oceanog- raphy Introduction Humpback Whale (Megaptera novaeangliae) is the most common rorqual along the west coast of Canada during the spring and summer, occurring in northern British Columbia (BC) waters princi- pally from May to September (COSEWIC 2011; Ford 2014). Most of the population that occurs in summer in northern BC waters winters around the Hawaiian Islands (Calambokidis et a/. 2001). Whales sighted in spring in BC waters may remain for the whole sum- mer or may pass through en route to summering grounds farther north (Ashe ef a/. 2013; Ford 2014). Most Humpback Whales are believed to be faithful to their summering areas, with the same individuals identified in particular parts of the summer range over several years (Rambeau 2008; Gabriele et al. 2017). Humpback Whale populations were heavily 1m- pacted by commercial whaling that took place along the BC coast between 1905 and 1967 (Trites et al. 2007). Since then, detailed observations between 1985 and 2014 in Glacier Bay, Alaska, showed that a humpback population summering there increased over that period at a mean 5% annually. A similar esti- mate, but based on fewer years, has been obtained for the population summering in inlets along the main- land coast of Hecate Strait (Ashe et a/. 2013), while an assessment of trends in BC waters by COSEWIC (2011) suggested an annual rate of increase in adult numbers of 4%. These trends reflect a population re- covery after severe reductions by commercial whal- ing in the period before 1970 (COSEWIC 2011). Since 1990, the Laskeek Bay Conservation So- ciety, a citizen science non-governmental organization based on the archipelago of Haida Gwaii, BC, has con- ducted observations of marine mammals in western Hecate Strait, in one of the three important Humpback Whale areas in BC waters identified by Dalla Rosa et al. (2012). Observations were made from a seasonal camp on East Limestone Island, a 40-ha island off the southeast corner of the much larger Louise Island, on the east coast of Haida Gwaii (Figure 1). In this paper, we summarize observations of Humpback Whales made over the period 1990-2018 from March to July. We analyze seasonal and inter- annual variation and compare our observations with those made elsewhere in the northeast Pacific. Given the large amount of inter-annual variation in our data, we compare them with variations in oceanographic A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 263 ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 264 THE CANADIAN FIELD-NATURALIST Vol. 133 A Boat-survey _ _ Area Louise Island =. c _ British 2. Columbia i waii NY Hecate q Wee. ~ ‘ a Wn ~~. Strait e ae Ral ‘ , pe C 0 270km “aa Bay a ‘ \ \ Laskeek \ Bay. z f a ie / ae / os SS / . { Cabin : / "3 _ Lookout Point ¥ wy 4 Lookout Survey Area Je & yy Point £ f ho » 7 Se oo r, aad 0 5 10km hay 0 0.5 ikm 4 1 J J FiGurE 1. Map of the study area showing: a. the arc of view from Lookout Point (dashed line) and the area within which boat surveys were conducted regularly (rectangle); b. the location of Haida Gwaii; c. detail of East Limestone Island. conditions, both in the northeast Pacific and more locally in BC waters, to improve our understanding of the factors influencing Humpback Whale occur- rence in western Hecate Strait. Methods Fieldwork The Laskeek Bay Conservation Society camp on East Limestone Island (Figure la—c) has been active in spring and early summer since 1990. Marine mam- mals were noted both systematically and incident- ally throughout the period when camp was occu- pied, for periods between 56 and 126 days (mean 88 days/year). Starting dates varied from 15 March to 5 May. In 1990, the first year of operations, camp was open 25 April—5 June, but thereafter, in all years up to 2004, camp opened before 10 April and closed be- tween 3 and 25 July. From 2005 to 2018, camp opened later, with starting dates between 21 April and 5 May and closure between 8 and 22 July (Table 1). Systematic timed observations of marine mam- mals were made for several hours each week from a point ~20 m above sea level (asl; maximum tidal range 4 m; Fisheries and Oceans Canada 2019) at the south- eastern tip of the island. If two or three observers were present, they watched together for 30 or 60 min, continuously, scanning waters within sight (an area of ~120 km? shown approximately in Figure la). When marine mammals were sighted, they were observed through a 25x60 spotting scope. Single observers scanned the area by dividing it into three sectors and spent 10 min on each sector in rotation. Watches were conducted during good visibility (usually >15 km), with sea conditions reflecting a Beaufort sea state of 3 or less (defined by World Meteorological Office as waves <1.25 m; National Oceans and Atmosphere Administration 2002). Incidental observations were made from several locations. The camp is located on the east shore of the island (Figure 1c), from which an arc of ~120° is visible in an east-northeast direction. Most observa- tions from camp were made from the cabin, ~5 m asl. People were present in camp for several hours each day. Incidental observations were made from other parts of the island shores and additional observations were also made from a small boat, used to survey for marine birds for 6—8 h every 10—15 days (area of rect- angle in Figure la), as well as while travelling be- tween islands for other fieldwork. Analysis To investigate seasonal variation in whale num- 2019 TABLE 1. Period during which the East Limestone Island camp was active in each year of the study. Year Start date End date Days of observation 1990 25 April 19 June 56 1991 26 March 14 June 81 1992 9 April 3 July 86 1993 9 April 10 July 98 1994 5 April 15 July 102 1995 25 March 15 July 113 1996 20 March 11 July 114 1997 15 March 11 July 119 1998 3 April 9 July 98 1999 2 April 25 July 115 2000 1 April 20 July 111 2001 22 March 25 July 126 2002 20 March 7 July 102 2003 20 March 4 July 99 2004 30 April 22 July 84 2005 22 April 22 July 92 2006 28 April 20 July 84 2007 28 April 13 July FF. 2008 5 May 16 July 73 2009 1 May 14 July PD 2010 1 May 9 July 70 2011 29 April 9 July 71 2012 4 May 12 July 70 2013 3 May 12 July 71 2014 1 May 11 July 72 2015 1 May 10 July 71 2016 30 April 22 July 84 2017 4 May 22 July 80 2018 4 May 20 July 78 bers, we used three statistics: (a) the proportion of observation days on which whales were seen, (b) the monthly sums of the number of whales seen each day (whales x days), and (c) the average number of whales seen on days when at least one was recorded. We in- cluded all years in this analysis, although no observa- tions were made in July 1990 and 1991 and, after 2003, no observations were made before 21 April. Because of variation in observing dates each year, only rec- ords from the 50-day period 1 May—20 June were used for inter-year trend analysis. Observations were made daily in every year during this period. We used the proportion of days on which one or more whales were seen during this 50-day period as our index of whale frequency (whale index, WI) for time-trend analysis. To examine the possible influence of large-scale oceanographic variation on the occurrence of Hump- back Whales in Laskeek Bay, we corrected the number of whales observed assuming an increas- ing population trend of 4% annually, as suggested by COSEWIC (2011). The resulting adjusted index of whale abundance is referred to as the “corrected whale index” (CWI): GASTON ET AL.. HUMPBACK WHALES IN LASKEEK BAY 265 CWI =(D,/D,) x 1.040°8 where D,, = days on which whales were sighted in a given year; D,= total days camp was occupied during 1 May—20 June; and y = year of observations. This index was compared with the following ocean climate indices: As a measure of the El Nifio/Southern Oscillation (ENSO), the Oceanic Nifio Index (ONI) for January— March, the three-month running mean of ERSST. v5 (extended reconstructed sea surface temperature anomalies in the Nifio 3.4 region; 5°N—5°S, 120-— 170°W), based on centred 30-year base periods up- dated every five years (National Weather Service n.d.). Sea surface temperatures in the northeast Pacific tend to be closely correlated with indices of the ENSO cycle (e.g., Nifio 3.4 index; Tseng et al. 2017). As a measure of the Pacific Decadal Oscillation (PDO), the H300-based PDO index (HPDO), defined as the projections of monthly mean H300 anom- alies from the National Centers for Environmental Prediction’s Global Ocean Data Assimilation System onto their first empirical orthogonal function vector in the North Pacific (20°—60°N), based on the 30-year period from 1981 to 2010 (GODAS n.d.). Years in which the ONI was —0.5 or lower for the first three months of the year were classified as “cold” (as defined at National Weather Service n.d.) and the CWI for these years was compared with the CWI for warmer years. Comparisons among days with and without whale sightings were made using the Fisher exact probability test. Tests for time trends were made using linear regression and the Pearson correlation coefficient. Statistics were performed using Statistica v. 7.1 (Statsoft, Inc., Tulsa, Oklahoma, USA). Mean values are given + | SE. Results Humpback Whales were seen in all but three years of the study, with sightings from early April to late July. They were recorded on 14% of the 2572 days that camp was occupied and on 20% of the 1673 days during the period 1 May—20 June. No humpbacks were seen in March and the frequency of sightings built up during April, with the buildup continuing longer in cold than in warmer years (Figure 2). The highest sighting frequency occurred in May, peaking 21-31 May in cold years (when whales were seen on 36% of days) and 1-10 May in other years (recorded on 21% of days). WIs were significantly higher in cold years than in others during 21-31 May and 1-10 June (Fisher exact test, P < 0.001 for both periods). No humpbacks were seen in 1990, 1991, or 1996. The highest frequencies for 1 May—20 June oc- curred in 2007 (WI = 56% of days), 2008 (63%), 266 0.40 o uw a o N ui a days with whales ~) ‘ in 11-20 21-31 1-10 11-20 21-30 1-10 11-20 April May March THE CANADIAN FIELD-NATURALIST o w oO Vol. 133 * @Cold Other °o No So 21-31 1-10 11-20 21-30 1-10 11-20 21-31 June July FiGuRE 2. Proportion of days when Humpback Whales (Megaptera novaeangliae) were seen in Laskeek Bay in relation to date, for years when the Oceanic Nifio Index was less than —0.5 during January—March (cold) and other years (1990-2018). *Proportion of days with whales was significantly greater in cold years than in other years (Fisher exact P < 0.001). 2016 (39%), and 2018 (41%). Six of the 10 lowest years occurred before 1999 (Figure 3). There was a significant positive correlation between year and the proportion of days with whales during 1 May— 20 June (r,, = 0.48, P = 0.009). A similar positive correlation was found for non-cold years (r,, = 0.48, P = 0.04) when analyzed separately. The correla- tion coefficient was similar, but non-significant for cold years (rz = 0.41, P = 0.24). The linear regres- sion slope for the proportion of days with whales over time was consistent with an annual rate of in- crease of 6%. Slopes were similar for cold and non- cold years when analyzed separately, but were closer 0.7 0.6 to a 4% rate of annual increase (Figure 4). Number of whales per day Summing daily counts, 1750 humpback sight- ings were recorded, 1602 during the period 1 May—20 June. Probably many of these involved the same ani- mals on different days, but we think it unlikely that many involved the same animal seen more than once on a given day. The highest number was recorded during May (1304, 75% of all sightings). Highest numbers of whales x days were recorded in 2003 (142), 2007 (213), and 2014 (233). The number sighted on days when at least one whale was seen aver- y = 0.0089x + 0.0479 R? =0.1975 0.5 0.4 0.3 Whale index (WI) 0.2 0.1 =e 0.0 1990 1993 1996 1999 2002 2005 2008 2011 2014 2017 FiGurE 3. Whale index (WI), 1.e., days when Humpback Whales (Megaptera novaeangliae) were seen in Laskeek Bay (11 May-—20 June) as a proportion of all days, during 1990-2018, showing linear regression. 2019 0.7 @ Cold years oOther 0.6 = 05 wn a qe ca = e re "S03 , SS e O So 0.2 3X e —_ 0.1 : = == _-02-7"7 0 ie ~~ 9 o ° 1990 1993 1996 1999 2002 Ficure 4. Proportion of days with whales (WI) in Laskeek GASTON ET AL.. HUMPBACK WHALES IN LASKEEK BAY 267 R? = 0.1397 -_ R? = 0.2004 _ —_ 0 — ° ° 2005 2008 2011 2014 2017 Bay (11 May—20 June) separated into years with Oceanic Nifio Index below —0.5 during January—March (cold years) and warmer years (other). aged 5.6 whales/day during 1 May—20 June and 2.3 whales/day outside that period (Figure 5). The num- ber of whales seen per day on days when at least one whale was seen did not differ significantly between cold years (4.5 + 1.8 whales/day) and other years (4.3 + 0.9 whales/day, 4, = 0.11, P =0.9). Effects of oceanography The proportion of days with whales was generally higher in years with negative ONI (cold years, 28% + 5%) than others (15% + 4%, t,, = 2.02, P=0.05). CWI was negatively correlated with ONI for January— March (r,, = 0.37, P = 0.037; Figure 6), but did not show any relation to the HPDO index (P > 0.10). The ONI accounted for 17% of variation in CWI (Fy, = 4.82, adjusted R? = 0.17, beta = —0.41). 9 o > ui a ~ oo | Whales per whale-day Ww cael ellie N fay 11-20 21-31 1-10 11-20 21-30 1-10 11-2 March April Discussion Despite substantial variation in the amount of ef- fort devoted to whale observations and the inevit- able fluctuations in viewing conditions created by weather, our results show a clear increasing trend in the frequency of Humpback Whale sightings in Laskeek Bay since 1990. The complete set of annual indices has a regression coefficient consistent with a 6% annual rate of increase, while dividing the years into those displaying colder relative oceanic condi- tions and others (average or warmer conditions), based on the ONI, suggests a rate of increase closer to 4% for both samples. Observations of marine mam- mals from nearby Reef Island (5 km ESE of East Limestone Island) during April-June of 1984-1989 @cold OOther 21-31 1-10 11-20 21-30 1-10 11-20 21-31 June July FiGurE 5. Number of whales seen per day in Laskeek Bay (11 May—20 June) on days when at least one was recorded (1990-2018). 268 THE CANADIAN FIELD-NATURALIST Vol. 133 1.0 0.9 0.8 0.6 0.5 Corrected whale index (CWI) =6 -4 -2 y = -0.0396x + 0.3141 R2 = 0.1675 Oceanic Nifio Index (ONI) for January-March Ficure 6. The corrected whale index (assumes a 4% annual rate of population increase) compared with El Nifio/Southern Oscillation conditions, represented by the Oceanic Nifio Index (ONI). Negative ONI is associated with colder than aver- age ocean conditions, while positive ONI is associated with warmer than average conditions (Laskeek Bay, 1990-2018). included sightings of Humpback Whales only in 1985 (up to three on 17 days), 1987 (one on a single day), and 1989 (up to five on six days; Gaston and Jones 1991). In all years, these observations extended from early April to mid-June, but all sightings fell between 2 May and 6 June (Gaston and Jones 1991). The pau- city of sightings during the 1980s supports the idea that numbers have increased substantially since then. Our results are consistent with those obtained else- where in BC waters (COSEWIC 2011; Ashe et al. 2013). An estimate by Fisheries and Oceans Canada (2009) suggested a mean rate of increase for the BC population of 4.1% a year, identical with ours once warm and cold years are separated. The appearance of large numbers of Humpback Whales in Queen Charlotte Strait and the inside passage off Vancouver Island since the early 2000s (Nichol et al. 2017) is also consistent with our findings. The absence of humpbacks in March and low numbers in the first 20 days of April may be partly accounted for by lower population size in the early years of the study, when most observations in March and April occurred. However, the number of days of observations after 10 July was biased toward recent years; thus, the decrease in number of sightings after mid-June is unlikely to have been influenced by the population trend. Some of the humpbacks recorded in Laskeek Bay may be migrating to summering areas farther north. The timing of peak numbers reported in Laskeek Bay fits well with data from Glacier Bay, Alaska, ~750 km by sea to the north of Laskeek Bay, where the peak arrival of humpbacks occurs in June (Gabriele et al. 2017), about three weeks after the peak in Laskeek Bay. This rate of travel (about 36 km/day) is comfortably within the migration speed of 48 km/ day observed for humpbacks by satellite telemetry (Lagerquist et al. 2008). However, it is possible that some or all of the whales seen in Laskeek Bay shift to other BC waters in July. Animals were frequently observed feeding in Laskeek Bay, both lunging at the surface and “flick feeding” (A.J.G. unpubl. data), which Ford (2014) mentions as common in waters off Moresby Island. It seems likely that most whales ob- served were feeding in the vicinity, causing them to pause in the area for a period. Inter-year variability in sighting frequency was high, with the proportion of days with humpback sightings during the period 1 May—20 June, varying from 0 to 60%. Part of this variation can be explained by oceanographic processes, with the Oceanic Nifio Index accounting for 17% of variation in the trend- corrected proportion of whale sightings. Seasonal trends in sightings, with sightings in cold years peak- ing later than those in other years, suggests that ocean conditions, influenced by large-scale processes, such as ENSO, may affect the suitability of inshore wat- ers along the western side of Hecate Strait for hump- back foraging. A similar effect of large-scale oceano- graphic forcing on Humpback Whales (in that case on diet) was reported by Fleming ef al. (2016). The fact that numbers seen on a given day were not affected by ONI suggests that much of the variation in obser- vation frequency probably relates to the rate at which the whales pass through the area, rather than being accounted for by fluctuations in the number of indi- 2019 viduals using the area. Given the much greater fre- quency of whale sightings in Laskeek Bay in recent years, we may be able to make more detailed obser- vations in future, perhaps with greater emphasis on photo-identification, giving us better understanding of the importance of Laskeek Bay waters to individ- ual Humpback Whales. Acknowledgements We thank past staff and volunteers of the Laskeek Bay Conservation Society for their contributions to the observations we catalogue here. Useful comments on the manuscript were provided by Christie McMil- lan and Jackie Hildering of the Marine Education and Research Society and by an anonymous reviewer. For permission to maintain the field camp at East Limestone Island, we thank British Columbia Parks and the Council of the Haida Nation. Literature Cited Ashe, E., J. Wray, C.R. Picard, and R. Williams. 2013. Abundance and survival of Pacific Humpback Whales in a proposed critical habitat area. PLoS ONE 8: e75228. https://doi.org/10.1371/journal.pone.0075228 Calambokidis, J., G.H. Steiger, J.M. Straley, L.M. Herman, S. Cerchio, D.R. Salden, J. Urban-Ramirez, J.K. Jacobsen, O. Von Ziegesar, K.C. Balcomb, C.M. Gabriele, M.E. Dahlheim, S. Uchida, G. Ellis, Y. Miyamura, P. Ladron de Guevara, M. Yamaguchi, F. Sato, S.A. Mizroch, L. Schlender, K. Rasmussen, J. Barlow, and T.J. Quinn. 2001. Movements and population structure of Humpback Whales in the North Pacific. Marine Mammal Science 17: 769-794. https:// dot.org/10.1111/j.1748-7692.2001.tb01298.x COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2011. COSEWIC assessment and status report on the Humpback Whale Megaptera novae- angliae in Canada. COSEWIC, Ottawa, Ontario, Canada. Dalla Rosa, L., J.K.B. Ford, and A.W. Trites. 2012. Distribution and relative abundance of humpback whales in relation to environmental variables in coastal Bri- tish Columbia and adjacent waters. Continental Shelf Research 36: 89-104. https://doi.org/10.1016/j.csr.2012. 01.017 Fisheries and Oceans Canada. 2009. Recovery poten- tial assessment of the Humpback Whale, Pacific Popu- lation. Canadian Science Advisory Secretariat Science Advisory Report 2009/048. Fisheries and Ocean Can- ada, Ottawa, Ontario, Canada. Fisheries and Oceans Canada. 2019. Copper Islands (#9724): 2019 tide tables. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Accessed 1 January 2019. https://www.tides.gc.ca/eng/data/table/2019/wlev_sec /9724. Fleming, A.H., C.T. Clark, J. Calambokidis, and J. Barlow. 2016. Humpback whale diets respond to vari- ance in ocean climate and ecosystem conditions in the California Current. Global Change Biology 22: 1214— 1224. https://doi.org/10.1111/gcb.13171 GASTON ET AL.. HUMPBACK WHALES IN LASKEEK BAY 269 Ford, J.K.B. 2014. Marine Mammals of British Columbia. Royal BC Museum, Victoria, British Columbia, Canada. Gabriele, C.M., J.L. Neilson, J.M. Straley, C.S. Baker, J.A. Cedarleaf, and J.F. Saracco. 2017. Natural his- tory, population dynamics, and habitat use of hump- back whales over 30 years on an Alaska feeding ground. Ecosphere 8: e01641. https://doi.org/10.1002/ecs2.1641 Gaston, A.J., and I.L. Jones. 1991. Seabirds and marine mammals recorded in western Hecate Strait, British Co- lumbia, in spring and early summer, 1984-1989. Can- adian Field-Naturalist 105: 550-560. Accessed 16 January 2020. https://biodiversitylibrary.org/page/34349068. GODAS (Global Ocean Data Assimilation System). n.d. Monthly HC300-based PDO. Climate Prediction Center, National Weather Service, College Park, Mary- land, USA. Accessed 1 January 2019. http://www.cpc. ncep.noaa.gov/products/GODAS/PDO body. html. Lagerquist, B.A., B.R. Mate, J.G. Ortega-Ortiz, M. Winsor, and J. Urban-Ramirez. 2008. Migratory movements and surfacing rates of humpback whales (Megaptera novaeangliae) satellite tagged at Socorro Island, Mexico. Marine Mammal Science 24: 815-830. https://doi.org/10.1111/).1748-7692.2008.00217.x National Oceans and Atmospheric Administration. 2002. WOCE Upper Ocean Data. Accessed 8 January 2020. https://www.nodc.noaa.gov/woce/woce_v3/wocedata_1/ woce-uot/document/wmocode.htm. National Weather Service. n.d. Cold & warm episodes by season. Climate Prediction Center, National Weather Service, College Park, Maryland, USA. Accessed 1 January 2019. http://origin.cpc.ncep.noaa.gov/products/ analysis_monitoring/ensostuff/ONI_vS. php. Nichol, L.M., S.P. Majewski, B.M. Wright, C.J. McMil- lan, J. Hildering, E.U. Rechsteiner, and J.K.B. Ford. 2017. Humpback whales, harbor seals, and Steller sea lions in British Columbia: population trends of formerly harvested marine mammals. Pages 115-121 in State of the physical, biological and selected fishery resources of Pacific Canadian marine ecosystems in 2016. Canadian Technical Report of Fisheries and Aquatic Sciences 3225. Edited by P.C. Chandler, S.A. King, and J. Boldt. Fisheries and Oceans Canada, Sidney, British Columbia, Canada. Rambeau, A.L. 2008. Determining abundance and stock structure for a widespread migratory animal: the case of humpback whales (Megaptera novaeangliae) in British Columbia, Canada. M.Sc. thesis, University of British Columbia, Vancouver, British Columbia, Canada. Trites, A.W., V.B. Deecke, E.J. Gregr, J.K.B. Ford, and P.F. Olesiuk. 2007. Killer whales, whaling, and sequen- tial megafaunal collapse in the North Pacific: a compar- ative analysis of the dynamics of marine mammals tn Ala- ska and British Columbia following commercial whal- ing. Marine Mammal Science 23: 751—765S. https://doi.org/ 10.1111/j.1748-7692.2006.00076.x Tseng, Y.H., R. Ding, and X. Huang. 2017. The warm Blob in the northeast Pacific—the bridge leading to the 2015/16 El Nifio. Environmental Research Letters 12: 054019. https://doi.org/10.1088/1748-9326/aa67c3 Received 16 March 2019 Accepted 23 December 2019 The Canadian Field-Naturalist A Tribute to Rudolph Frank Stocek, 1937-2018 DONALD F. MCALPINE!" and GRAHAM J. FORBES? 'New Brunswick Museum, 227 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada *University of New Brunswick, Fredericton, New Brunswick E3B 5A3 Canada Corresponding author: email: donald.mcalpine@nbm-mnb.ca McAlpine, D.F., and G.J. Forbes. 2019. A Tribute to Rudolph Frank Stocek, 1937—2018. Canadian Field-Naturalist 133(3): 270-275. https://doi.org/10.22621/cfn.v13313.2435 Rudolph (Rudy) Frank Stocek, aged 81, “the eagle man” of New Brunswick (Figure 1), passed away on 2 December 2018 at the Dr. Everett Chalmers Hospital, Fredericton, New Brunswick following a stroke. Rudy was born 5 June 1937 in Woodside, New York, just south of the Bronx, even then one of the most densely populated regions of the United States. However, summers at his grandparent’s farm in Millhurst, New Jersey, left Rudy with a deep love of the outdoors and a fascination with the natural history of the trees, fish, birds, and other wildlife that populated the surround- ing waterways and woodlands. The north woods beckoned, and Rudy set off to the University of Maine at Orono, graduating in 1959 with a B.Sc. in Wildlife Management and a minor in Forestry. During his undergraduate summers he worked for the Maine Department of Inland Fisheries and Game, undertaking stream and lake surveys, tag- ging fish, and reading fish scales. Athletic and active, Rudy played football and boxed in university. His athleticism would serve him well as both a field biolo- gist and an instructor in wildlife management. With a solid, practical, introduction to fisheries management during his summers, it probably seemed natural to pursue fisheries science at the Ontario Agricultural College in Guelph, Ontario, then affili- ated with the University of Toronto. Rudy com- pleted his M.Sc. in 1962 under the supervision of Dr. Hugh R. McCrimmon, a committed fish cul- turist, now remembered for his volume on carp in Canada (McCrimmon 1968). Although Rudy later became best known for his work on raptors, his in- terest in freshwater fish never left him. He designed and taught the first ichthyology course offered at the University of New Brunswick, and he reported the addition of Muskellunge (Esox masquinogy) to the Saint John River system. Even as he approached 70, Figure 1. Rudy Frank Stocek, 1937-2018. Photograph taken as part of a composite for the 1997 Maritime Forest Ranger School graduating class. Photo: J. Cummings. he documented the occurrence of Fat Head Minnow (Pimephales promelas) in New Brunswick. Brief stints as a wildlife biologist with the Metro- politan Toronto and Region Conservation Authority (1960-1961), and as Manager of the Tinicum Wildlife Preserve, Philadelphia (1961-1965), followed his M.Sc. In 1965 Rudy entered a Ph.D. program at the University of Western Ontario to study growth and development in young Canada Geese (Branta ca- nadensis). However, his marriage in 1960 (to Arlene, nee Wellhauser), and the two children that followed (Rudy Jr. born 1963, Lehanne born 1964), made a 1968 job opportunity in New Brunswick simply A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 270 ©The Authors. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 too good to pass up. Rudy and family moved east to the Maritimes, where Rudy took a position as an Instructor and Fish and Wildlife Biologist with the Maritime Forest Ranger School (MFRS; since 2003 the Maritime College of Forest Technology). Opened in 1946 as a co-operative venture of the New Brunswick—Nova Scotia governments and the forest industry, at its establishment the role of the MFRS was to re-train and integrate returning World War II veterans. However, by the time Rudy arrived in New Brunswick, MFRS was catering to students from wide and varied backgrounds. By 1973, Rudy’s ability to inspire students in- terested in pursuing careers that required a knowl- edge of fish and wildlife biology had become evi- dent. From then until his retirement in 2002, Rudy designed, directed, and with other instructors, deliv- ered, the MFRS wildlife technology program as his “day job” to hundreds of young men and women in- tent on becoming forest rangers, wildlife protection Officers, fish and wildlife technicians, park rangers, and wardens. But for one with Rudy’s energy and cu- riosity, teaching alone was not enough. As an inde- pendent fish and wildlife biologist, he also accepted contracts from various federal and provincial gov- ernment departments and private environmental con- sulting firms to investigate a variety of wildlife-re- lated issues in the Maritimes. A 1982 contract from the New Brunswick Environmental Assessment Branch that identified and assessed more than 90 environmentally sensitive areas in southern New Brunswick was an important precursor to current work by government and land trusts now intent on setting aside habitat in New Brunswick for conserva- tion purposes. In the broader context, and in a prov- ince dominated by industrial forestry, Rudy’s re- search and teaching can be seen as part of a rising tide of concern for North American wildlife and the environment that became evident in the 1970s. Until 1968, New Brunswick was one of the heaviest DDT (Dichlorodiphenyltrichloroethane) users in North America, a pesticide identified as a major cause of raptor declines across the continent and one that ap- pears to have had impacts on aquatic ecosystems in the province that are still evident (Kurek ef a/. 2019). With the establishment of the New Brunswick Endangered Species Act in 1976 (superseded by the New Brunswick Species at Risk Act in 2012) a short list of species was accorded protection. Although Bald Eagle (Haliaeetus leucocephalus) was assigned regionally endangered status, there were limited data to work from. There are two sources of Bald Eagles occupying New Brunswick. A resident population breeds in the province (Haliaeetus leucocephalus washingtonien- McALPINE AND ForRBES: A TRIBUTE TO R.F. STOCEK 27) sis), while immature birds fledged in the southeast- ern United States (Haliaeetus leucocephalus leuco- cephalus) disperse northward into New Brunswick in the late summer. In the early 1970s the impact of pes- ticide use in the United States was evident in New Brunswick, with a reduced number of southern birds available to disperse northward. Although legend- ary New Brunswick wildlife biologist Bruce Wright (1912-1975) had identified the Saint John River estu- ary as critical summer habitat for Bald Eagles fledged south of the province (Wright 1953), historically the species was never a common breeding bird in New Brunswick. A mere 12-15 pairs nested in the prov- ince by the early 1970s. Accordingly, Rudy was contracted by the New Brunswick Department of Natural Resources to undertake regular Bald Eagle surveys in New Brunswick. This was not Rudy’s first foray into rap- tor research. In 1973 Rudy had been engaged by the Canadian Wildlife Service to assess the status and re- productive success of Peregrine Falcon (Falco pere- grinus) and Osprey (Pandion haliaetus), along with Bald Eagle, across the Maritimes. Rudy would con- tinue annual New Brunswick Bald Eagle surveys for the next 25 years. When funds for aerial surveys be- came difficult to secure, undeterred, Rudy was able to take advantage of members of the Fredericton Flying Club and the 422nd Tactical Helicopter Squadron out of Canadian Forces Base Gagetown. In addition to determining numbers of breeding pairs in New Brunswick, Rudy also collected information on dis- tribution, nesting success, habitat requirements, and winter feeding patterns and developed management guidelines for individual Bald Eagle nesting sites (Figure 2). With a growing interest in the problems of rap- tor conservation in North America, Rudy soon found himself presenting his findings, both for Bald Eagles and Ospreys, at meetings of raptor specialists across North America. As an experienced educator and an engaging speaker, Rudy also lectured widely on Bald Eagle in the Maritimes to regional audiences. For the media, he became the go-to-guy for expert opinion on a bird with a high public profile, and huge signif- icance to the Indigenous community. Rudy went on to write the Bald Eagle account for the iconic federal government Hinterland Who’s Who series (Stocek 1992) and in 2006 summarized his decades of re- search on the bird in a book that ultimately received an independent publishers book award. While best known for his work on raptors, Bald Eagles in particular, Rudy also found time to work and publish on other Maritime wildlife, including Common Loon (Gavia immer), the elusive Cougar (Felis concolor), and Tree Swallows (Tachycineta bi- 2A2 FiGure 2. Rudy banding a Bald Eagle (Haliaeetus leuco- cephalus) nestling in southwestern New Brunswick, circa 1978. Photo: unknown. color). A useful and well-received field guide to New Brunswick trees and shrubs in winter helped ful- fil his commitment to teaching, but was also made a te PTA THE CANADIAN FIELD-NATURALIST Vol. 133 available to a wider general audience (Figure 3). A mainstay of provincial government committees deal- ing with species of conservation concern, in 1995 Rudy received the Career Achievement Award of the Atlantic Society of Fish and Wildlife Biologists and in 2010 an Award of Professional Excellence from the University of Maine Wildlife Program. Outgoing and gregarious, Rudy somehow still found time to play tuba and accordion in five local bands, stay active as a judge and organizer for the New Brunswick and Canadian gymnastics communities (including judg- ing at the 1976 Olympics in Montreal), and help man- age the local curling club! Today, as the biodiversity crisis deepens, there is a growing chorus calling for the revitalization of natural history (Schmidly 2005; Nature News 2014; Tewksbury et al. 2014). In the best possible way, Rudy was an “old school” wildlife biologist who never left natural history behind, even as he upped his game after graduate school with courses in com- puter programming, teaching and administration, and media communications. A first-rate field natural- ist, Rudy could identify trees and shrubs, and knew his fish, his birds, and his mammals. But his second love, after Arlene, his wife of 58 years, was the Bald Eagle. Although recent research has documented pre- cipitous declines in numbers of birds of many spe- cies in North America (Rosenberg ef a/. 2019), Rudy had the satisfaction of watching New Brunswick’s Bald Eagle populations, both migratory and resi- dent, recover and rise dramatically, and know that his work played a role in this. By the time the 1986-1990 if 2 maul Ficure 3. Rudy with a Maritime Forest Ranger School dendrology class in 2000, two years before his retirement. His 1991 winter field guide to trees and shrubs continues to be used in wildlife and forestry programs. Photo: Maritime School of Forest Technology. 2019 Maritimes Breeding Bird Atlas had been completed, 40 pairs of Bald Eagles were confirmed nesting in New Brunswick (Erskine 1992), and numbers have continued to rise. An astounding 92 pairs of Bald Eagles nested in New Brunswick during the second atlas period (2006-2010; Stewart et al. 2015). Bald Eagle populations in New Brunswick are now recog- nized as secure. There have been few success stories for North American wildlife since Europeans arrived on the continent over 400 years ago, but the recovery of the Bald Eagle is one of them. That recovery is tes- tament to the vision and the hard work of many, Rudy among them. Acknowledgements We are grateful to Lehanne Knowlton and Arlene Stocek for providing access to Rudy’s papers and for help filling in details related to Rudy’s life and scien- tific contributions. Lehanne Knowlton, Peter Pearce, and Arlene Stocek kindly commented on an early draft of the manuscript. Literature Cited Erskin, A.J. 1992. Atlas of breeding birds of the Maritime Provinces. Nimbus Publishing Limited and the Nova Scotia Museum, Halifax, Nova Scotia, Canada. Kurek, J., P.\W. MackKeigan, S. Veinot, A. Mercer, and K.A. Kidd. 2019. Ecological legacy of DDT archived in lake sediments from eastern Canada. Environmental Science Technology 53: 7316-7325. https://doi.org/10. 1021 /acs.est.9b01396 McCrimmon, H.R. 1968. Carp in Canada. Fisheries Research Board of Canada, Ottawa. Bulletin 165. Nature News. 2014. Natural decline. Few biology degrees still feature natural history. Is the naturalist a species in decline? Nature 508: 7-8. https://do1.org/10.1038/508 007b Rosenberg, K.V., A.M. Dokte, P.J. Blancher, J.R. Sauer, A.C. Smith, P.A. Smith, J.C. Stanton, A. Panjabi, L. Helft, M. Parr, and P.P. Marra. 2019. Decline of North American avifauna. Science 366: 120-124. https://doi. org/10.1126/science.aaw1313 Schmidly, D.J. 2005. What it means to be a naturalist and the future of natural history at American universities. Journal of Mammalogy 89: 449—456. https://doi.org/10. 1644/1545-1542(2005)86[449:wimtba]2.0.co;2 Stewart, R.L.M., K.A. Bredin, A.R. Couturier, A.G. Horn, D. Lepage, S. Makepeace, P.D. Taylor, M.A. Villard, and R. Mm. Whittam. 2015. Second atlas of breeding birds of the Maritime Provinces. Bird studies Canada, Environment Canada, Natural History Society of Prince Edward Island, Nature New Brunswick, New Brunswick Department of Natural Resources, Nova Scotia Bird Society, Nova Scotia Department of Natural Resources, and Prince Edward Island Department of Agriculture and Forestry, Sackville, New Brunswick, Canada. Stocek, R.F. 1992. Bald Eagle. Hinterland Who’s Who. Canadian Wildlife Service, Environment Canada, Ot- tawa, Ontario, Canada. Accessed 19 January 2020. McALPINE AND ForRBES: A TRIBUTE TO R.F. STOCEK DIS http://www.hww.ca/en/wildlife/birds/bald-eagle.html. (also issued in French under the title: Le pygargue a téte blanche). Tewksbury, J.J., J.G.T. Anderson, J.D. Bakker, T.J. Billo, P.W. Dunwiddie, M.J. Groom, S.E. Hampton, S.G. Herman, D.J. Levey, N.J. Machnicki, C. Martinez del Rio, M.E. Power, K. Rowell, A.K. Solomon, L. Stacey, S.C. Trombulak, and T.A. Wheeler. 2014. Natural his- tory’s place in science and society. BioScience 64: 300— 310. https://doi.org/10.1093/biosci/b1u032 Wright, B.S. 1953. The relation of Bald Eagles to breeding ducks in New Brunswick. Journal of Wildlife Manage- ment 17: 55-62. Bibliography of Rudolph Frank Stocek Stocek, R.F. 1962. Some aspects of the ecology of the Rain- bow Trout (Salmo gairdneri) and the Largemouth Bass (Micropterus salmoides) in a small lake. M.Sc. thesis, University of Toronto (Ontario Agricultural College), Guelph, Ontario, Canada. Stocek, R.F. 1965. Tinicum—A City Preserve. Frontiers 29: 136-141. Stocek, R.F., and H.R. McCrimmon. 1965. The co-exist- ence of Rainbow Trout (Salmo gairdneri Richardson) and Largemouth Bass (Micropterus salmoides Lace- pede) in a small Ontario lake. Canadian Fish Culturist 35: 37-58. Stocek, R.F. 1970. Observations on the breeding biology of the Tree Swallow. Cassinia 52: 3-20. Stocek, R.F. 1972a. Copulation in the Tree Swallow. Eastern Bird Banding News 35: 163-165. Stocek, R.F. 1972b. The occurrence of Osprey on electric power lines in New Brunswick. N.B. Naturalist 3: 19-27. Stocek, R.F. 1972c. Technical in service training for fish and wildlife personnel. Canadian Society of Wildlife and Fish Biologists, Atlantic Chapter Newsletter 2: 3-5. Stocek, R. 1973. The Tree Farm: there’s a lot [of wild- life]. The Maritime Farmer and Co-operative Dairyman, Sussex, New Brunswick, Canada. 20 March, No 12: 2, 17. Stocek, R.F. 1976a. A dwarf Tree Swallow egg. Canadian Field-Naturalist 90: 183-184. Accessed 26 January 2020. https://biodiversitylibrary.org/page/28045666. Stocek, R.F. 1976b. The 1975 North American Peregrine Falcon Survey—The Maritime Provinces. Canadian Field-Naturalist 90: 248. Accessed 26 January 2020. https://biodiversitylibrary.org/page/28045735. Stocek, R.F., and P.A. Pearce. 1978a. The Peregrine Fal- con in the Maritime Provinces. Manuscript Report. No. 36. Canadian Wildlife Service, Wildlife Toxicology Division, Ottawa, Ontario, Canada. Stocek, R.F., and P.A. Pearce. 1978b. The Bald Eagle and Osprey in the Maritime Provinces. Manuscript Report. No. 37. Canadian Wildlife Service, Wildlife Toxicology Division, Ottawa, Ontario, Canada. Stocek, R.F. 1979. Decline of summering Bald Eagles in central New Brunswick. Canadian Field-Naturalist 93: 443-445, Accessed 26 January 2020. http://biodiversity library.org/page/28063770. Stocek, R.F. 1981. Bird Related Problems on Electric Power Systems in Canada. Report 110-T-210. Canadian 274 Electrical Association Research, Montreal, Quebec, Canada. Stocek, R.F., and P.A. Pearce. 1981. Status and breed- ing success of New Brunswick Bald Eagles. Canadian Field-Naturalist 35: 428-433. Accessed 26 January 2020. https://biodiversitylibrary.org/page/28062135. Stocek, R.F. 1982. Environmentally Significant Areas in the Saint John Planning Region of New Brunswick. Environmental Services Branch, Environment New Brunswick, Fredericton, New Brunswick, Canada. Patch, J.R., and R.F. Stocek. 1983. Introduction. Pages 1-2 in Wildlife Habitat Management and Small Woodlots. Proceedings of the 14 June 1983 DNR Workshop. Fish and Wildlife Branch, New Brunswick Department of Natural Resources, Fredericton, New Brunswick, Canada. Stocek, R. 1983a. The Bald Eagle. New Brunswick Natural Resources. Department of Natural Resources, Frederic- ton, New Brunswick, Canada. Spring 1983: 14. Stocek, R.F. 1983b. Non-Game Birds and Woodlot Man- agement. Pages 49-73 in Wildlife Habitat Management and Small Woodlots. Proceedings of the 14 June 1983 DNR Workshop. Fish and Wildlife Branch, New Bruns- wick Department of Natural Resources, Fredericton, New Brunswick, Canada. Stocek, R.F., and P.A. Pearce. 1983. Distribution and re- productive success of Ospreys in New Brunswick, 1974— 1980. Pages 215-221 in Biology and Management of Bald Eagles and Ospreys. Proceedings of the Ist Sym- posium on Bald Eagles and Ospreys. Edited by D.M. Bird, N.R. Seymour, and J.M. Gerrard. McDonald Raptor Research Centre and Raptor Research Foun- dation, McGill University, Montreal, Quebec, Canada. Stocek, R.F. 1984. Snags and woodpeckers in the Uni- versity of New Brunswick forest. Technical Bulletin No. 117. Environment Canada, Canadian Forestry Ser- vice, Martimes Forest Research Centre, Fredericton, New Brunswick, Canada. Stocek, R.F. 1985a. The Bald Eagle in New Brunswick in the 80’s. Pages 44-47 in The Bald Eagle in Canada: Proceedings of Bald Eagle Days, 1983. Edited by J.M. Gerrard and T.N. Ingram. White Horse Palins Publish- ers, Headingly, Manitoba, Canada. Stocek, R. 1985b. Bald Eagles and winter feeding. N.B. Naturalist 14: 138-140. Stocek, R.F., and D. Cartwright. 1985. Birds as non-tar- get catches in the New Brunswick furbearer harvest. Wildlife Society Bulletin 13: 314-317. Stocek, R.F. 1986a. Spring weather and the local movement of Tree Swallows. Canadian Field-Naturalist 100: 134— 136. Accessed 26 January 2020. https://biodiversity library.org/page/28072209. Stocek, R. 1986b. Notes on Tree Swallow breeding biology I—Aggression directed towards human disturbance. N.B. Naturalist 15: 87-89. Stocek, R. 1987. Notes on Tree Swallow breeding biology II —Ageing nestlings. N.B. Naturalist 16: 19-20. Stocek, R. 1988a. Notes on Tree Swallow breeding biology I1IJ—Nest building and breeding success. N.B. Natura- list 16: 50-51. THE CANADIAN FIELD-NATURALIST Vol. 133 Stocek, R. 1988b. The Bald Eagle in New Brunswick. N.B. Naturalist 16: 81-82. Stocek, R. 1989a. An unusual Mourning Dove breeding re- cord. N.B. Naturalist 16: 116. Stocek, R.F. 1989b. The Common Loon in New Brunswick. Status Report prepared for the World Wildlife Fund (Canada), Toronto, Ontario, Canada. Stocek, R. 1990. The rise and fall of Passer domesticus. N.B. Naturalist 17: 60. Stocek, R. 1991a. The abundance of wintering Bald Eagles in New Brunswick. N.B. Naturalist 18: 4—5. Stocek, R.F. 1991b. New Brunswick trees and shrubs in winter: a field guide. New Brunswick Maritime Forest Ranger School, and Canada/New Brunswick Cooperation Agreement on Forestry Development, Fre- dericton, New Brunswick, Canada. Stocek, R.F. 1992a. Bald Eagle. Hinterland Who’s Who. Canadian Wildlife Service, Environment Canada, Ot- tawa, Ontario, Canada. Accessed 19 January 2020. http://www.hww.ca/en/wildlife/birds/bald-eagle.html. (also issued in French under the title: Le pygargue a téte blanche). Stocek, R. 1992b. Book Review. Fishes of the Atlantic Coast of Canada (W.B. Scott and M.G. Scott. University of Toronto Press and Canadian Bulletin of Fisheries and Aquatic Sciences 219, 1988, 731 pages). N.B. Naturalist 19: 23. Stocek, R.F. 1993. Book Review: Ecology and Man- agement of the Eastern Coyote (A.H. Boer [Editor]. Wildlife Research Unit, University of New Brunswick, Fredericton, 1992, 194 pages). The Daily Gleaner, Fre- dericton. January 8, 113(6): 8. Stocek, R.F. 1994. Importance of riparian zones as wildlife habitat. Horizons (newsletter of the Fundy Model Forest Network) 1(2): 3. Stocek, R.F. 1995a. Book Review: Common sense wild- life management: discourses on personal experiences (N.R. Dickinson. Settle Hills Publishing, 1993, 107 pages). Atlantic Society of Fish and Wildlife Biologists Newsletter 11(1): 4. Stocek, R.F. 1995b. The Cougar, Felis concolor, in the Maritime Provinces. Canadian Field-Naturalist 109: 19-22. Accessed 26 January 2020. https://biodiversity library.org/page/35456881. Stocek, R.F., P. Cronin, and P.D. Seymour. 1999. The Muskellunge, a recent addition to the ichthyofauna of New Brunswick. Canadian Field-Naturalist 113: 230-— 234. Accessed 26 January 2020. https://biodiversity library.org/page/34235092. Stocek, R.F. 2000a. Diet of wintering Bald Eagles in New Brunswick. Canadian Field-Naturalist 114: 605-611. Accessed 26 January 2020. https://biodiversitylibrary. org/page/34237111. Stocek, R. 2000b. Nest utilization by Bald Eagles in New Brunswick. N.B. Naturalist 26: 107-110. Stocek, R. 2000c. The Bald Eagle: A species at risk in New Brunswick? Eléments (Online Environmental Magazine) New Brunswick Environment Network. De- cember 2000. Accessed 19 January 2020. http://www. elements.nb.ca/theme/endangeredspecies/bald/eagle. htm. 2019 Stocek, R. 2002. An Eagle-Eyed Odyssey. N.B. Naturalist 28: 99. Stocek, R.F. 2003a. The Bald Eagle population in New Brunswick. Bird Trends: a report on results of National Ornithological Surveys in Canada. No. 9: 21-24. Stocek, R.F. 2003b. Fifty forestry-focused frivolities: per- plexing problems for the restless mind. Privately Printed, R.F. Stocek, Fredericton, New Brunswick, Canada. Hood D.J., and R.F. Stocek. 2005. The Fathead Minnow in New Brunswick. Canadian Field-Naturalist 119: 351— 354. https//doi.org/10.22621/cfn.v11913.144 McALPINE AND ForRBES: A TRIBUTE TO R.F. STOCEK 275 Stocek, R. 2005. The Bald Eagle in the St. John River Watershed. The River (Newsletter of the St. John River Society). Summer 2005: 1—2. Stocek, R. 2006. Through the Eye of an Eagle: The Bald Eagle in New Brunswick. Rudy Stocek and Taylor Printing Group, Fredericton, New Brunswick, Canada. Stocek, R., and R. Eldridge. 2009. Fall and winter diet of coastal Bald Eagles in New Brunswick. NB Naturalist 36: 69-71. Received 19 December 2019 Accepted 29 January 2020 The Canadian Field-Naturalist Book Reviews Book Review Editor’s Note: The Canadian Field Naturalist is a peer-reviewed scientific journal publishing papers on ecology, behaviour, taxonomy, conservation, and other topics relevant to Canadian natural history. In line with this mandate, we review books with a Canadian connection, including those on any species (na- tive or non-native) that inhabits Canada, as well as books covering topics of global relevance, including climate change, biodiversity, species extinction, habitat loss, evolution, and field research experiences. Currency Codes: CAD Canadian Dollars, USD United States Dollars, EUR Euros, AUD Australian Dollars, GBP British Pounds. BOTANY Seaweed Chronicles By Susan Hand Shetterly. 2018. Algonquin Books of Chapel Hill. 271 pages, 27.59 CAD, Cloth. Seaweed Chronicles is a blend of engaging popular science and interview-based narrative. It is highly place- based—the Gulf of Maine and surrounding area is the main geographic focus— but the nature of the sub- ject matter means that con- nections are made between ocean coasts of all kinds. Shetterly has been involved in the regulatory and re- search communities for decades, and although her knowledge of the subject matter is apparent and the book is clearly well researched, this is not a treatise on the biology of seaweed. Several species of sea- weed and the creatures that depend on them are high- lighted and sufficient background is provided to in- form the uninitiated, but this is primarily a book of stories, lives lived in a relationship with algae. Each chapter focusses on one or a few related sub- jects, typically presented from the experience of a topic specialist via direct quotes and background in- formation. The writing is usually conversational in tone and covers integrated topics such as the cod fishery collapse, island sheep farming, potato gar- dening, invasive species, and eider ducks. I started reading this book while living in landlocked Ontario, but after living on the East Coast for a month, I found it significantly more engrossing. That is not to say that those without a coastal context will not enjoy Seaweed Chronicles, but I suspect having at least SEAWEED CHRONICLES A World: at the Waters Edge SUSAN HAND SHETTERLY casual contact with the ocean will only improve the reading experience. Shetterly does provide a short primer in the front matter on the algal species that feature prominently, but it would benefit from a measure of visual context—if you don’t know the species in question, it may help to start your reading with a quick Internet image search. While the author does refer to specific facts and findings, the book it- self does not contain references. What Seaweed Chronicles does exceptional- ly well is tell the stories of individual people. Per- spectives represented include those of researchers, farmers, harvesters, policymakers, and conserv- ationists, and their lived experiences are the foun- dation of the book’s content and structure. In this relatively short work, Shetterly delves into more di- mensions of seaweed than you ever knew existed— as habitat, foodstuffs, artisanal martini decoration, animal forage, restoration tool, fertilizer, and so on. Some stories will probably grip you more than others depending on your personal context, but the writing is accessible and there is almost certainly something within that will pique your interest. For those with a coastal upbringing or fond place-based memories, Seaweed Chronicles provides an enjoyable stroll along the water with a good teacher to reveal new layers of understanding. For those less familiar, it of- fers a window into a macroalgae world that is foun- dational to the health of our oceans. Either way, it is worth the read. HEATHER A. CRAY Halifax, NS, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 276 2019 ORNITHOLOGY Birds of Saskatchewan BooK REVIEWS DEL Edited by Alan R. Smith, C. Stuart Houston, and J. Frank Roy. 2019. Nature Saskatchewan. 765 pages, 79.95 CAD, Cloth. For over three-quarters of a century, Nature Sas- katchewan (formerly the Saskatchewan Natural Hist- ory Society) has promoted investigation of the natu- ral history of the province and surrounding areas by both amateurs and profes- sionals. Many of the re- sults have appeared in the quarterly Blue Jay, and in the Society’s many Special Publications. Birds have been the focus of much of this work. Birds of Saskatchewan is the culmination of these efforts. It began as the dream of the late Manley Callin (1911- 1985). His bequest helped fund its production and his dream is fulfilled by more than a decade of effort by the three editors, 107 authors of one or more species accounts, 69 photographers, and 24 reviewers. Readers will find much of interest and value on every page of this large book (30 x 23 cm; 3 kg). The 437 accounts cover all species occurring regularly in the province, plus extinct, accidental, and hypo- thetical species. There is much more than just spe- cies accounts in this book. The first section is an “Introduction to the Province”. It begins with a pres- entation of the seven “Natural Vegetation Zones in Saskatchewan”, describing their natural flora and listing their typical and special bird species. One or more beautiful photos illustrate each zone and give lie to the common belief that the province has noth- ing but flat wheat fields. “Human History and the Flora and Fauna of Saskatchewan” reviews the in- fluence of humans on the plants and animals, espe- cially since the beginning of European settlement in 1872. It describes the negative impacts of human activity such as agriculture and resource extraction on many avian species, and the positive effects on Species that have invaded or prospered in response to such activity. Efforts to protect and sustain bird populations are described, from the establishment of the first bird sanctuary in North America at Last Mountain Lake in 1887 to recent concerns about the federal government’s divestiture of the Prairie Farm Rehabilitation Administration pastures and its 1m- pact on populations of threatened grassland bird spe- cies. “The Ornithological History of Saskatchewan” begins with Henry Kelsey’s 1690 observation of Passenger Pigeons and Sir John Richardson’s orni- thological collections in 1827. A “who’s who” of pro- fessional ornithologists (John Macoun, A.C. Bent, Francis Harper, W.E. Clyde Todd, George M. Sutton, W. Earl Godfrey, and J. Dewey Soper, among others) visited the province between 1880 and 1947. Their Specimens are found in many of the major muse- ums in North America. The roles of resident natural- ists, bird banders, and institutions and organizations such as the Royal Saskatchewan Museum and Nature Saskatchewan are reviewed briefly. The section ends with a description of a century of “citizen science” which has contributed greatly to this book through Christmas Bird Counts (CBC), the Prairie Nest Records Scheme (PNRS), Breeding Bird Surveys (BBS), and through recent electronic activities such as Saskbirds (https://twitter.com/hashtag/saskbirds) and eBird (https://ebird.org/home). Accounts for species that occur regularly in the province are about four pages in length, and packed with information. Each begins with a brief in- troduction to the species and a description of the North American range. A small map of the prov- ince is colour-coded to indicate seasonal distribution. “History” summarizes records prior to extensive set- tlement (1924). “Status” reviews relative abundance and population trends based on BBS data and the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) designation, where appropri- ate. Sections on “Spring”, “Breeding”, “Fall”, and “Winter” summarize information on migration dates and breeding records. The final sections describe “Saskatchewan Research” and “Banding” (including names of banders with the number banded and recov- eries). One to five photos showing plumages, nests, young, and, in many cases, behaviour, illustrate each account. Accounts for permanent residents and win- ter visitors include a table summarizing CBCs for dif- ferent vegetation zones. Accounts for common water- fowl species include maps illustrating the recoveries of birds banded in the province, clearly demonstrat- ing the role of Saskatchewan as the “duck factory” of North America. Accounts for accidental (44) and hypothetical (42) species are a half-page or less in length. They summarize records for accidentals and available evidence for species whose occurrence in 278 Saskatchewan has not been documented with photo- graphs or sound recordings. Most of the accounts include a shaded “Interest Box”. Some provide special information (e.g., a short biography of Bernard Rogan Ross, for whom Ross’s Goose and Gull were named) or give taxonomic in- formation (e.g., the convoluted history of the scientific name for the Olive-sided Flycatcher). However, most recount specific experiences that contributors have had with the species in question. The late L.B. Potter describes (in the 1922 volume of The Canadian Field- Naturalist) the abundance of Sage-grouse in south- western Saskatchewan in the first decades of the 20th century. He notes a tameness which led them to tres- pass into the garden and eat the lettuce plants. Editor Alan Smith remembers a night in the 1960s that he spent sleeping on the prairie wool at the Matador Field Station, only to be woken in the early morning by a Vesper Sparrow who landed on his hip and used him as a song perch. Such vignettes capture the pleasures, rewards, and memories that we all derive from our ac- tivities in the natural world. Special mention must also be made of the pho- tographs that grace almost every page of this book. Many of these are nothing short of spectacular—it would be impossible to select a favourite! The Bo- hemian Waxwing on the cover is a good example. The photographers and the Photo Selection Committee de- serve congratulations for their efforts. Eight appendices follow the species accounts and cover various topics, including a list of bird banders mentioned in the accounts, a summary of results of CBCs from 1913 to 2016, a map of BBS localities (none in the northern third of the province), and a THE CANADIAN FIELD-NATURALIST Vol. 133 useful gazetteer of place names mentioned in the ac- counts. Appendix B includes biographical sketches of 168 now-deceased individuals who contributed to our knowledge of Saskatchewan ornithology. It includes explorers, early collectors, and professional ornithol- ogists who have worked in the province. But most contributors were farmers, ranchers, teachers, phy- siclans, accountants, homemakers, etc. who shared a common love of natural history and birds. On al- most every page, I found the name of someone who encouraged or supported a bird-crazy teenager on my path to a career in ornithology. I know that many oth- ers of my cohort (including the senior editor) shared this experience, and that it continues today, guaran- teeing that Saskatchewan ornithology will thrive in the 21st century. The book ends with a “Literature Cited” section spanning 49 pages and including approximately 2500 entries (my estimate)! I suspect that there few (if any) publications relevant to the avifauna of Saskatchewan that have been overlooked. Future researchers now have a single place to access relevant citations cov- ering information on observations and research from 1690 to 2016. Birds of Saskatchewan is an important record of the history and current state of the avian fauna of the province. It brings together a wealth of information that will be useful to both bird enthusiasts and future scholars. Beyond that, it is a delight to move through the pages, sampling both the data and the biological details contained. Its price is well worth the rewards of exploring a remarkable book. M. Ross LEIN University of Calgary, Calgary, AB, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 BooK REVIEWS 219 Feed the Birds: Attract and Identify 196 Common North American Birds By Chris Earley. 2019. Firefly Books. 296 pages, 29.95 CAD / 24.95 USD, Paper. Chris Earley’s book, Feed the Birds: Attract and Iden- tify 196 Common North American Birds, couldn't J identify have come at a better time. yee In October 2019, a land- ae mark paper was published documenting the catastro- & phic decline in North Amer- ican avifauna. Rosenberg et al. (2019: 120) wrote es their analyses indicate ‘ net loss approaching 3 billion birds, or 29% of 1970 abundance”. With an overall decline of avian species, even common species, we need to do everything we can to help our feathered friends. Feed the Birds is an easy-to-use book geared to- ward aiding in bird observation and study. The book begins with a brief introduction, asking questions like “why feed birds?” and delving into possible answers, such as citizen science, involving kids in birding, and photography. After the introduction, the book is di- vided into two major sections: (1) attracting and feed- ing birds and (2) identifying birds. The first section has four chapters: “Feeding Wild Birds” (what and how to feed); “Creating a Bird Friendly Backyard” (natural foods, water, shelter, nesting boxes, preda- tors, etc.); “Bird Feeder Building Plans”; and “Bird Behavior and Biology” (recognizing individual birds, nature’s predators, predator detectors, adaptions). The second section (comprising half the book) is focussed on bird identification, from hummingbirds and wood- peckers to grosbeaks and orioles. The book ends with a blank “birds at my feeder list”, works cited, further reading, photo credits, and an index. The first section is a joy to read. The writing is not long-winded and gets to the point. The para- graphs are packed with information, and each page is loaded with colour photos to support the text. For ex- ample, feeding wild birds is much more than tossing out a simple hardware store seed mix. Certain birds require certain kinds of food; different types of seeds attract different species of birds and some bird food types aren’t even seeds, like suet, fruits and jellies, nectar, and mealworms. What do you do with these foods? Toss them on the ground? Not always. There is a whole philosophy behind bird feeders and dispens- ers and, depending on what you want to attract to your yard, these feeders are critical. Creating a bird- friendly yard can also enhance the avian biodiversity in your area. Supplying a water source, nesting areas, shelters, and bird-attracting plants will no doubt in- Feed «Birds Attract and crease the number of birds in your yard. There are many things one needs to consider such as native versus non-native plants, berry-bearing shrubs and trees, plants that support invertebrates, and flowers that at- tract hummingbirds and other nectar feeders. Other must-read sections include how cats impact birds and the balance between enjoying the squirrels in your yard and when they become too much of a pest at the expense of your bird friends. Birds crashing into win- dows is another issue, and the book addresses this as well. As you watch the birds in your yard, you will be- gin to take note of various behaviours that may seem baffling at first, but with careful study, the mysteries of bird interactions, aggressive displays, and court- ship begin to tease out and demystify. Chapter 4, “Bird Behavior and Biology”, is a fascinating and welcome addition to your reading adventure. Various topics are covered, including feather maintenance, feeding behaviour, threat displays, nestling care, and bird intelligence. It’s always a joy to watch jays and crows figure out bird feeders as well as harass pesky squirrels. Of course, the biggest draw to attracting birds to your yard is bird identification. It’s an incredible thrill to keep a tally of bird species (life lists) and to add a rare bird from time to time. One huge reward is knowing that the bird likely arrived in your yard sim- ply because of the extra effort made to convert a once barren space into a bird paradise. The book discusses tips on how to identify birds, from size and shape, to beak morphology, bird movement, and field marks. The book includes a bird quick-find guide that assists in figuring out bird species, even if you only see the bird for a few moments. The guide will then direct you to a general bird group that will help with further identification. The general groups make up the last half of the book. Each group has species accounts, typically a page each, that provide additional infor- mation about the bird. Account topics include a gen- eral introduction to the bird, what natural foods they prefer, what feeds should be available in your yard, size comparisons, field identification marks, and a range map. All accounts are supported by several full colour photos of the bird species and sometimes ad- ditional side-by-side photo comparisons of closely re- lated or resembling species. Along the bottom of each account is a small call-out box with a photo and nat- ural history tidbit or anecdote about the bird. For ex- ample, in the American Robin account, the author discusses his observation of a robin “anting”, that is, 280 allowing ants to crawl on its feathers, crushing them, and rubbing the ants through the feathers. Anting is thought to be a form of chemical application to help repel external parasites. Overall, Feed the Birds is a must-read for those interested in attracting birds to their yard. With the general decline in bird species in North America and elsewhere, now is the time to create as many bird friendly spaces as possible. Chris Earley’s book is a one-stop shop for all you need to know to move for- ward with this wildlife enhancing concept. It’s easy to do and, with minimal effort, we can enjoy bird watching without even leaving the house. Be sure to participate in citizen science projects, like iNatural- THE CANADIAN FIELD-NATURALIST Vol. 133 ist or eBird, and let’s do our part to conserve and pre- serve avian biodiversity. Acknowledgement: I thank Susan Hagen for im- proving the review manuscript. Literature Cited Rosenberg, K.V., A.M. Dokter, P.J. Blancher, J.R. Sauer, A.C. Smith, P.A. Smith, J.C. Stanton, A. Panjabi, L. Helft, M. Parr, and P.P. Marra. 2019. Decline of the North American avifauna. Science 366: 120-124. https://doi.org/10.1126/science.aaw1313 Howarp O. CLARK, JR. Colibri Ecological Consulting, LLC, Fresno, California, USA ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 ZOOLOGY BooK REVIEWS 281 The North Atlantic Right Whale: Disappearing Giants. Revised and Updated Edition By Scott Kraus, Marilyn Marx, Heather Pettis, Amy Knowlton, and Kenneth Mallory. 2019. Fitzhenry & Whiteside. 140 pages, 24.95 USD, Paper. North Atlantic Right Whales (NARW; Fubalaena glaci- alis) have been in the news quite a bit over the last three years in Canada, be- = ginning in 2017 with the === deaths of 12 NARW in the Beitsssigapageiones = Gulf of St. Lawrence. This may not seem like a huge number, but with around 400 individuals left, mortal- ity events like this are noteworthy. The Government of Canada acted surprisingly quickly, enacting vessel slowdown measures and fisheries closures to reduce the risk of ship strikes and entanglements, respect- ively. This management strategy apparently worked, with no NARW found dead in Canadian waters in 2018. But 2019 was a dire season again, with eight or nine NARW found dead in Canadian waters. The National Oceanic and Atmospheric Administration of the United States have even labelled these mor- talities as Unusual Mortality Events. Disappearing Giants outlines the plight of NARW, paying particu- lar attention to the recent Unusual Mortality Event that the population underwent. This book provides a useful, relatively concise overview of the conserva- tion issues surrounding this species, and could be es- pecially interesting for Canadian readers who want to learn more about this species following the recent deaths in the Gulf of St. Lawrence. Disappearing Giants is a clearly written, non- technical overview of NARW, filled with wonder- ful photos of the whales. Close to 50% of the book is filled with photos, so beyond the interesting content, it would be a great book to leave out on a coffee table. This book is written by researchers from the New England Aquarium, some of whom, including main author Kraus, have been studying NARW since the 1970s. The book educates readers about NARW, with a brief chapter on evolution, followed by their hist- ory with humans, starting with centuries of whaling that devastated the population, to current research and The North Atlantic Right Whale DISAPPEARING GIANTS threats to the species. While all of the 14 species of baleen or great whales (Mysticetes) were target spe- cies for whalers, the ‘right whales’ to hunt were the three species of right whales (Eubalaena spp., includ- ing NARW) and Bowhead Whales (Balaena mystice- tus, Family Balaenidae) because they were easier to re- cover after they were killed: when they die, they tend to float on the surface, unlike rorqual whales (Family Balaenopteridae) that typically sink once they die. The commercial hunt for NARW ended in 1935 and, at that point, it was thought that only 100 whales were left. The population has recovered since then, but not as well as other right whale species, such as Southern Right Whales (Eubalaena australis). A main reason for this difference is that NARW live along the Atlantic coast of North America, where they are constantly ex- posed to ship traffic and active fishing grounds, lead- ing to continued human-caused mortality. Disappearing Giants doesn’t just focus on the bleak history of NARW. It ends on a chapter called “Hope for the Future”, where the authors describe reasons why we shouldn’t give up on NARW, and should continue working towards helping this spe- cies recover. The authors outline recent management initiatives that have been quite effective in reducing mortalities of NARW and, perhaps most importantly, describe the collaborative nature of NARW con- servation initiatives, where like-minded people have come together to address conservation issues sur- rounding NARW. These collaborations are a crucial aspect of the recovery of this species, and do indeed give me hope that the conservation issues surround- ing NARW are solvable, which will hopefully lead to recovery. WILLIAM D. HALLIDAY Wildlife Conservation Society Canada, Whitehorse, YT, and Department of Biology, University of Victoria, Victoria, BC ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 282 THE CANADIAN FIELD-NATURALIST Vol. 133 Yellowstone Cougars: Ecology Before and During Wolf Restoration By Toni K. Ruth, Polly C. Buotte, and Maurice G. Hornocker. 2019. University Press of Colorado. 336 pages, 65.00 USD, Cloth, 53.00 USD, E-book. Yellowstone Cougars is an @ academic-style book writ- | ten primarily for biologists and wildlife managers. It contains a treasure trove of data on Cougar (Puma con- color) and is the first book written on an apex carni- vore to examine their ecol- ogy before and after the re- covery of another keystone predator—in this case, Gray Wolf eae eS Given its scientific focus, the reading is very dense and time consuming with an impressive amount of data within its 300+ pages. The 8.25 x 10.25 inch (20.8 x 26.1 cm) hardcover contains small font and double columns per page so the book really felt like over 500 pages! Each chapter took me a couple hours to read given the length along with the technical infor- mation on each page. However, everything you want to know about Cougars in the world’s first national park is told here. This book and Cougar: Ecology and Conservation (2010, University of Chicago Press) are now the two reference books for this animal to which other works will be compared. Ruth and her colleagues conducted a 14-year study—seven years before wolves returned to Yel- lowstone National Park (1987-1994) and seven years (1998-2005) covering the tail end of wolf reintroduc- tion to the beginning of a recovered population. The book is broken into five main sections, the first con- sisting of three introductory chapters discussing their methods and the study area, followed by three mid- dle sections on Cougar diet and their competition with wolves (five chapters), landscape use (four chap- ters), and Cougar population characteristics pre- and post-wolf recovery (four chapters), and the final sec- tion contains two synthesis chapters on carnivores and humans. Each of the three middle sections has an introductory chapter which frames where the next three to four chapters will take us. Each chapter ends with a convenient summary of the most pertinent in- formation from that section, making it easier to digest the scientific information presented in that chapter— I often read those sections first before starting each chapter, then re-read it after I finished a given chapter. The authors spent an amazing amount of time in the field, marking 87-94% of the adult Cougars an- nually with 80 total Cougars radio-collared and ear- tagged (including all age classes) during pre-wolf studies and 88-93% post-wolf adult Cougars collared annually with 83 total tagged post-wolf (pp. 28, 69). Cougars were treed by hounds and then darted by biologists to sedate them. They were then followed so their movements and predation patterns could be recorded, with 40-50 kills found annually (p. 34). The researchers collected about 12 000 VHF (very high frequency) radio locations and over 19 500 GPS (global positioning system) points on these animals over 14 years. Their study area consisted mainly of the Northern Range of Yellowstone because Cougars only seasonally lived in the remaining 75—80% of the park due to deep snow and ungulates leaving those areas in winter, except for Bison (Bison bison) which they did not prey on (p. 58). We learned that Cougars were at the bottom of the large carnivore hierarchy, with wolves and Grizzly Bears (Ursus arctos) dominating them (pp. 93, 116), wolves most commonly, though rarely, killing them and bears most frequently usurping their kills (p. 244). Elk (Cervus canadensis) were the staple prey for Cougars and wolves throughout both study per- iods, with Mule Deer (Odocoileus hemionus) second for Cougars (p. 50). Calf Elk were the most import- ant prey class throughout the study for Cougars (p. 52). This remained the case even when Elk decreased at the end of the study owing to recovery of carni- vore populations, including bears, causing the system to change from bottom-up regulation of Elk before wolves to top-down post-wolf (p. 117). In general, wolves were superior at exploiting Elk adults and Cougars at exploiting Elk calves. Given their high niche overlap (82%, pp. 243-244), the sympatric car- nivores were unwittingly engaged in exploitation competition for a common food source (pp. 119, 244). Cougars survived by avoiding competitors, mainly wolves, by living in more forested and rougher terrain (p. 62), which contained a lower density of prey (p. 66). When prey Cougars killed was not taken over by competitors, Cougars often spent two to five days at a carcass before moving on to travel and eventu- ally kill again, usually three to four days after leav- ing their previous food cache (pp. 71—73). The auth- ors believed that Cougars benefited from using areas of lower densities of prey as this reduced potentially fatal encounters with wolves (pp. 179, 240, 242). Even so, wolves killed at least three adult Cougars and five kittens during the study (pp. 180, 208, 212). Interestingly, Cougar home ranges and core areas were more stable after wolf restoration compared to before (pp. 134-136). While Cougars used less area (females 10—46% and males 43—65%) on the landscape 2019 when wolves were back, they overlapped with more conspecifics sharing non-defended areas (i.e., females and philopatric daughters; pp. 137-138). In avoiding open and flat areas when wolves were back on the land- scape, the authors repeatedly stressed the importance of forested and rough terrain for Cougars (e.g., pp. 159, 176, 181, 235). The heterogenous habitats of northern Yellowstone likely makes carnivore coexistence pos- sible as each species used different areas (pp. 181, 242). Some unexpected findings of the study were, de- spite wolves engaging in exploitation (eating simi- lar prey) and interference (direct killing) competition with Cougars, the cats had similar litter sizes (aver- aging 2.9 cubs) throughout the study (p. 216) and kit- ten survival actually increased post-wolf (pp. 216, 202, 205) with less infanticide by adult male Cougars (p. 212). Because territories were more stable post- wolf, kittens actually stayed with their mothers for five months longer (12—14 versus 17-19 months) than before wolves came back (pp. 204, 220, 240), and liv- ing in small groups of adult-sized Cougars likely of- fered enhanced protection, intimidation, and vigi- lance from other predators (p. 230). In addition, the Cougar population increased post-wolf with about 30—40 total Cougars living in northern Yellowstone despite using a smaller percentage of overall habi- tat in the park (pp. 226, 230). Part of this can be ex- plained by Cougars recolonizing the area in the 1980s (p. 21) and then becoming saturated on the land- scape as wolves came back. Densities of Cougars of two adults and 3.9 total per 100 km? in the study area were actually on the high end compared to other Cougar populations (p. 225). With this fully occu- pied area, females—with no room to stay near their mothers—averaged the same dispersal distance as young males (70 km; p. 221) and females moved more home range diameters away than did males (pp. 224, 254). Similarly, while 35% of females before wolves were philopatric only 11% were so post-wolf (p. 220). These young dispersers contributed to Yellowstone being a source population to nearby areas (pp. 255— 256). Source populations are helping Cougars recover and colonize long vacant areas like the midwestern and even eastern United States (Way 2017: 249). Yellowstone Cougars is comparable to the in- credibly detailed and well-researched books Desert Puma (2001, Island Press) and Mountain Lions of the Black Hills (2018, Johns Hopkins University Press; Way 2018) in that it does a superb job of describing an in-depth long-term study on Cougars in a specific region. Yellowstone Cougars includes: 10 pages of Appendices explaining their study variables; 10 pages of “Notes” which are detailed statistics described in the chapters but shown at the end of the book to avoid too much detail in each section; an impressive 30 BooK REVIEWS 283 pages of double-columned references; and a seven- page index. It takes six pages at the beginning of the book to list the titles of the illustrations, including 118 figures and 60 tables. Many of those figures are black and white pictures of wild Cougars from the study, impressive because Cougars are notoriously diffi- cult to photograph. My only complaint was that there was no map displaying dispersal distances from the source population when the authors discussed emi- gration (pp. 219, 221). Also, I did notice a few errors on some of the figures, including wrong labels in the charts (e.g., Figure 11.1 on p. 155, Figure 11.14 on p. 173, Figure 15.4 on p. 207, and Figure 16.4 on p. 230). The reading material is labourious to go through thoroughly but is vital to understanding Cougar ecol- ogy in Yellowstone. I found Part 5, Carnivores and Humans: Competition and Coexistence, to be par- ticularly important because it provided a synthesis of the book and offered management and conserva- tion recommendations for the big cats. I was a little disappointed with the last chapter (18) in that it de- scribed management and conservation of Cougars but did not actually offer any concrete management op- tions for state agencies. For instance, their data (see Figure 18.1, p. 253) showed that many female Cougars killed by hunters left orphaned offspring that died via starvation (p. 250). The authors do suggest manage- ment regimes where non-parklands also include areas closed to hunting to mimic natural populations (p. 250). These areas can be managed adaptively through rest rotation, whereby periods of hunting alternate with periods of rest (p. 248). However, without any specific suggestions of where these could occur, my experience with carnivore management suggests that even with involving citizens in a bottom-up approach (p. 258) it is difficult to envision state wildlife agen- cies doing anything other than continuing with kill- ing the maximum sustainable amount of a species— even an ecologically important predator. For enthusiasts of Yellowstone or carnivores, this 1s an important book. Unfortunately, and like many aca- demic-style texts, Yellowstone Cougars is expensive. However, the book is truly a benchmark in detailing the life history of an elusive and difficult to study species. Literature Cited Way, J.G. 2017. [Book Review] Heart of a Lion: A Lone Cat’s Walk Across America. Canadian Field-Naturalist 131: 82—84. https://doi.org/10.22621/cfn.v13111.1972 Way, J.G. 2018. [Book Review] Mountain Lions of the Black Hills: History and Ecology. Canadian Field- Naturalist 132: 200-201. https://doi.org/10.22621/cfn.v 13212.2189 JONATHAN (JON) Way Eastern Coyote/Coywolf Research, Osterville, MA, USA ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 284 OTHER THE CANADIAN FIELD-NATURALIST Vol. 133 How to Give Up Plastic: A Guide to Changing the World, One Plastic Bottle at a Time By Will McCallum. 2018. Penguin Random House. 224 pages, 15.00 USD, Paper. Plastic. No matter where we look, it is everywhere, whether we see it or not. | The impact that plastic has & been having on wildlife and § ecosystems has extended Raga past the environmental field § and has exploded into main- &: stream media. It can almost feel like we are bombarded by the various ways to gi become more eco-friendly and adopt a zero-waste life- style, leaving many people feeling like aoe contribu- tions will be minimal at best. As someone who is al- ways trying to reduce their footprint and encouraging others to do the same, this book was one I was par- ticularly excited to check out. How to Give Up Plastic begins with a wake-up call. The first two chapters focus entirely on the plas- tic problem and include many astonishing statistics and research that I was unfamiliar with. It was very interesting to learn the history of how plastics became so prevalent in our lives, how they have evolved, and how our recycling systems are not what we, as regu- lar consumers, believe them to be. These chapters paint a somewhat glum picture of where our reliance on plastic has brought us, countered in the next chap- ter, “Stories of Hope and Success”, showing how one person or one group created a huge difference in their community and beyond. Throughout the book you can find mini interviews relating the experiences of people leading the charge in the fight against plas- tics. I thoroughly enjoyed reading their views, tips, and reasons for doing the work that they do. The next five chapters take us through different areas in our homes and lives. Each chapter breaks its area down into the most common items one might use (for example, the bathroom chapter includes sec- tions on lip balm, shampoo, make-up, and hair re- moval). Some of the categories mention businesses that are targetting certain waste forms by creating al- ternatives. This information is very helpful in giving you a place to begin searching for items that suit your lifestyle; however, it could go quickly out of date as businesses come and go. While most of this advice is available through internet searches, having it all in one location to read through puts the bigger picture together and allows you to see where you want your plastic-free life to begin. As someone who has begun changing my lifestyle to lessen my waste and use of single-use plastics, I was happy to find many items that weren’t on my radar and a few new blog sug- gestions! Many of the chapters end with a work page where you can list your plastic-free plan based on the topics covered. Chapters 10 and 11 take you from targetting your individual plastic use to your workplace and com- munity. They are full of ideas, from getting people motivated to using your vocabulary to engage others to join the cause. The chapter on community gives a step-by-step guide to running your own clean-up, writing an effective letter to your members of gov- ernment, and hosting a protest. These are activities that I think many people would like to be involved in and this allows them to take the next step in the actual planning process. This is a great book for those who are relatively new to being plastic-free. It has tips and tricks for your everyday life and acts as an easy access, easy- to-read guide to start making your plastic-free plan. What felt like almost a blog-type format kept the con- tent engaging and easy to read. I appreciated that the author consistently highlighted the need for systemic change, beginning with the industry, and an under- standing that different realities exist for different people and can inhibit their ability to fully give up plastics. I typically expect books like this to be rela- tively preachy, but I also appreciated the fact that the author did a good job of being non-judgemental while still giving solid advice and statistics on the repercus- sions of plastic use. I would recommend this book to anyone interesting in beginning to reduce the amount of plastic in their life or anyone currently on the plas- tic-free journey as it may provide some topics they haven't thought of yet. TIANNA BURKE Conservation Biologist, Georgian Bay Biosphere Reserve, Parry Sound, ON ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 Plastic Soup: An Atlas of Ocean Pollution BooK REVIEWS 285 By Michiel Roscam Abbing. 2019. Island Press. 136 pages, 27.00 CAD, Cloth or E-book. Plastic is in almost every item we use and own. Its convenience as a relative- ly cheap and durable ma- terial has become rather inconvenient from an en- vironmental standpoint. For a few years now I have per- sonally tried to make my household less reliant on plastic and am continuous- ly surprised at just how dif- ficult it is. Whether it 1s pur- chasing food, soap, pet products, or craft supplies, it has been incredibly difficult to manoeuver in a world where things are not individually wrapped or contain plastic. Plastic Soup: An Atlas of Ocean Pollution is a short book with a lot of impact, both by the writ- ten content and the visual content. Author Michiel Roscam Abbing 1s a political scientist actively work- ing on the plastic soup problem since 2011 alongside the Plastic Soup Foundation. Trying to put an end to increasing amount of plastic pollution, the Plastic Soup Foundation works to tackle plastic issues at the source, something that is focussed on in this book. Plastic Soup is separated into two distinct parts; the first, “On the Map”, focusses on the plastic crisis: its creation, the effects plastic has on ecosystems and wildlife, and some of the major items contributing to the plastic problem. The second half of the book, “Off the Map”, focusses on solving the plastic crisis, high- lighting research and initiatives around the world, the introduction of laws, and even how art is bringing this issue into the forefront. More than just an atlas of pollution, this book is also an atlas of hope. I found it so interesting to learn how many different stakeholders across the world are reducing the use of plastic. Being someone who loves food, the idea of lasered food to reduce packaging and stickers was especially of interest. This book also challenged the optics of plastic itself and some of the plastic solutions that are becoming popular. I found PLASTIC MICHIEL ROSCAM ABBING it interesting to read that, technically, plastic reduces food waste because it helps to extend the shelf life of many items. Similarly, it takes fewer emissions and less water to produce than paper does, another com- mon packaging item. However, while this may seem like plastic is an obvious solution, the total lifecycle of the product says otherwise, emphasizing the need to think critically and in terms of lifecycles. Critical thinking came up again in Chapter 7, “Between Belief and Hope”, which tackles subjects such as recycling, bioplastics, and creating products out of ocean plastic. While these ideas may seem bril- liant, they may be better than the actual results. Many books tend to focus on what you can do at an individ- ual level and, while this book points out the roles our purchasing and lifestyle choices play, I was impressed by the author’s emphasis on change at a level greater than a household. It helped me realize what more I can be doing at home and provided ideas I can push through to my local municipality and government. I really enjoyed how in this first section the topics moved from obvious plastics, such as balloons, to plastics that we cannot see, such as microplastics. The author did a great job informing the reader of the problems and delivering the scientific evidence in a way that reaches all audiences. It was easy to understand, and the written content was enhanced by stunning photography and infographics. The for- mat of the book makes it easy to read, providing a valuable tool for people seeking to learn more about the plastics issue. Throughout, heart-wrenching im- ages bring home very effectively the message of the damage plastic has done to our planet and wildlife. Even if you don’t read the book, the images alone will make you want to change your habits! I would recom- mend this book to anyone who is interested in learn- ing more about plastics or for those, like me, who are trying to teach others and could use a resource jam- packed with information. TIANNA BURKE Conservation Biologist, Georgian Bay Biosphere Reserve, Parry Sound, ON ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 286 THE CANADIAN FIELD-NATURALIST Vol. 133 Mama’s Last Hug: Animal and Human Emotions By Frans de Waal. 2019. W.W. Norton. 336 pages, 36.95 USD, Paper. Humans have held them- selves superior to all other life forms for millennia. Dating back to Aristotle, this attitude in Western cul- tures was crystalized in the biblical notion that ‘man will have dominion over the earth and all the creatures therein’. The consequences of this belief, and the subse- quent actions over succeed- ing millennia, have been disastrous for the animals, as well as the earth itself. While ecologists, environmentalists, and most stu- dents of the life sciences are increasingly recogniz- ing, defining, and warning us of these consequences, the notion of human supremacy is one that still re- mains strong. Why this should be so is a key question that primatologist Frans de Waal addresses in this, his 12th book, a “companion” to his Are We Smart Enough to Know How Smart Animals Are?, pub- lished by Norton in 2016. As de Waal explains in his Acknowledgments, “[e]ven though these two books treat emotions and cognition separately, in real life they are fully integrated” (p. 279). The arguments for animal intelligence and emotional lives presented by de Waal are compelling, supported by the increas- ing research in these areas and the rich anecdotal evi- dence gathered during his own long experience with primates and from other primatologists. Mama’s Last Hug begins with just such a story. We meet Mama as a 50-year-old Chimpanzee on her death bed. A researcher who had spent much time with her but who had not seen her in several years appeared for a final visit. On seeing him, Mama was transformed, from a listless animal on its way out to an excited, expressive creature that greeted her old friend effusively. How this could be seen in any other way as an emotional response is the mystery that de Waal seeks to unravel. After a brief Prologue, the book continues for seven chapters. The first, “Mama’s Last Hug”, relates the story noted above; the next three discuss various emotions. Many of us conflate emotions with feelings, but de Waal distinguishes between them, defining feelings as interior states that we can describe using language and emotions as the deeply rooted, initially subconscious states that emerge into consciousness during various situations. The distinction is helpful, allowing him to address the idea that because ani- mals don’t have words to express emotion, they do not feel emotion; they simply react behaviourally to various stimuli in instinctual ways. This idea is not to be underestimated in its force—centuries of animal research have been premised on it. Chapters 2 and 3 present evidence of positive emotions in animals— laughter, empathy, sympathy—while Chapter 4, “Emotions That Make Us Human”, deals with nega- tive emotions, including disgust, guilt, and shame. These chapters contain many instances, observed in the wild or concluded from ingenious experiments, demonstrating the reality of animal emotional lives. Two themes running through these chapters, and in- deed the book, are the continuity between the behav- ioural responses of apes and humans, and the con- tinuing, though diminishing, resistance of scientists to accept or, more accurately, to write as though they accept, that apes have emotions just as humans do. These chapters are the foundation for the more difficult, controversial discussions in the next three, Chapter 5, “Will to Power — Politics, Murder, War- fare”, Chapter 6, “Emotional Intelligence — On Fair- ness and Free Will”, and Chapter 7, “Sentience — What Animals Feel”. If you cannot accept that animals have emotions, then it will be next to impossible to accept, as argued in Chapters 5 and 6, that animals have complex political relations, can engage in mur- der and warfare, or choose to act with fairness, and have the capacity to think through the consequences presented at times decisions are required. But the evi- dence is strong, the stories compelling. If one accepts evolutionary continuity between apes and humans, de Waal’s conclusions are inescapable. Chapter 7 is the capstone of the book. It begins by exploding the long-held belief that human superi- ority is based on the size of our brains and number of neurons therein. Recent research has shown that elephants have more of both than we do! And the re- lated myth that consciousness is a property of hu- mans alone gets similar treatment. Not only that, but instinct as sole explainer of animal actions is it- self relegated to the dustbin of historical ideas. In the process of making these remarkable conclusions, de Waal discusses “three reasons (apart from press- ing ecological ones) that humans should respect all forms of life: the inherent dignity of all living things, the interest every form of life has in its own exist- ence and survival, and sentience and the capacity for suffering” (p. 245; italics in the original). He admits that assigning dignity to all forms of life is based on our subjective evaluations, the danger we must guard against is falling back into the ancient concept of what the Elizabethans called the great chain of being. 2019 It is more readily seen that living things have an inter- est in remaining alive. While this is obvious from the reactions of animals, from mammals to arthropods, it remains true of plants, which, science is discovering, have incredibly complex defensive systems. The big reason for respecting all forms of life, however, is sentience, the idea, impossible to con- firm with scientific certainty, that animals have con- scious experience of their emotions. Surely this must be an essentially human capacity. Well, not so surely, it turns out. All creatures, from cells to fungi, plants, and animals, have some capacity for sentience, or the ability to adjust their experienced conditions. But “Ts]entience in the narrow sense implies subjective feeling states, such as pain and pleasure” (p. 248). It is de Waal’s view that all living creatures, with and without brains and central nervous systems, should be considered as “sentient in the sense of having subject- ive feeling states” (p. 249). And this form of sentience resulted, de Waal believes, in the development of con- sciousness “relatively early in evolution” (p. 255). The acceptance of these ideas is still ongoing, al- though science has come a long way from the early days of research into “affective neuroscience”, a disci- pline founded by Jaak Panksepp, who “was ahead of his time...” (p. 256). In Panksepp’s day, relates de Waal, funding for such research was difficult to come by, so strong was the opposition to animal emotions and intelligence, particularly in the field of psychol- ogy, dominated by Skinnerian behaviourism. My first degree was in psychology during the heyday of this movement, which I rejected instinctively. I took a per- sonal delight in reading de Waal’s description of the movement’s demise. Unfortunately, its lingering leg- acy is the “gap between humans and all other spe- cies, which only widened with time” (p. 260). The re- sults of that gap are still being promulgated in books BooK REVIEWS 287 celebrating human exceptionalism, but meanwhile, “Tb]ehaviourism is dying a slow death” (p. 262). And about time. As noted in my review (Cottam 2018) of Through a Glass Brightly, people who reject the no- tion that humans are animals need to elevate their concepts of what animals are. Personally, I find it comforting to think that rather than dwelling on some fictional peak, we humans are connected with all liv- ing matter, part of the great natural cycle of life and death, the only ‘eternity —should we manage not to destroy the earth—that we can know. Much of the evidence in this book is derived from field experience, whether in the jungles and other habitats where the animals live, or in the hu- mane environments in which many research ani- mals now reside, relatively free to interact in their normal social ways. It’s highly readable ‘popular’ science at its best, but the topics are huge and crit- ically important, the concepts revolutionary if we accept them. Thus it provides some hope that we humans will realize that continuing to consider our- selves superior to all other forms of life is just what it takes to destroy our own. Editor’s note: I used an advance reading copy to review this book. The final publication will differ somewhat—it will be indexed, for example, and has a different cover—so page numbers for quotations in this review may not exactly match those in the pub- lished version. Literature Cited Cottam, B. 2018. [Book review] Through a Glass Brightly: Using Science to See Our Species as We Really Are. Canadian Field-Naturalist 132: 202—203. https://doi.org/ 10.22621/cfn.v132i2.2191 BARRY COTTAM Corraville, PE, and Ottawa, ON ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 288 North Pole: Nature and Culture THE CANADIAN FIELD-NATURALIST Vol. 133 Michael Bravo. 2019. Reaktion Books. 254 pages and 111 illustrations, 62 in colour. 24.95 USD, Cloth or E-book. Michael Bravo, Head of Circumpolar History and Public Policy Research at the Scott Polar Research Institute, Cambridge, Uni- ted Kingdom, has written a rather unusual book. As the title suggests, it deals with the North Pole, but it is extraordinarily eclectic, ranging from classical writ- ings on the polar regions, through the speculations of renaissance geographers, to accounts of polar exploration in the 18th through 20th centuries, and includes diversions into different sorts of poles (astronomical, geographical, magnetic) and polar exploration in cartoons and satirical writ- ing. We meet Madame Blavatsky, Scipio Africanus, Herakles, and Baron Munchausen, among many oth- ers. Some are characters we might expect to see at the North Pole (Peary, Amundsen, Nansen), while others come as a total surprise (Mary Shelley and Frankenstein, Ptolemy, Bal Gangadhar Tilak). This is not a book for those who primarily want factual information about the North Pole, although some of that is included. It is more likely to appeal to those who enjoy a ramble through miscellaneous po- lar ‘factoids’. Among the great names of polar travel, Peary gets quite a bit of space, although the contro- versy about where he actually got to is referenced but not described in detail, and Cook only gets passing mention. Steffanson, although never attempting to approach the pole, gets fairly extensive treatment, but I felt that Nansen got rather short shrift. There is much in the book to be cherished regarding the impact of the pole on literature and art, and there are some lovely and, I suspect, little-known, images. However, I was constantly asking myself whether the book is really serious or a very well-disguised send- up of arcane scholarship. For example, after mention- ing the section in Winnie-the-Pooh where Pooh finds a pole (he “just found it”) and Christopher Robin an- nounces that it must be the North Pole, Bravo makes the following suggestion (p. 158): Milne, diverging from ethnonationalists who elevated the status of the North Pole to that of an ur-site of Aryan origins, recognised it for what it was, the essential point of origin in a mathematical projection but philosophically no more special than anywhere else ... The book is very attractively produced and illus- trated on wonderful glossy paper. It is very entertain- ing to thumb through and browse and only the most diligent student of things polar is likely to be famil- iar with all the material covered. However, the Pooh excerpt given above is just a rather extreme example of the book’s generally over-erudite and, to my mind, over-elaborate, approach to the topic. Recommended for generalists and romantics. Not recommended for those only wanting information on polar exploration. TONY GASTON Ottawa, ON ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 NEw TITLES Prepared by Barry Cottam NEw TITLES 289 Please note: Only books marked ¢ or * have been received from publishers. All other titles are listed as books of potential interest to subscribers. Please send notice of new books to the Book Review Editor. +Available for review *Assigned Currency Codes: CAD Canadian Dollars, AUD Australian Dollars, USD United States Dollars, EUR Euros, GBP British Pounds. BIOLOGY Biodiversity and Evolution. Edited by Philippe Grand- colas and Marie-Christine Maurel. 2018. ISTE Press— Elsevier. 284 pages, 127.50 USD, Cloth, 97.75 USD, E-book. The Biology of Parasites. By Richard Lucius, Bri- gitte Loos-Frank, Richard P. Lane, Robert Poulin, Craig W. Roberts, and Richard K. Grencis. Translated by Renate FitzRoy and Ron Shankland. 2017. Wiley- Blackwell. 472 pages, 118.75 CAD, Cloth. Biology of Sex. By Alex Mills. 2018. University of Toronto Press. 432 pages, 170.00 CAD, Cloth, 77.00 CAD, Paper, 62.00 CAD, E-book. Is That a Bat? A Guide to Non-Bat Sounds En- countered During Bat Surveys. By Neil Middleton. 2020. Pelagic Publishing. 288 pages, 59.39 CAD, Paper. The Evolutionary Biology of Species. Oxford Se- ries in Ecology and Evolution. By Timothy G. Bar- raclough. 2019. Oxford University Press. 288 pages, 90.00 CAD, Cloth, 45.95 CAD, Paper. Also available as an E-book. Field and Laboratory Methods in Animal Cog- nition: A Comparative Guide. Edited by Nereida Bueno-Guerra and Federica Amici. 2018. Cambridge University Press. 456 pages, 137.95 CAD, Cloth, 62.95 CAD, Paper, 44.00 USD, E-book. Fundamentals of Microbiome Science: How Mi- crobes Shape Animal Biology. By Angela E. Doug- las. 2018. Princeton University Press. 248 pages, 39.50 USD, Cloth. Also available as an E-book. Mate Choice: The Evolution of Sexual Decision Making from Microbes to Humans. By Gil G. Rosenthal. 2017. Princeton University Press. 648 pages, 55.00 USD, Cloth. Also available as an E-book. Quantitative Biology: Theory, Computational Methods, and Models. Edited by Brian Munsky, William S. Hlavacek, and Lev S. Tsimring. 2018. MIT Press. 728 pages, 70.00 USD, Cloth. Social Evolution and Inclusive Fitness Theory: An Introduction. James A.R. Marshall. 2019. Princeton University Press. 216 pages, 27.95 USD, Paper. Cloth and E-book editions published in 2015. BOTANY Applied Tree Biology. By Andrew D. Hirons and Peter A. Thomas. 2018. Wiley Blackwell. 432 pages, 62.00 CAD, Paper, 49.99 CAD, E-book. The Great Fen: A Journey Through Time. By Alan Bowley. Illustrated by Oliver Bowley. Foreword by Tony Juniper. 2019. Nature Bureau Pisces Publications. 194 pages, 27.50 GBP, Cloth. The Nature of Plants: An Introduction to How Plants Work. By Craig N. Huegel. 2019. University Press of Florida. 288 pages, 24.95 USD, Paper. The Sphagnum Species of the World. Bibliotheca Botanica Series, Volume 162. By Dierk Michaelis. 2019. Schweizerbart Science Publishers. 435 pages and 219 plates, 159.00 EUR, Cloth. Sprout Lands: Tending the Everlasting Gift of Trees. By William Bryant Logan. 2019. W.W. Norton. 384 pages, 27.95, Cloth, 17.95 USD, Paper. Trees of Life. By Max Adams. 2019. Apollo. 256 pages, 9.99 GBP, Paper. Witness Tree: Seasons of Change with a Century- Old Oak. By Lynda V. Mapes. 2017. Bloomsbury USA. 240 pages, 27.00 USD, Cloth, 18.90 USD, E-book. CLIMATE CHANGE Biodiversity and Climate Change: Transforming the Biosphere. Edited by Thomas E. Lovejoy and Lee Hannah. Foreword by Edward O. Wilson. 2019. Yale University Press. 416 pages, 40.00 USD, Paper. Floating Coast: An Environmental History of the Bering Strait. By Bathsheba Demuth. 2019. W.W. Norton. 416 pages, 27.95 USD, Cloth, 17.95 USD, Paper. Invasive Species and Global Climate Change. Edit- ed by Lewis Ziska and Jeffery Dukes. 2019. CABI. 366 pages, 70.00, Paper. Cloth edition published in 2014. 290 Living in the Anthropocene: Earth in the Age of Humans. Edited by W. John Kress and Jeffrey K. Stine. Foreword by Elizabeth Kolbert. Afterword by Edward O. Wilson. 2017. Smithsonian Books in asso- ciation with Smithsonian Institution Scholarly Press. 208 pages, 34.95 USD, Cloth. Losing Earth: A Recent History. By Nathaniel Rich. 2019. Farrar, Strauss and Giroux. 224 pages, 25.00 USD, Cloth, 16.00 USD, Paper. Structures of Coastal Resilience. By Catherine Seavitt Nordenson, Guy Nordenson, and Julia Chap- man. 2018. Island Press. 248 pages, 80.00 USD, Cloth, 45.00 USD, Paper or E-book. When The Seas Rise: Global Changes and Local Impacts. By Heather Dewar. 2016. University Press of Florida. 120 pages, 14.95 USD, Paper. CONSERVATION & ECOLOGY Abundant Earth: Toward an Ecological Civiliza- tion. By Eileen Crist. 2020. University of Chicago Press. 288 pages, 105.00 USD, Cloth, 35.00 USD, Paper. Also available as an E-book. Climate Change Impacts on Urban Pests. CABI Climate Change Series 10. Edited by Partho Dhang. 2017. CABI. 200 pages, 132.80 USD, Cloth. Community-Based Control of Invasive Species. Edited by Paul Martin, Theodore R. Alter, Donald W. Hine, and Tanya M. Howard. 2019. CABI. 288 pages, 105.00 USD, Cloth. Complex Ecology: Foundational Perspectives on Dynamic Approaches to Ecology and Conserva- tion. Edited by Charles G. Curtin and Timothy F.H. Allen. 2018. Cambridge University Press. 594 pages, 140.00 USD, Cloth, 54.99 USD, Paper. Conservation Biology. By Bradley Cardinale, Rich- ard Primack, and James Murdoch. 2019. Sinauer As- sociates, an imprint of Oxford University Press. 672 pages, 114.95 CAD, Cloth. Also available as an E-book. Ecological Effects of Electricity Generation, Stor- age and Use. By Peter Henderson. 2018. CABI. 240 pages, 45.00 USD, Paper. Ecoviews Too: Ecology for All Seasons. By Whit Gibbons and Anne R. Gibbons. 2017. University of Alabama Press. 224 pages, 24.95 USD, Paper or E-book. Evolutionary Community Ecology. Monographs in Population Biology, Volume 58. By Mark A. McPeek. 2017. Princeton University Press. 324 pages, 60.00 USD, Cloth. THE CANADIAN FIELD-NATURALIST Vol. 133 The Fall of the Wild: Extinction, De-Extinction, and the Ethics of Conservation. By Ben A. Minteer. 2018. Columbia University Press. 192 pages, 28.00 USD, Cloth, 27.99 USD, E-book. Forests Adrift: Currents Shaping the Future of Northeastern Trees. By Charles D. Canham. 2020. Yale University Press. 240 pages, 28.00 USD, Paper. Marine Conservation. By P. Keith Probert. 2017. Cambridge University Press. 517 pages, 99.99 USD, Cloth, 54.99 USD, Paper. Open Ecosystems: Ecology and Evolution Beyond the Forest Edge. By William J. Bond. 2019. Oxford University Press. 192 pages. 75.00 CAD, Cloth. Also available as an E-book. The Rights of Nature: A Legal Revolution That Could Save the World. By David R. Boyd. 2017. ECW Press. 312 pages, 12.95 CAD, Paper, 14.99 CAD, E-book. ENTOMOLOGY Bees of Australia: A Photographic Exploration. By James Dorey. 2018. CSIRO Publishing. 222 pages, 49.99 AUD, Paper. Courtship and Mating in Butterflies. By R.J. Cannon. 2020. CABI. 384 pages and 242 colour pho- tographs, 160.00 USD, Cloth. The Dark Side of the Hive: The Evolution of the Imperfect Honey Bee. By Robin Moritz and Robin Crewe. 2018. Oxford University Press. 203 pages, 80.00 CAD, Cloth. Also available as an E-book. The Discovery of a Visual System—The Honeybee. By Adrian Horridge. 2019. CABI. 296 pages, 120.00 USD, Cloth. Dragonflies & Damselflies: A Natural History. By Dennis Paulson. 2019. Princeton University Press. 224 pages, 29.95 USD, Paper. The Economics of Integrated Pest Management of Insects. Edited by David W. Onstad and Philip Crain. 2019. CABI. 232 pages, 120.00 USD, Cloth. Forest Insect Population Dynamics, Outbreaks, and Global Warming Effects. By A.S. Isaev, V.G. Soukhovolsky, O.V. Tarasova, E.N. Palnikova, and A.V. Kovalev. 2017. Wiley and Scrivener Publishing. 304 pages, 281.00 CAD, Cloth. Insects and Society. By Timothy D. Schowalter. 2019. CRC Press. 306 pages and 157 colour illus- trations, 87.99 GBP, Cloth, 43.99 GBP, Paper. Also available as an E-book. 2019 Insect Behavior: From Mechanisms to Ecological and Evolutionary Consequences. Edited by Alex Cordoba-Aguilar, Daniel Gonzalez-Tokman, and Isaac Gonzalez-Santoyo. 2018. Oxford University Press. 404 pages, 100.00 USD, Cloth, 49.95 USD, Paper. Insect Collection and Identification: Techniques for the Field and Laboratory. By T.J. Gibb and C. Oseto. 2019. Academic Press. 342 pages, 76.95 GBP, Paper. Insect Conservation: A Global Synthesis. By Michael J. Samways. 2019. CABI. 600 pages, 205.00 USD, Cloth, 90.00 USD, Paper. Insects Did It First. By Gregory S. Paulson and Eric R. Eaton. 2018. Xlibris Corporation. 156 pages, 29.99 USD, Cloth, 19.99 USD, Paper, 3.99 USD, E-book. Insect Mouthparts: Form, Function, Development and Performance. Edited by Harald W. Krenn. 2019. Springer Nature. 695 pages, 159.99 USD, Cloth, 119.99 USD, E-book. Life Cycles of British & Irish Butterflies. By Peter Eeles. Foreword by Chris Packham. 2019. NatureBureau. 400 pages and 1300 colour photos and maps, 35.00 GBP, Cloth. True Bugs of the World (Hemiptera: Heteroptera): Classification and Natural History. Monograph Series Volume 8. By Randall T. Schuh and Christine Weirauch. 2020. Siri Scientific Press. 800 pages, 129.99 GBP, Cloth. Stingless Bees of Mexico: The Biology, Manage- ment and Conservation of an Ancient Heritage. By José Javier G. Quezada-Euan. 2018. Springer International Publishing. 304 pages, 169.99 USD, Cloth, 159.99 USD, Paper, 119.00 USD, E-book. Urban Landscape Entomology. By David Held. 2019. Academic Press. 224 pages, 79.96 USD, Paper or E-book. Wasp. By Richard Jones. 2019. Reaktion Books. 208 pages and 114 illustrations, 12.95 GBP, Paper. HERPETOLOGY Fossil Frogs and Toads of North America. Life of the Past. By J. Alan Holman. 2018. University of Indiana Press. 261 pages, 30.00 USD, Paper. Originally published in 2013. Salamanders: Habitat, Behavior and Evolution. Edited by Rashid Gerasimov. 2019. Nova Science Publishers. 154 pages, 82.00 USD, Paper. ICHTHYOLOGY Age and Growth of Fishes: Principles and Tech- niques. Edited by Michael C. Quist and Daniel A. NEw TITLES 291 Isermann. 2017. American Fisheries Society. 359 pages, 79.00 USD, Cloth. The Behavior and Ecology of Pacific Salmon and Trout. Second Edition. By Thomas P. Quinn. 2018. University of Washington Press in association with the American Fisheries Society. 554 pages, 60.00 USD, Paper. From Catastrophe to Recovery: Stories of Fishery Management Success. Edited by Charles C. Krueger, William W. Taylor, and So-Jung Youn. 2019. American Fisheries Society. 530 pages, 79.00 USD, Paper. Fishes of the Salish Sea. Volume One: Puget Sound and the Straits of Georgia and Juan de Fuca. By Theodore Pietsch and James Wilder Orr. Illustrated by Joseph R. Tomelleri. 2019. Heritage House. 1032 pages, 179.00 CAD, Cloth. Managing Centrarchid Fisheries in Rivers and Streams. Edited by Michael J. Siepker and Jeffrey W. Quinn. 2019. American Fisheries Society. 270 pages, 79.00 USD, Paper. The Ocean Ecology of Pacific Salmon and Trout. Edited by Richard J. Beamish. 2018. American Fisheries Society. 1090 pages, 98.00 USD, Cloth. Paddlefish: Ecological, Aquacultural, and Regu- latory Challenges of Managing a Global Resource. Edited by Jason D. Schooley and Dennis L. Scarnec- chia. 2019. American Fisheries Society. 290 pages, 79.00 USD, Paper. Trout and Char of the World. Edited by Jeffrey L. Kershner, Jack E. Williams, Robert E. Gresswell, and Javier Lobon-Cervia. 2019. American Fisheries Society. 800 pages, 79.00 USD, Paper. ORNITHOLOGY Kingfisher. By Ildiko Szabo. 2019. Reaktion Books. 208 pages and 90 colour plates, 19.95 USD, Paper. Owls of the World: A Photographic Guide. Second Edition. By Heimo Mikkola. 2019. Firefly Books. 528 pages, 49.95 CAD, Flexibound Cloth, 44.95 CAD, Flexibound Paper. Red Coats and Wild Birds: How Military Orni- thologists and Migrant Birds Shaped Empire. Flows, Migrations, and Exchanges Series. By Kirsten A. Greer. 2020. University of North Carolina Press. 190 pages, 90.00 USD, Cloth, 29.95 USD, Paper, 22.99 USD, E-book. A Season on the Wind: Inside the World of Spring Migration. By Kenn Kaufman. 2019. Houghton 292 Mifflin Harcourt. 288 pages, 26.00 USD, Cloth, 14.99 USD, E-book. White Feathers: The Nesting Lives of Tree Swal- lows. 2020. By Bernd Heinrich. Houghton Mifflin Harcourt. 256 pages, 27.00 USD, Cloth. ZOOLOGY Bat Surveys for Professional Ecologists: Good Practice Guidelines. Third Edition. Edited by Jan Collins. Foreword by Julia Hanmer and Kit Stoner. 2016. Bat Conservation Trust. 103 pages, Paper. Non- printable PDF available at https://cdn.bats.org.uk/ pdf/Resources/Bat_Survey_Guidelines_2016_ NON_ PRINTABLE. pdf? mtime=20181115113931. Black Bears: A Natural History. Second Edition. By Dave Taylor. 2020. Fitzhenry and Whiteside. 288 pages, 40.00 USD, Paper. Biodemography: An Introduction to Concepts and Methods. By James R. Carey and Deborah A. Roach. Foreword by James W. Vaupel. 2020. PUP. 480 pages, 60.00 USD, Cloth. Biology and Conservation of Musteloids. Edited by David W. Macdonald, Chris Newman, and Lauren A. Harrington. 2018. Oxford University Press. 672 pages, 137.50 CAD, Cloth, 115.95 CAD, Paper. The Champions of Camouflage. By Jean-Philippe Noél. 2019. Firefly Books. 160 pages and 110 colour photographs, 35.00 CAD, Cloth. Extinction and Evolution: What Fossils Reveal about the History of Life. By Niles Eldredge. 2019. Firefly Books. 256 pages and 160 colour plates, 29.95 CAD, Paper. Originally published in 2014. Guide to Venomous and Medically Important Invertebrates. By David E. Bowles, James A. Swaby, and Harold J. Harlan. 2018. CSIRO Publishing. 237 pages, 59.99 AUD, Paper. Handbook of Whales, Dolphins and Porpoises of the World. By Mark Carwardine. 2020. Princeton University Press. 528 pages, 1000 colour illustra- tions, and 90 maps, 35.00 CAD, Paper. Narwhal: Revealing an Arctic Legend. Edited by William W. Fitzhugh and Martin T. Nweeia. 2017. IPI Press and Arctic Studies Center, National Museum of Natural History (Smithsonian Institution). 261 pages, 30.00 USD, Paper. This book is the companion to the exhibit “Narwhal: Revealing an Arctic Legend” at the Smithsonian Institution’s National Museum of Natural History. THE CANADIAN FIELD-NATURALIST Vol. 133 *Voices of Marine Mammals: William E. Schevill and William A. Watkins: Pioneers in Bioacoustics. 2019. New Bedford Whaling Museum. 125 pages and flexi-disc insert of audio recordings, 29.99 USD, Paper. OTHER All Things Harmless, Useful, and Ornamental: Environmental Transformation through Species Acclimatization, from Colonial Australia to the World. By Pete Minard. 2019. University of North Carolina Press. 208 pages, 90.00 USD, Cloth, 32.95 USD, Paper, 25.99 USD, E-book. Animal Welfare in a Changing World. Edited by Andrew Butterworth. 2018. CABI. 320 pages, 140.00 USD, Cloth, 75.00 USD, Paper. Being a Scientist: Tools for Science Students. By Michael H. Schmidt. 2019. University of Toronto Press. 320 pages, 63.75 CAD, Cloth, 36.95 CAD, Paper, 26.95 CAD, E-book. That Wild Country: An Epic Journey through the Past, Present, and Future of America’s Public Lands. By Mark Kenyon. 2019. Little A [Amazon Publishing imprint]. 300 pages, 31.12 CAD, Cloth, 14.42 CAD, Paper. Extinction Studies: Stories of Time, Death, and Generations. Edited by Deborah Bird Rose, Thom van Dooren, and Matthew Chrulew. Foreword by Cary Wolfe. 2017. Columbia University Press. 256 pages, 90.00 USD, Cloth, 30.00 USD, Paper or E-book. The Fruitful City: The Enduring Power of the Ur- ban Food Forest. By Helena Moncrieff. 2018. ECW Press. 224 pages, 22.95 CAD, Paper, 16.99 CAD, E-book. Geology of New Brunswick and Prince Edward Island Field Guide. 2019. Boulder Books. 300 pages, 34.95 CAD, Paper. Modern Plant Hunters: Adventures in Pursuit of Extraordinary Plants. By Sandy Primrose. 2020. Pimpernel Press. 272 pages, 30.00 GBP, Cloth. North America’s Galapagos: The Historic Channel Islands Biological Survey. By Corinne Heyning Laverty. 2019. University of Utah Press. 384 pages, 29.95 USD, Paper, 24.00 USD, E-book. Overrun: Dispatches from the Asian Carp Crisis. By Andrew Reeves. 2019. ECW Press. 384 pages, 22.95 CAD, Paper, 16.99 CAD, E-book. Phylogenetic Diversity: Applications and Chal- lenges in Biodiversity Science. Edited by Rosa A. 2019 Scherson and Daniel P. Faith. 2018. Springer Inter- national Publishing. 224 pages, 159.99 USD, Cloth or Paper, 119.00 USD, E-book. Sampling Theory for the Ecological and Natural Resource Sciences. By David G. Hankin, Michael S. Mohr, and Kenneth B. Newman. 2019. Oxford University Press. 368 pages, 105.00 CAD, Cloth, 55.00 CAD, Paper. Also available as an E-book. Smitten by Giraffe: My Life as a Citizen Scientist. Footprints Series, No. 22. By Anne Innis Dagg. 2016. McGill-Queen’s University Press. 256 pages, 34.95 CAD, Cloth. Also available as an E-book. Swamp: Nature and Culture. By Anthony Wilson. 2017. Reaktion Books. 248 pages, 14.95 GBP, Paper. NEw TITLES 293 Synergistic Selection: How Cooperation Has Shaped Evolution and the Rise of Humankind. By Peter Corning. 2018. World Scientific. 304 pages, 78.00 USD, Cloth, 29.95 USD, Paper, 19.95 USD, E-book. The Theory of Evolution: Principles, Concepts, and Assumptions. Edited by Samuel M. Scheiner and David P. Mindell. 2020. UCP. 464 pages, 120.00 USD, Cloth, 45.00 USD, Paper. Also available as an E-book. Why Study Biology by the Sea? Edited by Karl S. Matlin, Jane Maienschein, and Rachel A. Ankeny. 2019. University of Chicago Press. 344 pages, 135.00 USD, Cloth, 45.00 USD, Paper or E-book. The Canadian Field-Naturalist News and Comment Upcoming Meetings and Workshops Alberta Chapter of The Wildlife Society Conference The Alberta Chapter of The Wildlife Society Con- ference to be held 13-15 March 2020 at The Norse- men Inn, Camrose, Alberta. The theme of the con- ference is: “Species on the Move’. Registration is currently open. More information is available at https://www.actws.ca/conference/. Entomological Society of America, Joint North Central & Southwestern Branch Meeting The 2020 Joint North Central & Southwestern Branch Meeting of the Entomological Society of America to be held 15—18 March 2020 at The Skirvin Hilton Oklahoma City, Oklahoma City, Oklahoma. Eastern Bird Banding Association Meeting The 97th meeting of the Eastern Bird Banding Asso- ciation to be held 27—29 March 2020 at the Hilton Garden Inn Mystic/Groton, Groton, Connecticut. The theme of the conference is: ‘Using Bird Science Registration is currently open. More information is available at https://www.entsoc.org/2020-joint- northcentral-southwestern-branch-meeting. to Inform Conservation’. Registration is currently open. More information is available at https://ebba 2020.org/, Entomological Society of America, 2020 Joint Eastern & Southeastern Branch Meeting The 2020 Joint Eastern & Southeastern Branch Meeting of the Entomological Society of America to be held 29 March—1 April 2020 at the Sheraton Atlanta Hotel, Atlanta, Georgia. Registration is cur- rently open. More information is available at https:// www.entsoc.org/2020-joint-eastern-southeastern- branch-meeting. American Fisheries Society, Western Division and Washington-British Columbia Chapters Annual Meeting The annual meeting of the Western Division and Washington-British Columbia Chapters of the Amer- ican Fisheries Society to be held 13-17 April 2020 at the Pinnacle Harbourfront Hotel, Vancouver, Bri- tish Columbia. The theme of the conference is: Northeast Natural History Conference The 20th Northeast Natural History Conference to be held 17-19 April 2020 at the Hilton Stamford Hotel, Stamford, Connecticut. Registration is cur- Northeast Fish & Wildlife Conference The 76th annual Northeast Fish & Wildlife Con- ference, hosted by the New Jersey Division of Fish and Wildlife, to be held 19-21 April 2020 at the Ocean Place Resort, Long Branch, New Jersey. ‘Crossing Boundaries and Navigating Intersections’. Registration is currently open. More informa- tion is available at https://wa-bc.fisheries.org/2020- meeting/. rently open. More information is available at https:// www.eaglehill.us/NENHC_2020/NENHC2020. shtml. The theme of the conference is: ‘The Power of Partnerships’. Registration is currently open. More information is available at http://www.neafwa.org/ conference.html. Biodiversity Without Boundaries 2020 (NatureServe) Biodiversity Without Boundaries 2020 to be held 19— 22 April 2020 at the Richmond Marriott, Richmond, Virginia. Registration is currently open. More in- formation is available at https://www.natureserve. org/news-events/events/biodiversity-without- boundaries-2020. 294 2019 NEWS AND COMMENT 295 Entomological Society of America, Pacific Branch Meeting The 104th annual meeting of the Pacific Branch of the Entomological Society of America to be held 19— 22 April 2020 at The Centennial Hotel, Spokane, Washington. Registration is currently open. More information is available at https://www.entsoc.org/ pacific/2020-branch-meeting. Wild Canis spp. of North America: a pictorial representation There has been considerable discussion of hy- bridization in the genus Canis in North America with the general consensus that the western Coyote (Canis latrans), Eastern Timber Wolf (Canis lycaon), and Gray Wolf (Canis [upus) hybridized to produce the Eastern Coyote/Coywolf (Canis latrans var. or Canis latrans x lycaon) and Great Lakes Wolf (Canis lupus x lycaon) in eastern North America (Rutledge et al. 2012, 2015; Way 2013; Way and Lynn 2016; Heppenheimer ef a/. 2018). Way (2013) described the five types of wild canids (Canis spp.; foxes excluded) in North America and noted that these canid groups were useful even with the few studies that claim that the Eastern Timber Wolf is not a distinct species but rather a hybrid between western Coyotes and Gray Wolves (von Holdt et a/. 2011, 2016), despite the lack of field evidence that these two species mate and pro- duce viable offspring (e.g., see Mech et al. 2014). A comprehensive review of the taxonomy of wolves in North America supports the Eastern Timber Wolf as a distinct taxon (Chambers ef al. 2012) as has most of the research on canids in eastern North America (see references in Rutledge ef a/. 2015, but see vonHoldt 2011 countering this). With this “Canis soup” of different, but closely related, species (there is gene flow from C. /ycaon to C. Jupus and from C. lycaon to C. latrans [Way 2013; Heppenheimer et al. 2018]), distinct species status for any canid com- plicates conservation efforts (including C. /upus in eastern North America). Furthermore, the degree of hybridization and terminology associated with these hybrids can be confusing for the layperson, for exam- ple, Way and Lynn’s (2016) use of the term Coywolf Ws Red Wolf Canis rufus Eastern Coyote/Coywolf Canis latrans x lycaon Western Coyote Canis latrans Eastern Timber Wolf Canis lycaon versus Wheeldon and Patterson’s (2017) use of the term Eastern Coyote. Accordingly, we created a pictorial representation of Canis spp. in North America showing the six main types of canids: western Coyotes, Eastern Coyotes/ Coywolves, Red Wolves, Eastern Timber Wolves, Great Lakes Wolves, and Gray Wolves (Figure 1). Because of the frequent separation of Red Wolf (Canis rufus), Eastern Timber Wolf, and Gray Wolf in analyses (e.g., von Holdt et al. 2011; Chambers et al. 2012) we show these canids separately even though others believe Red Wolf and Eastern Timber Wolf are the same species at opposite ends of their range (Wilson et a/. 2000). This drawing represents average body sizes of one canid compared to another; however, it is important to realize the limitations of these average depictions. Even within a given type, males and females differ in size and there is consider- able variation—where the size of one might be sim- ilar or even larger than the one adjacent. They may be difficult to tell apart in the field, not only from a distance, but even when captured, especially where their ranges overlap (e.g., in and around Algonquin Provincial Park, Ontario). This is further exempli- fied by Newsome ef al. (2015) noting that even larger western Gray Wolves and smaller western Coyotes (which share no size overlap; Figure 1) are often diffi- cult to tell apart from a distance and someone ‘shoot- ing a coyote can sometimes result in a dead wolf’. Natural expansion or recolonization of a range is a confounding factor (e.g., Eastern Timber Wolves or Great Lakes Wolves dispersing into southern Canada and the northeastern USA are just claimed to be \ \ ——) Great Lakes Wolf Canis lupus x lycaon ae al Gray Wolf Canis lupus FiGurE 1. Wild Canis of North America. These drawings are intended to represent average body sizes of one canid com- pared to another. But within a given type, males and females differ in size and there 1s considerable variation such that the size of one might be similar or even larger than the one adjacent making them difficult to tell apart in the field, especially where ranges overlap. Also, while Red and Eastern Timber Wolf are considered separate here, many studies have indi- cated that they are possibly the same species (Canis lycaon) living on opposite ends of their eastern North American range. Drawings: J.L. Hirten. 296 heavy Eastern Coyotes). Often genetic testing is the only way to differentiate among Canis spp. in eastern North America (Rutledge et al. 2012). Recent research acknowledges the importance of hybridization among closely related species and in the case of eastern wolves there is a need for man- aged introgression that focusses on preserving any eastern wolf genetic material in any genome regard- less of their potential mosaic ancestry composition (Heppenheimer ef a/. 2018). If such an effort priori- tizes and maintains individuals that carry admixed genomes, as Heppenheimer ef al. (2018) suggest, then more common animals like the Eastern Coyote would be an important source of greater genetic variation and potential adaptive capacity. It is our hope that this diagram (Figure 1) is a useful guide to show the vari- ation and types of Canis spp. in North America with a Specific focus in eastern North America. Literature Cited Chambers, S.M., S.R. Fain, B. Fazio, and M. Amaral. 2012. An account of the taxonomy of North American wolves from morphological and genetic analyses. North American Fauna 77: 1-67. https://doi.org/10.3996/nafa. 77.0001 Heppenheimer, E., R.J. Harrigan, L.Y. Rutledge, K.-P. Koepfli, A.L. DeCandia, K.E. Brzeski, J.F. Benson, T. Wheeldon, B.R. Patterson, R. Kays, P.A. Hohenlohe, and B.M. von Holdt. 2018. Population genomic analy- sis of North American eastern wolves (Canis lycaon) supports their conservation priority status. Genes 9: 606: 1-18. https://doi.org/10.3390/genes9120606 Mech, L.D., B.W. Christensen, C.S. Asa, M. Callahan, and J.K. Young. 2014. Production of hybrids between western gray wolves and western coyote. PLoS ONE 9(2): e88861. https://doi.org/10.1371/journal.pone.0088861 Newsome, T.M., J.T. Bruskotter, and W.J. Ripple. 2015. When shooting a coyote kills a wolf: mistaken identity or misguided management? Biodiversity and Conservation 24: 3145-3149. Rutledge, L.Y., S. Devillard, J.Q. Boone, P.A. Hohenlohe, and B.N. White. 2015. RAD sequencing and genomic simulations resolve hybrid origins within North American Canis. Biology Letters 11: 20150303. https:// doi.org/10.1098/rsbl.2015.0303 Rutledge, L.Y., P.J. Wilson, C.F.C. Klutsch, B.R. Pat- terson, and B.N. White. 2012. Conservation genom- THE CANADIAN FIELD-NATURALIST Vol. 133 ics in perspective: a holistic approach to understand- ing Canis evolution in North America. Biological Conservation 155: 186-192. https://doi.org/10.1016/).bio con.2012.05.017 von Holdt, B.M., J.A. Cahill, Z. Fan, I. Gronau, J. Robinson, J.P. Pollinger, B. Shapiro, B.J. Wall, and R.K. Wayne. 2016. Whole-genome sequence analy- sis shows that two endemic species of North American wolf are admixtures of the coyote and gray wolf. Sci- ence Advances 2: e1501714. https://doi.org/10.1126/sci adv.1501714 von Holdt, B.M., J.P. Pollinger, D.A. Earl, J.C. Knowles, A.R. Boyko, H. Parker, E. Geffen, M. Pilot, W. Je- drzejewski, B. Jedrzejewska, V. Sidorovich, C. Gre- co, E. Randi, M. Musiani, R. Kays, C.D. Bustamante, E.A. Ostrander, J. Novembre, and R.K. Wayne. 2011. A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids. Genome Research 21: 1294-1305. https://doi.org/10.1101/gr.1163 01.110 Way, J.G. 2013. Taxonomic implications of morpholo- gical and genetic differences in Northeastern Coyotes (Coywolves) (Canis latrans x C. lycaon), Western Coyotes (C. latrans), and Eastern Wolves (C. lycaon or C. lupus lycaon). Canadian Field-Naturalist 127: 1-16. https://doi. org/10.22621/cfn.v12711.1400 Way, J.G., and W.S. Lynn. 2016. Northeastern coyote/coy- wolf taxonomy and admixture: a meta-analysis. Canid Biology and Conservation 19: 1-7. Accessed 11 October 2019. http://canids.org/CBC/19/Northeastern_coyote_ taxonomy. pdf. Wheeldon, T.J., and B.R. Patterson. 2017. Comment on “northeastern coyote/coywolf taxonomy”. Canid Biology and Conservation 20: 14-15. Accessed 11 October 2019. https://canids.org/CBC/20/Comment%20 _on_Way_and_Lynn_2016.pdf. Wilson, P.J., S. Grewal, I.D. Lawford, J.N.M. Heal, A.G. Granacki, D. Pennock, J.B. Theberge, M.T. Theberge, D.R. Voigt, W. Waddell, R.E. Chambers, P.C. Paquet, G. Goulet, D. Cluff, and B.N. White. 2000. DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf. Canadian Journal of Zoology 78: 2156-2166. https://doi.org/10.1139/z 00-158 JONATHAN G. Way Eastern Coyote Research, Osterville, MA, USA JUSTINE LEE HIRTEN Fairfield, CT, USA 2019 NEWS AND COMMENT 297 In Memoriam: Francis Cook (3 March 1935-3 January 2020) Francis Cook was the longest serving editor of The Canadian Field-Naturalist. He was editor of the journal for 34 years, from 1962 to 1966 and from 1981 to 2010. In total, Francis edited 35 volumes of The Canadian Field-Naturalist. He helped hundreds of researchers publish their work in the journal. In addition to his work on The Canadian Field- Naturalist, Francis Cook was the Curator of Herpeto- logy at what is now the Canadian Museum of Nature from 1960 to 1993, aside from a two-year educational leave to work on a Ph.D. at the University of Manitoba. Francis had a passion for herpetofauna that lasted a lifetime. He spent decades gathering data on the nat- ural history of local amphibians near his home. In 2018, Francis was awarded the Order of Canada for his research on amphibians and reptiles and for being the long-time editor of The Canadian Field- Naturalist. He was also honoured by the Ottawa Field-Naturalists’ Club. He was selected as Member of Year in 1990 and 2010 for his efforts editing The Canadian Field-Naturalist, and he was made an Honorary Member of the Club in 1998 “For service to the Club and herpetological work”. Francis’s ex- ceptional contributions to our understanding of the natural history of amphibians and reptiles (detailed in Halliday and Seburn 2018; Seburn and Halliday 2018) were honoured in special issues (volume 132, issues 1 and 2) of The Canadian Field-Naturalist, with the content of those issues dedicated to studies on Canadian amphibians and reptiles. Francis Cook died in Kemptville on 3 January 2020. Memorial donations may be made to The Can- adian Field-Naturalist if desired; you may direct an e-transfer to treasurer@ofnc.ca with a note “Re: The Canadian Field Naturalist in Memory of Francis Cook”. Literature Cited Halliday, W.D., and D.C. Seburn. 2018. Introduction to the Special Issue on herpetology in Canada. Canadian Field-Naturalist 132: 1-3. https://doi.org/10.22621/cfn.v 13211.2113 Seburn, D.C., and W.D. Halliday. 2018. The publications of Francis Cook. Canadian Field-Naturalist 132: 99-102. https://do1.org/10.22621/cfn.v132i2.2169 OFNC PUBLICATIONS COMMITTEE The Canadian Field-Naturalist Editors’ Report for Volume 132 (2018) Mailing dates for the four issues in volume 132 were: 24 October 2018, 31 January 2019, 2 May 2019, and 31 July 2019. Summaries of the distribution of paid subscribers to The Canadian Field-Naturalist for 2018 are provided in Table 1, along with com- parison numbers for volume 131. This list does not include free copies distributed to Honorary Ottawa Field Naturalists’ Club (OFNC) members or online access, which is included in OFNC membership dues. Institutional subscribers potentially represent many thousands of users. The number of articles published in volume 132 increased by 10 over the number pub- lished in volume 131 while the number of notes de- creased by 10, with the same number of manuscripts published both years (Table 2). Not surprisingly, 25/45 (56%) of the manuscripts in 132 were on am- phibians and reptiles, given the first two issues of 132 were Special Issues: studies on Canadian amphibi- ans and reptiles in honour of Dr. Francis Cook. The three manuscripts in the “other” category were on al- vars, Arctic slumps, and fungi (Table 2). The number of book reviews and new titles published in volume 132 were slightly up and down, respectively, over the numbers in volume 131 (Table 3). The total number of pages published increased by 36 for volume 132 over volume 131 (Table 4) with articles contributing 69% to the page count and 76% of manuscripts published (Table 2). There were no thematic collections (editor- selected compilations of previously published contri- butions in both The Canadian Field-Naturalist and the regional OFNC publication, Trail & Landscape, on a central theme with internet links to each article) nor articles on Greatest Canadian Field Naturalists, the latter of which were included in News and Com- ment in 131. TABLE 2. Number of research articles and notes published in The Canadian Field-Naturalist, Volume 132 (Volume 131), by major field of study. Subject Articles Notes Total Mammals 3 (3) 0 (8) 3 (11) Birds 3 (10) 2 (6) 5 (16) Amphibians and Reptiles 19 (3) 6 (2) 25 (5) Fishes 0 (1) 1 (2) 1 (3) Plants 4 (3) 1 (2) 5 (5) Insects 1(1) 0 (0) 1 (1) Non-insect invertebrates 1 (2) 1(1) 2 (3) Other 3 (1) 0 (0) 3 (1) Total 34(24) 11 (21) 45(45) TABLE 3. Number of reviews and new titles published in the Book Review section of The Canadian Field-Naturalist, Volume 132 (Volume 131), by topic. Reviews New Titles Zoology 26 (15) 148 (155) Botany 7 (7) 24 (43) Miscellaneous 9 (18) 122 (111) Total 42 (40) 294 (309) Sixty-five manuscripts were submitted to The Canadian Field-Naturalist in 2018, eight more than in 2017; there were also two initial enquiries about suit- ability of topics for submission, one of which submit- ted formally in 2019. All except one manuscript was submitted using the Online Journal System, some (n = 9) after an initial email submission. Thirteen of the 65 were for the Special Issues on Canadian amphib- ians and reptiles. Only 11 of the 65 submitted manu- scripts were not accepted for publication upon initial submission or review or were insufficiently revised to warrant publication. The remainder, 83.1%, were ac- cepted or are undergoing revision and review. In 2017, TABLE 1. The 2018 (2017) circulation of The Canadian Field-Naturalist. Compiled by Eleanor Zurbrigg from the subscrip- tion list for 132(4). This list does not include copies distributed to Honorary Members or online access which is included in OFNC membership fees. Subscriber Type Canada USA Other Total OFNC Members 43 (51) 1 (4) 0 (0) 44 (55) Subscriptions: Individual 26 (26) 7 (7) 0 (0) 33 (33) Institutional 66 (73) 90 (106) 12 (12) 168 (191) Total 135 (150) 98 (117) 12 (12) 245 (279) 298 ©The Ottawa Field-Naturalists’ Club 2019 EpDITORS’ REPORT TABLE 4. Number of pages per section published in The Canadian Field-Naturalist, volume 132 (131), by issue. 1 2 Editorials/Editors’ Report* 3 (0) 4 (0) Articles 51(67) 86 (47) Notes 12 (7) 2 (17) Thematic Collections 0 (5) 0 (8) Tributes 0 (0) 0 (0) Book Reviewst 18 (14) 18 (15) News and Comment 2 (2) 2 (1) Reportst 12 (19) 0 (0) Erratum 0 (0) 0 (0) Index —() -C©) Total 98 (114) 112 (88) *Includes introductions to Special Issue Parts I and I. tIncludes reviews and new titles. 299 Issue 5 ; Total 3 (2) 0 (1) 10 (3) 74 (47) 94 (42) 305 (203) 11 (30) 12-27) 37 (81) 0 (0) 0 (0) 0 (13) 0 (0) 0 (0) 0 (0) 18 (15) 8 (13) 62 (57) 2 (6) 2 (12) 8 (21) 0 (0) 0 (0) 12 (19) 0 (0) 0 (0) 0 (0) =e) 8 (9) 8 (9) 108 (100) 124 (104) 442 (406) {Includes Annual Business Meeting Minutes, Annual Committee Reports, and Awards, including the James Fletcher Award for best paper published in the volume; Financial Statements are only available online beginning with 132. 89.5% of the 57 submissions were accepted for publi- cation and either published or underwent further re- vision and review. Guest Editors William Halliday and Dave Se- burn received the manuscripts submitted for the Spe- cial Issues, assigned reviewers, handled the re- view process, and passed the accepted manuscripts to Dwayne Lepitzki, Editor-in-Chief, and Amanda Martin, the Assistant Editor, for the rest of the pub- lication process. Amanda edited content, proofread galleys, and sent and received author order and trans- fer of copyright forms; she also prepared the News and Comment. Sandra Garland and John Wilmshurst proof-read and copy edited manuscripts. Wendy Cotie typeset galleys, provided corrections for page proofs, and created pdfs. Barry Cottam, Book Review Editor, requested books for review, selected reviewers, ed- ited submitted reviews, and prepared the new titles listings. Ken Young sent page charge invoices to au- thors and tracked the budget while Eleanor Zurbrigg managing subscriptions and mailed printed cop- ies. William Halliday, Online Journal Manager and Webmaster, provided digital content to subscribers, posted tables of contents, abstracts, and pdfs on The Canadian Field-Naturalist website, and prepared the Index. Our Associate Editors managed manuscripts, provided reviews and recommendations, and guided authors through the revisions process. Dave Seburn, our Map Editor, reviewed and provided suggestions for all the maps. The Publication Committee, chaired by Jeff Saarela and consisting of Annie Bélair, Dan Brunton, Carolyn Callaghan, Paul Catling, Barry Cottam, William Halliday, Diane Kitching, Dwayne Lepitzki, Amanda Martin, Karen McLachlan Hamil- ton, Dave Seburn, Ken Young, and Eleanor Zurbrigg effectively guided the operation of the journal. We are indebted to our very dedicated team. The following Associate Editors managed, as- sessed, and reviewed manuscripts published in vol- ume 132: R. Brooks, University of Guelph, emeritus, Guelph ON (2 manuscripts); P.M. Catling, Agriculture and Agri-Food Canada, retired, Ottawa ON (6); F. Chapleau, University of Ottawa, Ottawa ON (1); J. Foote, Algoma University, Sault St. Marie ON (4); W. Halliday, University of Victoria, Victoria BC (18); D. Lepitzki, Banff AB (1); D.F. McAlpine, New Brunswick Museum, Saint John NB (1); J. McCracken, Bird Studies Canada, Port Rowan ON (1); G. Mowat, Government of British Columbia, Nelson BC (1); DW. Nagorsen, Mammalia Biological Consulting, Victoria BC (2); J.M. Saarela, Canadian Museum of Nature, Ottawa ON (1); D. Seburn, Canadian Wildlife Federation, Ottawa ON (6); J. Skevington, Agriculture and Agri-Food Canada, Ottawa ON (1). As with many other journals, Associate Editors are at times having difficulty finding suitable review- ers; without dedicated Associate Editors and review- ers there would be no journal. As such, a heart-felt thanks and gratitude is extended to the following who reviewed manuscripts published in volume 132 (number of manuscripts reviewed >1 in parentheses): Carl Anthony, John Carroll University; Andréanne Beardsell, Université du Québec a Rimouski; Chri- stine Bishop, Environment and Climate Change Canada; Sean Blaney, Atlantic Canada Conservation Data Centre (2); Ernie Brodo, Research Associate, Canadian Museum of Nature; Ron Brooks, University of Guelph (2); Dan Brunton, Brunton Consulting (2); Jacob Burkhart, University of Missouri; William Busby, Kansas Biological Survey; Rob Cannings, Royal British Columbia Museum; Pauline Catling, North-South Environmental Inc.; Tony Chubbs, De- partment of National Defence; Stephen Clayton, New Brunswick Museum; Justin Congdon, University of 300 Georgia; Joe Crowley, Ontario Ministry of Natural Resources and Forestry; David Cundall, Lehigh Uni- versity; Christina Davy, Ontario Ministry of Natur- al Resources and Forestry; Kendra Driscoll, New Brunswick Museum; Marco Festa-Bianchet, Univer- sity of Sherbrooke (2); Neil Ford, University of Texas at Tyler; Robert Forsyth, Kamloops BC; Roseanna Gamlen-Greene, University of British Co- lumbia; Scott Gillingwater, Upper Thames River Conservation Authority (2); Peter Gogan, US Geo- logical Survey; Patrick Gregory, University of Vic- toria (2); Gareth Griffith, Aberystwyth University; Samuel Hache, Environment and Climate Change Canada; Gavin Hanke, Royal British Columbia Museum; Allan Harris, Northern Bioscience Eco- logical Consulting; Virgil Hawkes, LGL Ltd.; Tim Haxton, Ontario Ministry of Natural Resources and Forestry (2); Stephen Hecnar, Lakehead University (2), Eric Hellquist, New York Botanical Garden; Tom Herman, Acadia University (2); Bob Inman, Montana Fish, Wildlife and Parks; Gregory Jongsma, New Brunswick Museum; Karl Larsen, Thompson Rivers University (2); Jackie Litzgus, Laurentian University; Eric Lofroth, BC Conservation Data Centre, retired; Teresa Lorenz, US Department of Agriculture Forest Service; Stephen MacFarlane, University of Toronto; John Maunder, The Rooms Provincial Museum; David McCorquondale, Cape Breton University; Liam McGuire, Texas Tech Uni- versity; David Mifsud, Herpetological Resource Management; Joseph Mitchell, Florida Museum of Natural History; Steve Mockford, Acadia University; Patrick Moldowan, University of Toronto (2); Mason Murphy, Miami University; Jeff Nekola, University of New Mexico; Annegret Nicolai, Université de Rennes; Michael Oldham, Ontario Natural Heritage Information Centre (2); Martin Ouellet, Amphibia- THE CANADIAN FIELD-NATURALIST Vol. 133 Nature; Brittany Ousterhout, National Great Rivers Research and Education Center; Kristiina Ovaska, Biolinx Environmental Research Ltd.; Steve Paiero, University of Guelph; James Paterson, University of Ottawa (2); Cynthia Paszkowski, University of Alber- ta; Ray Poulin, Royal Saskatchewan Museum; George Powell, University of Calgary; Tanya Pulfer, Ontario Nature; Jennie Rausch, Environment and Climate Change Canada; Don Reid, Wildlife Conservation Society Canada; Scott Redhead, Agriculture and Agri-Food Canada; Matt Reudink, Thompson Rivers University; Anton Reznicek, University of Michigan; Tony Roberts, US Fish and Wildlife Service; Pamela Rutherford, Brandon University; Taza Schaming, Cornell University; Fred Schueler, Fragile Inheritance; Cory Sheffield, Royal Saskatchewan Museum; Brian Slough, Whitehorse YT; Brian Smith, Black Hills State University; Chris Somers, University of Regina; Duane Stevenson, National Oceanic Atmospheric and Administration, National Marine Fisheries Service; David Swanson, University of South Dakota; Carl Taylor, American Museum of Natural History. The journal was printed at Gilmore Printers, Ottawa. Thanks to Guylaine Duval of Gilmore Printers for overseeing production and printing. We are grate- ful to The Ottawa Field-Naturalists’ Club President Diane Lepage and the club’s Board of Directors for their support of the journal. We are also grateful to all of the individual subscribers and authors who support our team as we strive to provide a high-quality scien- tific journal on natural history, field biology, and ecol- ogy. Finally, we thank our families/partners for being patient and supportive throughout many long days, evenings, and weekends of working on the journal. Dwayne LepitzKx1, Editor-in-Chief AMANDA ManrTIN, Assistant Editor TABLE OF CONTENTS (concluded) Volume 133, Number 3 Tributes A Tribute to Rudolph Frank Stocek, 1937-2018 | DONALD F. MCALPINE and GRAHAM J. FORBES Book Reviews BoTANY: Seaweed Chronicles ORNITHOLOGY: Birds of Saskatchewan—Feed the Birds: Attract and Identify 196 Common North Ameri- can Birds Zoo.LoGy: The North Atlantic Right Whale: Disappearing Giants. Revised and Updated Edition—Yellow- stone Cougars: Ecology Before and During Wolf Restoration OTHER: How to Give Up Plastic: A Guide to Changing the World, One Plastic Bottle at a Time—Plastic Soup: An Atlas of Ocean Pollution—Mama’s Last Hug: Animal and Human Emotions—North Pole: Nature and Culture NEw TITLES News and Comment Upcoming Meetings and Workshops Alberta Chapter of The Wildlife Society Conference—Entomological Society of America, Joint North Central & Southwestern Branch Meeting—Eastern Bird Banding Association Meeting—Entomological Society of America, 2020 Joint Eastern & Southeastern Branch Meeting—American Fisheries Society, Western Division and Washington-British Columbia Chapters Annual Meeting—Northeast Natural History Conference—Northeast Fish & Wildlife Conference—Biodiversity Without Boundaries 2020 (NatureServe)—Entomological Society of America, Pacific Branch Meeting Wild Canis spp. of North America: a pictorial representation In Memoriam: Francis Cook (3 March 1935-3 January 2020) Editors’ Report Mailing date of the previous issue 133(2): 5 December 2019 2019 270 276 2 281 284 289 294 29> 297 298 THE CANADIAN FIELD-NATURALIST Volume 133, Number 3 First recorded co-occurrence of Valvata lewisi Currier, 1868 and Valvata lewisi ontariensis Baker, 1931 (Gastropoda: Valvatidae) from Alberta, Canada, with notes on morphometric and genetic variability ROBERT P. HINCHLIFFE, CHERYL TEBBY, and TYLER P. COBB Spotless burnsi pattern in Northern Leopard Frog (Lithobates pipiens) in Maine Scott B. LINDEMANN, Davip E. PUTNAM, MALCOLM L. HUNTER, JR., and TREVOR B. PERSONS Axanthism in Green Frogs (Lithobates clamitans) and an American Bullfrog (Lithobates cates- beianus) in Maine Scott B. LINDEMANN, AIDAN M. O’BRIEN, TREVOR B. PERSONS, and PHILLIP G. DEMAYNADIER Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt new to Atlantic Canada SEAN R. HAUGHIAN and THOMAS H. NEILY Lichens and allied fungi of Sandbar Lake Provincial Park, Ontario HANNA R. Dorva_ and RICHARD TROY MCMULLIN Do turtle warning signs reduce roadkill? Davip C. SEBURN and HANNAH McCurbDy-ADAMS Surveys for terrestrial gastropods in the Kootenay region of British Columbia, with new records and range extensions KRISTIINA OvASKA, LENNART SOPUCK, and JENNIFER HERON Conspecific cues encourage Barn Swallow (Hirundo rustica erythrogaster) prospecting, but not nesting, at new nesting structures ANDREW J. CAMPOMIZZI, ZOE M. LEBRUN-SOUTHCOTT, and KRISTYN RICHARDSON Seasonal movements of White-tailed Deer (Odocoileus virginianus) in the Rocky Mountains of British Columbia TREVOR A. KINLEY Sharp-tailed Grouse (Zympanuchus phasianellus) population dynamics and restoration of fire- dependent northern mixed-grass prairie ROBERT K. MurpuHy and KAREN A. SMITH Humpback Whale (Megaptera novaeangliae) observations in Laskeek Bay, western Hecate Strait, in spring and early summer, 1990-2018 ANTHONY J. GASTON, NEIL G. PILGRIM, and VIVIAN PATTISON 2019 189 193 196 199 206 216 221 235 246 253 263 (continued inside back cover) ISSN 0008-3550 The CANADIAN FIELD-NATURALIST A JOURNAL OF FIELD BIOLOGY AND ECOLOGY Promoting the study and conservation of northern biodiversity since 1880 Volume 133, Number 4 ¢ October—December 2019 Ottawa Field-Naturalists’ Club Club des naturalistes d’Ottawa The Ottawa Field-Naturalists’ Club FOUNDED 1863 (CURRENT INCORPORATION 1879) Patron Her Excellency the Right Honourable Julie Payette, C.C., C.M.M., C.O.M., C.Q., C.D. Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encour- age Investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. 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Cover: Polyergus bicolor, a slave-making ant, was recently discovered for the first time in Alberta, a substantial range expansion for the species. It was found parasitizing Formica podzolica, a new host for the species. See note in this issue by Christine Sosiak et al., pages 309-312. Photo: C. Sosiak, 21 July 2017, near Sylvan Lake, Alberta. The Canadian Field-Naturalist Note A successfully breeding, partially leucistic American Robin (7urdus migratorus) Nina M. ZITANt’*, LEANNE A. GRIEVES!, and R. GREG THORN! ‘Department of Biology, University of Western Ontario, London, Ontario N6A 5B7 Canada “Corresponding author: nzitani@uwo.ca Zitani, N.M., L.A. Grieves, and R.G. Thorn. 2019. A successfully breeding, partially leucistic American Robin (Turdus migratorius). Canadian Field-Naturalist 133(4): 301-304. https://do1.org/10.22621/cfn.v13314.2141 Abstract American Robin (7urdus migratorius) is the most abundant and broadly distributed thrush in North America. Both sexes likely engage in mate choice, and there is some evidence of assortative mating based on breast colour in this species. Over two breeding seasons, we documented a case of partial leucism, primarily of the breast feathers, in a male American Robin in London, Ontario, Canada. We report evidence that the leucistic robin was capable of successful breeding. How the fit- ness of leucistic versus normal robins compares and how leucism influences mate choice in this and other species remain to be explored. Key words: Ornithology; colouration; leucism; sexual selection; fitness; breeding; American Robin; Turdus migratorius The colouration of birds is a result of light inter- acting with either the nanostructure of the integu- ment or cellular pigments, and sometimes a combi- nation of the two (Prum 2006). In birds, melanin is the most common pigment. A variety of feather and skin colour is attributable to two forms of melanin, eumelanin (grey to black colours) and phaeomelanin (some yellows and reds, and browns by admixture of eumelanin). The other major source of pigments in birds is carotenoids derived from their diet. Melanins are not derived from food but are produced by ani- mals endogenously. Early in embryonic development, neural crest-derived melanoblasts migrate to the skin and the newly forming feathers. The melanoblasts differentiate into melanocytes and begin synthesizing melanin by the end of the first week of development (Bharti et al. 2006; McGraw 2006). A multitude of mutations can cause white feathers where there should be feathers coloured by pigments, and there is much confusion in the literature and among birders about the correct names for such col- our aberrations. We follow van Grouw (2006, 2013), who provided a summary of the most frequently oc- curring colour aberrations and a much-needed guide to standardize their naming. Leucism is defined as the partial or total lack of both melanins in feathers and skin as a result of the heritable failure of melanoblasts to migrate to the proper area of the body. Melanocytes and the resulting colours are absent in those areas, and the feathers appear white. Birds may be partially leu- cistic, with only some white feathers, or totally leucis- tic, with all white feathers. Importantly, melanocytes and eye pigment cells differ in their embryological or- igin and leucistic birds have normally coloured eyes (Bharti et al. 2006; van Grouw 2013). Wild birds with leucism may face a number of challenges; however, evidence of a detrimental effect of leucism is inconclusive. In one study, the mortal- ity of leucistic young was double that of young with normal plumage (Reese 1980). In another, a leucis- tic adult was not accepted into a conspecific group (Corréa et al. 2017). In contrast, a leucistic adult was frequently accompanied by conspecifics in an- other study (Cestari and Vernaschi Vieira da Costa 2007). Several studies report no evidence that leu- cism affects adult breeding performance (Owen and Skimmings 1992; Forrest and Naveen 2000). American Robin (7urdus migratorius) is North America’s largest, most abundant, and widely distrib- uted thrush. Typically, adult male American Robins have deep greyish to dark-brown upper parts, a black- ish head, white crescents above and below the eye, white undertail coverts, and, in most eastern popu- lations, white tips on the outer retrices. The under A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 301 ©The Ottawa Field-Naturalists’ Club 302 parts and breast are a rich rufous colour ( Vanderhoff et al. 2016; Figure la,b). Females appear similar but with a paler grey crown and mantle, more white on the ventrum, and a paler breast (Figure la,b). Adult plumages vary little throughout the year; however, males have darker crowns, less white on the ventrum, and darker breasts in spring compared with autumn (Vanderhoff et a/. 2016). Turdus migratorius (male) =— a Americ Turdus migratorttt (female) THE CANADIAN FIELD-NATURALIST Vol. 133 There is great interest in documenting the behav- iour of leucistic birds to further understand the effects of aberrant plumages and the diverse roles plumage colouration plays in the lives of birds. Here we pre- sent evidence of a partially leucistic male American Robin successfully breeding on a ~0.2-ha private property in northwest London, Ontario (43.00°N, 81.29°W) during the 2016 and 2017 breeding seasons. American Robin Turdus migratorius (male) Turdus migratorius (female) = FiGurE 1. American Robin (7urdus migratorius). a,b. Male (top) and female specimens showing normal plumage colour- ation, collected in Strathroy, Ontario, 1932, Western University Zoological Collections: a. dorso-lateral view; b. ventral view. c—e. Partially leucistic male American Robin, London, Ontario: c. 30 June 2016, d. 27 May 2017, e. 5 June 2017. f. American Robin embryo, below nest site of partially leucistic robin, 18 June 2017. Photos: Nina M. Zitani. 2019 Beginning in late spring 2016, a male American Robin with aberrant white feathers and normal eye colouration was observed multiple times. A photo of the robin was taken on 30 June 2016 (Figure Ic). Later in the summer, the leucistic robin was observed mating with a female of normal plumage and subse- quently feeding a fledgling on a lawn. In 2017, the leu- cistic robin was first observed on 23 April. Over the course of the spring, the robin was observed repeat- edly, and photos were taken on 27 May 2017 (Figure 1d) and 5 June 2017 (Figure le). By early June, the leucistic robin and a mate with normal colouration were observed bringing nest materials into a large, woody climbing hydrangea (Hydrangea sp.). On 18 June 2017, a nearly fully developed embryo was found smashed on a rock below the nest site (Figure If). Throughout the season, several Brown-headed Cowbirds (Molothrus ater) were observed in the area. On 5 August 2017 at 2000, the leucistic robin was ob- served on a lawn 0.25 m from a vocalizing fledgling. Shortly thereafter, the leucistic robin approached and fed the fledgling. On several occasions, the leucistic robin was observed singing normally. The plumage colouration of this leucistic robin was as follows: the typically greyish upperparts of the body were mixed with patches of white, particularly on the mantle and lesser, median, and greater cov- erts. There appeared to be a greater-than-normal pro- portion of white around the eye and throat. The usu- ally rich rufous breast was heavily marked by white feather patches. The eyes of the robin were black. The lack of colouration in typically pigmented areas that we observed in this bird and normally coloured eyes are characteristic of partial leucism (van Grouw 2006, 2013). Because of the characteristic markings of this bird, we were confident in all cases that our ob- servations were of the same individual (Figure 1c—e). The occurrence of leucism in natural populations of wild birds rarely exceeds 1% (Bensch et al. 2000). Gross (1965) reported that American Robin had the highest rate (8.2%) of “albinism” among North American birds he surveyed; his tally included not only leucism but all forms of pale aberrations. When strictly defined, leucism in American Robin has been reported less often than albinism and melanism (Vanderhoff et al. 2016). Plumage colouration has long been associated with sexual selection (Darwin 1871), with females typically preferring brightly coloured males (e.g., Safran et al. 2005), likely because plumage is of- ten condition-dependent (Hamilton and Zuk 1982). Leucistic birds may appear duller or less attractive to prospective mates and, consequently, may have lower reproductive success and overall fitness compared ZITANI ET AL.: LEUCISTIC AMERICAN ROBIN 303 with normally-coloured individuals, especially those with brightly coloured plumage. In species where the sexes share the same traits (e.g., breast colour in American Robin), mutual sex- ual selection can occur if both sexes benefit from dis- criminating among potential mates based on these traits (Rowe and Weatherhead 2011). The partially leucistic male robin we observed had a large propor- tion (~40—50% of breast area) of white feathers on his breast. To our knowledge, there are no data on how leucism might influence mate preference in American Robin; however, because robins apparently exhibit positive assortative mating with respect to breast col- our (Rowe and Weatherhead 2011), we expect this leucistic male would be more likely to mate with a paler female. In conclusion, our report documents a rare case of partial leucism in American Robin, and provides ev- idence that leucistic robins are capable of successful breeding. How the fitness of leucistic versus normal robins compares remains to be explored. Given the mixed results in the literature on the impacts of leu- cism, more studies are needed to understand the main- tenance of leucism in natural populations and the im- pacts of this plumage abnormality on wild birds. Author Contributions Writing — Original Draft: N.Z. and L.G.; Writing — Review & Editing: N.Z., L.G., and R.G.T.; Concep- tualization: N.Z.; Investigation: N.Z and R.GT. Acknowledgements We acknowledge that this work was conducted on the traditional lands of the Anishinaabek, Haudenosaunee, Lunaapéewak, and Attawandaron peoples. The Zoological Collections, Department of Biology, Western University, provided the male and female specimens of American Robin, and the cam- era, a Finepix S1 (Fujifilm, Mississauga, Ontario, Canada), and we thank Elizabeth MacDougall- Shackleton for helpful discussion. Literature Cited Bensch, S., B. Hansson, D. Hasselquist, and B. Nielsen. 2000. Partial albinism in a semi-isolated population of Great Reed Warblers. Hereditas 133: 167-170. https:// doi.org/10.1111/j.1601-5223.2000.t01-1-00167.x Bharti, K., M.T. Nguyen, S. Skuntz, S. Bertuzzi, and H. Arnheiter. 2006. The other pigment cell: specifica- tion and development of the pigmented epithelium of the vertebrate eye. Pigment Cell Research 19: 380-394. https://doi.org/10.1111/j.1600-0749.2006.00318.x Cestari, C., and T. Vernaschi Vieira da Costa. 2007. A case of leucism in Southern Lapwing (Vanellus chilen- sis) in the Pantanal, Brazil. Boletin SAO 17: 145-147. Corréa, L.L.C., N. Horn, C. dos Santos Bruckmann, and MLV. Petry. 2017. Leucism in Vanellus chilensis 304 (Molina, 1872) (Birds: Charadriiformes) in Pampa bi- ome, southern Brazil. Oecologia Australis 21: 219-221. https://doi.org/10.4257/oeco.2017.2102.14 Darwin, C. 1871. The Descent of Man, and Selection in Re- lation to Sex. John Murray, London, United Kingdom. Forrest, S.C., and R. Naveen. 2000. Prevalence of leucism in pygocelid penguins of the Antarctic Peninsula. Water- birds 23: 283-285. Gross, A.O. 1965. The incidence of albinism in North American birds. Bird-Banding 36: 67—71. Hamilton, W.D., and M. Zuk. 1982. Heritable true fitness and bright birds: a role for parasites? Science 218: 384— 387. https://doi.org/10.1126/science.7123238 McGraw, K.J. 2006. Mechanics of melanin-based colora- tion. Pages 243—294 in Bird Coloration Volume 1. Me- chanisms and Measurements. Edited by G.E. Hill and K.J. McGraw. Harvard University Press, Cambridge, Massachusetts, USA. Owen, M., and P. Skimmings. 1992. The occurrence and performance of leucistic Barnacle Geese Branta leuc- opsis. Ibis 134: 22-26. https://doi.org/10.1111/j.1474-919 X.1992.tb07224.x Prum, R.O. 2006. Anatomy, physics, and evolution of struc- tural colors. Pages 295-353 in Bird Coloration Volume 1. Mechanisms and Measurements. Edited by GE. Hill THE CANADIAN FIELD-NATURALIST Vol. 133 and K.J. McGraw. Harvard University Press, Cambridge, Massachusetts, USA. Reese, J.G. 1980. Demography of European mute swans in Chesapeake Bay. Auk 97: 449-464. Rowe, K.M., and P.J. Weatherhead. 2011. Assortative mating in relation to plumage traits shared by male and female American Robins. Condor 113: 881—889. https:// doi.org/10.1525/cond.2011.100207 Safran, R.J., C.R. Neuman, K.J. McGraw, and LT. Lovette. 2005. Dynamic paternity allocation as a func- tion of male plumage color in barn swallows. Science 309: 2210-2212. https://doi.org/10.1126/science.1115090 Vanderhoff, N., P. Pyle, M.A. Patten, R. Sallabanks, and F.C. James. 2016. American Robin (7urdus migrator- ius), Version 2.0. In The Birds of North America. Edited by P.G. Rodewald. Cornell Lab of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.462 van Grouw, H. 2006. Not every white bird is an albino: sense and nonsense about color aberrations in birds. Dutch Birding 28: 79-89. van Grouw, H. 2013. What color is that bird? The cause and recognition of common color aberrations in birds. British Birds 106: 17-29. Received 21 September 2018 Accepted 14 February 2020 The Canadian Field-Naturalist Occurrence of anthropogenic litter in nestling Tree Swallows (Tachycineta bicolor) STEPHANIE WALSH!, JENNIFER HAUGHTON’, LEE BELLAN|, ISABELLE GOSSELIN|, 9 Amy FESTARINI*, DAvip LEE!?, and MARILYNE STUART! 9 9 ‘Canadian Nuclear Laboratories, Chalk River Laboratories, 286 Plant Road, Chalk River, Ontario KOJ 1JO Canada *Algonquin College, 186 Frank Nighbor Street, Pembroke, Ontario K8A 4M5 Canada *University of Waterloo, Waterloo, Ontario N2L 3G1 Canada “Corresponding author: amy.festarini@cnl.ca Walsh, S., J. Haughton, L. Bellan, I. Gosselin, A. Festarini, D. Lee, and M. Stuart. 2019. Occurrence of anthropogenic lit- ter in nestling Tree Swallows (Jachycineta bicolor). Canadian Field-Naturalist 133(4): 305-308. https://doi.org/10. 22621 /cfn.v13314.2221 Abstract While undertaking a study of the effects of strontium-90 on Tree Swallow (Jachycineta bicolor) near Chalk River, Ontario, we noticed the presence of anthropogenic litter (pieces of metal, glass, and plastic, and paper, plastic, and foil wrappers, >I mm in size) in the nestlings. Although combustible litter (pieces of plastic and wrappers) were not quantified before the nestlings were incinerated in 2014 and 2015, gizzards were dissected in 2016. Litter (>1 mm diameter) was found in 30% of the 74 nestlings examined. This material is most likely provided to nestlings, along with food (insects) and natural grit (sand, stones, and mollusc shells), which we also found, by parent birds; however, it could lead to internal injuries and/or harmful substances being absorbed by the young birds. Key words: Tree Swallow; Tachycineta bicolor; nestling; grit; environmental impact; anthropogenic litter; metal; glass; plastic; paper Introduction Insoluble and soluble natural grit (sand, stones, and mollusc shells) is an important component of many avian diets, as it improves the process of grind- ing foods, such as seeds, plant material, and insects, in the gizzard (Barrentine 1980; Best and Gionfriddo 1991; Gionfriddo and Best 1995). In atricial species, grit is provided by parents. The amount and size of grit consumed by a species is believed to depend on the specific diet of the bird (Gionfriddo and Best 1995). Tree Swallow (Jachycineta bicolor), an insec- tivorous species, requires grit for efficient digestion (Mayoh and Zach 1986), and adults have been found to feed grit to nestlings as young as three days of age (Mayoh and Zach 1986). If anthropogenic litter (e.g., pieces of metal, glass, and plastic as well as paper, plastic, and foil wrap- pers) is present near nesting locations, it too could be fed to nestlings. However, ingested anthropogenic lit- ter could lead to internal injury, and/or harmful sub- stances from the materials could be absorbed by the nestlings (Bellrose 1975; Trost 1981; Azzarellow and Van Vleet 1987; Fry et al. 1987; Laist 1987; Cola- buono ef a/. 2010). Herein, we report on the inges- tion of anthropogenic litter by Tree Swallow nestlings near Chalk River, Ontario, Canada. Methods In preparation for a strontium-90 (Sr-90) study described in Lee et al. (2019), nest boxes were in- stalled on the 4000-ha property of the Canadian Nuclear Laboratories’ Chalk River Laboratories (46.052578°N, 77.360890°W; Figure 1) in suitable Tree Swallow nesting habitats (wetland and shore- line; De Steven 1980; Robertson and Rendell 1990). Tree Swallows will readily inhabit nest boxes and tolerate human disturbances, making them an ideal bird for biomonitoring and research (De Steven 1980; Mayoh and Zach 1986; Robertson and Rendell 1990). Monitoring of the nest boxes began in late April or early May of each year, and observations of nesting, egg laying, clutch size, hatchings, nestling growth, and fledging were documented. When nestlings were 12 days old (as determined from known hatch dates), one nestling from each nest box with at least four young was collected (on average a nest box would contain six nestlings). In all, 74 12-day-old nestlings 305 ©The Ottawa Field-Naturalists’ Club 306 Upper Bass Lake we (1,2,5) oe f &. ~ «Twin Lake tos (0,2,2) \ THE CANADIAN FIELD-NATURALIST FiGur_E 1. Number of Tree Swallow (Zachycineta bicolor) nestlings (2014, 2015, 2016) collected at each location on the Vol. 133 _ Waterfront \ | “i ; foipte aux Baptémes (2,3,2) o* Perch Lake’ | am (11,12,8) Canadian Nuclear Laboratories’ Chalk River property. Source: Chalk River Laboratories, Chalk River, Ontario, Canada. 46°03'00.2"N, 77°21'51.7"W. Google Earth Imagery date: 20 August 2019. Data providers: DigitalGlobe 2019. Accessed: November 2019. were taken, euthanized, their external surfaces veri- fied clean, and frozen. In 2013, carcasses were incinerated for the deter- mination of Sr-90 (Lee et a/. 2019). Frozen carcasses were thawed overnight in a refrigerator, then dehy- drated overnight in an oven, at 105°C. After cool- ing to room temperature, carcasses were heated to 250°C for 2 h and to 450°C for 16 h, with the 16 h in- cineration performed a second time to ensure com- plete ashing. After incineration, samples were cooled to room temperature. The ash was gently milled us- ing a spatula, and any material (i.e., stone, glass, and metal fragments) larger than about 1 mm in diameter was removed. Beginning in 2014, natural grit materi- als and anthropogenic litter larger than about 1 mm in diameter observed in the ashes were noted and pho- tographed. In 2016, on thawing of the carcasses, the gizzard contents of each bird were examined visually for ma- terials (e.g., insects, shells, plastic items) that would be incinerated during the ashing process. Observations were noted and the material was returned to the car- cass before each carcass was dried and incinerated as above; non-combustible materials larger than about 1 mm in diameter were removed and photographed af- ter the ashing process. Although the general type of litter was noted, pieces were not measured. Results In addition to small stones, metal, and/or glass fragments were found in five of 24 nestlings in 2014 and in 10 of 26 nestlings in 2015 (Table 1). Because the gizzards of nestlings collected in 2014 and 2015 were not examined before the nestlings were inciner- ated, results for combustible materials, such as plas- tic, are not available. In 2016, the gizzards were ex- amined prior to incineration and we observed flying insects mixed with small stones, sand, grass, and mollusc shells, as well as anthropogenic materials, including pieces of metal and glass, sections of wrap- pers (most often pieces of shiny cigarette and chew- ing gum wrappers up to ~1 cm wide) in seven of 24 birds (Table 1). Figures 2 and 3 provide examples of litter collected from nestlings sampled in 2014—2016. The presence of anthropogenic material in the nestlings occurred most often along the Ottawa River shoreline and around Perch Lake, where human ac- tivities are more prominent (Figure 1; Tables 2 and 3). Such material was seldom found in nestlings col- lected from more remote areas. In 2016, no significant differences (t-test, t= 1.146, P=0.281) were found between the weights of 12-day- old nestlings with (average 21.0 g, range 19.5—22.2 g) and without (21.8 g, 20.5—22.8 g) anthropogenic litter in their gizzards. 2019 WALSH ET AL.: ANTHROPOGENIC LITTER IN NESTLING TREE SWALLOWS 307 TABLE 1. Types of litter (>1 mm diameter) found in nestling Tree Swallows (Zachycineta bicolor), 2014-2016. Year No. nestlings Metal 2014 24 12.5 2015 26 30.8 2016 24 12.5 % nestlings containing fragments Glass Wrapper Plastic 42 n/a* n/a 11.5 n/a n/a 20.8 4.2 4.2 *n/a = not available because these materials would have been incinerated. FiGureE 2. Examples of metal turnings found in the whole body ashes of a Tree Swallow (Jachycineta bicolor) nest- ling. Photo: Jennifer Haughton. Ficure 3. Examples of glass fragments found in the whole body ashes of a Tree Swallow (Jachycineta bicolor) nest- ling. Photo: Jennifer Haughton. TABLE 2. Locations of nestling Tree Swallows (Zachycineta bicolor) with anthropogenic litter (>1 mm diameter) in their gizzards, 2016. No. nestlings containing anthropogenic fragments Location No. nestlings Metal Baggs Road 1 0 Maskinonge Lake 1 1 Upper Bass Lake 5 0 Twin Lake 2 0 Perch Lake 8 0 Pointe aux Baptemes 2 1 Waterfront 5 L* *Both fragment types were in the same nestling. Glass Wrapper Plastic 0 0 0 0 0 0 1 0 0 0 0 0 2 0 0 0 0 1 1 fet 0 TABLE 3. Locations of nestling Tree Swallows (Zachycineta bicolor) containing glass and metal pieces (>1 mm diameter) in 2014 and 2015. Location No. nestlings Baggs Road Maskinonge Lake Upper Bass Lake Twin Lake Perch Lake 2 Pointe aux Baptemes 5 Waterfront 12 WN WN Ww *Both fragment types were found in one nestling. Discussion Anthropogenic litter was found in 30% of 74 nestling Tree Swallows collected in 2014-2016 near Chalk River, Ontario. We consider this to be an un- derestimate, because it does not include litter frag- ments <1 mm in diameter or combustible litter for two of the three years of the study. Barrentine (1980) reported grit in 80% of Barn No. nestlings containing anthropogenic fragments Metal Glass 0 0 0 0 0 0 0 0 4 2 3 1 4* 1* Swallow (Hirundo rustica) nestlings sampled, pro- viding evidence that grit is an important dietary fac- tor during the growth of swallow nestlings and a cause for concern for birds that nest in areas where erit-like anthropogenic material may be present. Mayoh and Zach (1986) found that Tree Swallows had a greater percentage of anthropogenic litter in their “stomachs” than did House Wrens (7roglodytes 308 aedon) at the same age. This may be because swal- lows forage along shorelines and nearby roads (in a ~400 m feeding radius during the nestling period), where greater amounts of anthropogenic litter are generally found. Barrentine (1980) showed that while swallows consumed grit of various colours, sizes, and compositions, they have a clear preference for light-coloured objects between 1 and 3 mm in size. Considering metals are generally light in colour, and glass, plastic, and wrapper materials can also be a light colour, swallows could be intentionally choos- ing human-made materials over natural grit. Anthropogenic litter can be domestic or indus- trial. The presence of metal turnings in Tree Swallow nestlings was a unique finding that is particularly rel- evant to industrial areas. The potential detrimental effects of anthropogenic materials on birds are well known. For example, the ingestion of metal pieces by waterfowl can result in lead poisoning (Bellrose 1975; Trost 1981), and the occurrence and impacts of plastic ingestion by bird species, especially marine birds, are prevalent (see for example Provencher ef a/. 2014). Reported adverse health effects include: proventricular impactions, ul- cerative lesions (Azzarellow and Van Vleet 1987; Fry et al. 1987); digestive tract blockages, stomach lining damage, appetite suppression (Azzarellow and Van Vleet 1987; Laist 1987); exposure to polychlorinated biphenyls and organochlorine pesticides (Colabuono et al. 2010); and lowered steroid hormone levels, de- layed ovulation, and reproductive failure (Azzarellow and Van Vleet 1987). We have documented the presence of anthropo- genic litter in young Tree Swallows, in an environ- ment previously considered to be relatively litter free. While we observed that the ingestion of litter did not significantly impact the weights of the nestlings, po- tential risks of ingestion of anthropogenic litter on Tree Swallow nestlings remain to be investigated. Author Contributions Writing — Original Draft Preparation: S.W.; Writ- ing—Review & Editing: L.B., A.F.,1LG., .H.,D.L.,MS., and S.W.; Methodology: D.L. and MLS.; Investigation: L.B., A.F., LG, and J.H.; Resources: L.B.; Data analy- sis: A.F. and I.G.; Visualization: L.B.; Project Admin- istration: M.S.; Supervision: D.L. and MLS. Acknowledgements This work was funded through Canadian Nuclear Laboratories’ research and development programs. All animal work was conducted in accordance with a collection permit issued by Environment and Climate Change Canada (#CA 0315) and an Animal Care Protocol approved by the Chalk River Animal Care THE CANADIAN FIELD-NATURALIST Vol. 133 Committee (in compliance with the guidelines estab- lished by the Canadian Council on Animal Care). Literature Cited Azzarellow, M.Y., and E.S. Van Vleet. 1987. Marine birds and plastic pollution. Marine Ecology Progress Series 37. 295-303. https:/doi.org/10.3354/meps037295 Barrentine, C.D. 1980. The ingestion of grit by nest- ling Barn Swallows. Journal of Field Ornithology 51: 368-371. Bellrose, F.C. 1975. Impact of ingested lead pellets on water- fowl. International Waterfowl Symposium 1: 163-167. Best, L.B., and J.P. Gionfriddo. 1991. Characterization of grit use by cornfield birds. Wilson Bulletin 103: 68—82. Colabuono, F.I., S. Taniguchi, and R.C. Montone. 2010. Polychlorinated biphenyls and organochlorine pesti- cides in plastics ingested by seabirds. Marine Pollution Bulletin 60: 630-634. https:/doi.org/10.1016/j.marpol bul.2010.01.018 De Steven, D. 1980. Clutch size, breeding success, and pa- rental survival inthe Tree Swallow Uridoprocne bicolor). Evolution 34: 278-291. https:/do1.org/10.2307/2407392 Fry, D.M., S.I. Fefer, and L. Sileo. 1987. Ingestion of pla- stic debris by Laysan Albatrosses and Wedge-tailed Shearwaters in the Hawaiian Islands. Marine Pollu- tion Bulletin 18: 339-343. https://doi.org/10.1016/S002 5-326X(87)80022-X Gionfriddo, J.P., and L.B. Best. 1995. Grit use by House Sparrows: effects of diet and grit size. Condor 97: 57-67. https://doi.org/10.2307/1368983 Laist, D.W. 1987. Overview of the biological effects of lost and discarded plastic debris in the marine environment. Marine Pollution Bulletin 18: 319-326. https:/doi.org/ 10.1016/S0025-326 X(87)80019-X Lee, D.R., J. Haughton, A. Valente, L. Bellan, M. Stuart, D. Beaton, H. Chen, I. Gosselin, and A. Festarini. 2019. Effects of 90Sr on tree swallow nestlings near groundwater contaminant plumes. Health Physics 117: 267-277. https://doi.org/10.1097/HP.0000000000001076 Mayoh, K.R., and R. Zach. 1986. Grit ingestion by nestling Tree Swallows and House Wrens. Canadian Journal of Zoology 64: 2090-2093. https:/doi.org/10.1139/z86-319 Provencher, J.F., A.L. Bond, A. Hedd, W.A. Montevecchi, S.B. Muzaffar, S.J. Courchesne, H.G. Gilchrist, S.E. Jamieson, F.R Merkel, K. Falk, J. Durinck, and M.L. Mallory. 2014. Prevalence of marine debris in marine birds from the North Atlantic. Marine Pollution Bulletin 84: 411-417. https://doi.org/10.1016/j.marpolbul.2014.04.044 Robertson, R.J., and W.B. Rendell. 1990. A comparison of the breeding ecology of a secondary cavity nesting bird, the Tree Swallow (Jachycineta bicolor), in nest boxes and natural cavities. Canadian Journal of Zoology 68: 1046-1052. https:/do1.org/10.1139/z90-152 Trost, R.E. 1981. Dynamics of grit selection and retention in captive Mallards. Journal of Wildlife Management 45: 64-73. https:/doi.org/10.2307/3807874 Received 1 February 2019 Accepted 25 February 2020 The Canadian Field-Naturalist Note First record and new host record of the obligate dulotic ant, Polyergus bicolor (Hymenoptera: Formicidae), in Alberta, Canada CHRISTINE E. SostaAk'*, MARI WEST”, and JAMES R.N. GLASIER? 'Federated Department of Biology, New Jersey Institute of Technology/Rutgers-Newark, Newark, New Jersey 07102 USA "Department of Entomology, University of California Riverside, Riverside, California 92521 USA 3Métis Nation of Alberta, Environment Division, Edmonton, Alberta TSG X05 Canada “Corresponding author: ces43@njit.edu Sosiak, C.E., M. West, and J.R.N. Glasier. 2019. First record and new host record of the obligate dulotic ant, Polyergus bicolor (Hymenoptera: Formicidae), in Alberta, Canada. Canadian Field-Naturalist 133(4): 309-312. https://do1. org/10.22621/cfn.v13314.2381 Abstract We describe the discovery of Polyvergus bicolor, an obligate slave-making ant species, as a new provincial record in Alberta. This species was previously known mostly from eastern Canada and the northeastern United States and has been sparsely collected: only once in the past 50 years. Polyergus bicolor was discovered parasitizing Formica podzolica, which is also a new host for the species. This discovery marks a significant expansion of both range and host for P. bicolor. Key words: Polyergus bicolor; dulotic parasitism; range expansion; host expansion; Alberta; Formica podzolica Polyergus (Latreille 1804) is a predominantly hol- arctic genus of ants that contains 14 species, 11 of which are present in North America (Trager 2013). All Polyergus display obligate dulotic behaviour (slave-making), making them a remarkable genus that has received a good deal of research interest. Colony foundation occurs when a mated Polyergus queen enters a Formica nest, kills the queen, and usurps her role, with Formica workers taking care of her and her brood (Hdlldobler and Wilson 1990). To maintain Formica worker populations in the colony, Polyergus workers locate a host nest, and then raid it for pupae, prepupae, and occasionally last-instar lar- vae. When the Formica pupae mature to adults in the Polyergus nest, they accept that nest as their own, and perform the majority of tasks within the colony (Trager 2013). Host Formica species vary, depending on the Polyergus species: some Polyergus will para- sitize only one Formica species, while others are ca- pable of parasitizing multiple species. Generally, the host species is from the Formica fusca group or the Formica pallidefulva group (Trager 2013). In western North America, Polyergus is over- whelmingly represented by Polyergus mexicanus (Trager 2013; Glasier et al. 2016); in Idaho, P. brev- iceps is also present (Wheeler 1917; Smith 1947; Trager 2013). (Note: there are generally no accepted common names for ants.) Polyergus bicolor was previously reported as restricted to eastern North America: Ontario to Illinois (Smith 1947; Wheeler 1968; Trager 2013). It was reported as far west as Saskatchewan and Montana by Wheeler (1917) as Polyergus rufescens bicolor. It was only confirmed as far west as the Dakotas by Trager (2013), who raised it to the status of species. Trager noted that he was un- able to collect any P. bico/or during the course of his study within its historical range, save for one collec- tion made in Wisconsin. In the last 50 years, he had found no P. bicolor collection records from its histor- ical range (Trager 2013). We first found P. bicolor in Alberta in sum- mer 2017. We collected two colonies in Jarvis Bay Provincial Park, on Sylvan Lake, while col- lecting and observing Formica colony behaviour. Jarvis Bay Provincial Park is a drywood boreal for- est characterized by mostly deciduous stands dom- inated by Trembling Aspen (Populus tremuloides Michaux), Balsam Poplar (Populus balsamifera L.), Black Spruce (Picea mariana (Miller) Britton, Sterns and Poggenburgh), and White Spruce (Picea glauca (Moench) Voss); prior records of P. bicolor note that it nests mostly in mesic forest, generally in rotten A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 309 ©The Ottawa Field-Naturalists’ Club 310 stumps or fallen logs, thus habitat similar to Jarvis Bay (Trager 2013). The specimens were collected by hand around the provincial park campsite after mistaking them for a species of the Formica rufa or Formica sanguinea Species groups. They were found in domed dirt and debris mounds with the host species Formica pod- zolica, identified using published keys (Francoeur 1973; Glasier et al. 2013). Our Polyergus specimens were identified using Trager’s revised key to global Polyergus species (Trager 2013). They differ from P. mexicanus, the other known Polyergus species in the area, by the degree of dark colouration on the abdo- men and a complete lack of pilosity on both the vertex of the petiole and the pronotum (Glasier et al. 2013; Trager 2013). A second collection occurred in July 2018 near Hay Lakes, Alberta, an area dominated by mixed de- ciduous woodlands (Trembling Aspen and Balsam Poplar) similar to Sylvan Lake. They were collected from a rounded mound within a grass meadow and were also using F: podzolica as a host. Polyergus bi- color has been formally recorded parasitizing both Formica neorufibarbis and Formica subaenescens, but not F- podzolica. The mounds in which we found P. bicolor were unlike their normal reported nesting sites, but this could be the result of their using a dif- ferent host species. This discovery represents a significant expansion of P. bicolor’s previously known range, although it supports Wheeler’s (1917) reports of P. bicolor in Saskatchewan as P. r. bicolor. Although the habi- tat where we found P. bicolor in Alberta is similar to the type of habitat from which it was previously known, the climate of Alberta is distinct from that of southern Ontario and the northeastern United States. The expansion of host species to include F: podzol- ica is also notable; Polyergus may use one or several hosts species but tends to show high fidelity to one host for a given population. Within a Polyergus spe- cies, if different populations are using different hosts, they are often highly specialized to their own host species. Populations show distinct chemical and ge- netic divergence from one another, perhaps reflect- ing incipient speciation (Torres et al. 2018). Because newly mated Polyergus queens typically stay with the host species of their parent colony, this fidel- ity is passed down from generation to generation (Hoélldobler and Wilson 1990). Formica podzolica is widespread throughout North America and its range overlaps with that of P. bicolor in the northeastern United States (Wheeler and Kannowski 1994; Ellison et al. 2007); thus, it is difficult to say where host ex- pansion took place. Further genetic work would shed light on potential divergence between P. bicolor pop- THE CANADIAN FIELD-NATURALIST Vol. 133 ulations in western and eastern North America, con- tingent on their host species. Voucher specimens Canada, Alberta: Sylvan Lake Jarvis Bay, 52.347°N, 114.091°W and 52.345°N, 114.089°W, hand collected, 21 July 2017, C. Sosiak (Figure 1, personal collections of Christine Sosiak and Mari West). Canada, Alberta: 4 km SE of Hay Lakes, Aspen Parkland, 53.165°N, 113.014°W, hand collected, with F-. podzolica, 27 July 2018, J.R.N. Glasier (Strickland Museum and personal collection of J.R.N. Glasier). Strickland Museum accession numbers: P. bicolor specimens UASM396245, UASM396246; F. podzol- ica specimens UASM396247, UASM396248. Author Contributions Writing — Original Draft: C.E.S.; Writing — Re- view & Editing: C.E.S., M-W., and J.R.N.G.; Species Collection: C.E.S., M.W., and J.R.N.G.; Species Iden- tification: C.E.S. and J.R.N.G.; Funding Acquisition: MW. Acknowledgements We collected Polyergus specimens while con- ducting research funded by Alberta Conservation Association Grants in Biodiversity. Additional sup- port for this work was provided in part by University of California Riverside, National Science Foundation Research Traineeship for Integrated Computational Entomology, award 1631776. We thank John Acorn, James Trager, Jessica Purcell, and Phillip Barden, as well as an anonymous reviewer, for their helpful comments on a draft of this manuscript. Literature Cited Ellison, A.M., S. Record, A. Arguello, and N.J. Gotelli. 2007. Rapid inventory of the ant assemblage in a tem- perate hardwood forest: species composition and assess- ment of sampling methods. Environmental Entomology 36: 766-775. https://doi.org/10.1093/ee/36.4.766 Francoeur, A. 1973. Révision taxonomique des especes néarctiques du groupe Fusca, genre Formica (Formic- idae, Hymenoptera). Mémoires de la Société Entomo- logique du Québec 3. Entomological Society of Quebec, Quebec, Canada. Glasier, J.R.N., J.H. Acorn, S.E. Nielsen, and H. Proc- tor. 2013. Ants (Hymenoptera: Formicidae) of Alberta: a key to species based primarily on the worker caste. Canadian Journal of Arthropod Identification 22: 1— 104. https://doi.org/10.3752/cjai.2013.22 Glasier, J.R.N., S. Nielsen, J.H. Acorn, L.H. Borysenko, and T. Radtke. 2016. A checklist of ants (Hymenoptera: Formicidae) of Saskatchewan. Canadian Field-Naturalist 130: 40-48. https://do1.org/10.22621/cfn.v13011.1791 Hdlldobler, B., and E.O. Wilson. 1990. The Ants. Belknap (Harvard University Press), Cambridge, Massachusetts, USA. SOSIAK ET AL.: PROVINCIAL RECORD FOR POLYERGUS BICOLOR Figure 1. Lateral a. and frontal b. views of a Polyergus bicolor specimen collected in Jarvis Bay Provincial Park, Sylvan Lake. Photos: Christine Sosiak. BIZ Latreille, P.A. 1804. Tableau méthodique des insectes. Page 179 in Nouveau dictionnaire d’histoire naturelle. Edited by Société de Naturalistes et d’Agriculteurs. Dé- terville, Paris, France. Smith, M.R. 1947. A study of Polyergus in the United States, based on the workers (Hymenoptera: Formic- idae). American Midland Naturalist 38: 150-161. Torres, C.W., M.A. Tonione, S.R. Ramirez, J.R. Sapp, and N.D. Tsutsui. 2018. Genetic and chemical diver- gence among host races of a socially parasitic ant. Ecol- ogy and Evolution 8: 11385-11398. https://doi.org/10.10 02/ece3.4547 Trager, J.C. 2013. Global revision of the dulotic ant genus Polyergus (Hymenoptera: Formicidae, Formicinae, For- THE CANADIAN FIELD-NATURALIST Vol. 133 micini). Zootaxa 3722: 501-548. https://doi.org/10.11646/ zootaxa.3722.4.5 Wheeler, G.C., and P.B. Kannowski. 1994. Checklist of the ants of Michigan (Hymenoptera: Formicidae). Great Lakes Entomologist 26: 297-310. Wheeler, J. 1968. Male genitalia and the taxonomy of Poly- ergus. Proceedings of the Entomological Society of Washington 70: 156-164. Wheeler, W.M. 1917. The mountain ants of western North America. Proceedings of the American Academy of Arts and Science 52: 457-569. Received 24 October 2019 Accepted 24 December 2019 The Canadian Field-Naturalist Tall grass prairie ecosystem management—a gastropod perspective ANNEGRET NICOLA“, ROBERT G. FORSYTH?, MELISSA GRANTHAM“, and Cary D. HAMEL‘ 'Université Rennes, UMR CNRS 6553 EcoBio, Station Biologique Paimpont, Paimpont 35380 France "Western University, Department of Biology, 1151 Richmond Street North, London, Ontario N6A 5B7 Canada ’New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada “Nature Conservancy of Canada, Manitoba Region, Suite 200 - 611 Corydon Avenue, Winnipeg, Manitoba R3L OP3 Canada “Corresponding author: annegret.nicolai@univ-rennes1 .fr Nicolai, A., R.G. Forsyth, M. Grantham, and C.D. Hamel. 2019. Tall grass prairie ecosystem management—a gastropod perspective. Canadian Field-Naturalist 133(4): 313-324. https://doi.org/10.22621/cfn.v13314.2217 Abstract Less than 5% of the original tall grass prairie in North America remains. A portion of this remnant, composed of wetland, grassland and forest, is protected by the Nature Conservancy of Canada (NCC) in southern Manitoba. This heterogene- ous ecosystem has rich biodiversity; however, gastropods have not been surveyed in Canada’s tall grass prairie. We studied gastropods in Prairie, Wet Meadow, Forest, and Wet Forest habitats of the Manitoba Tall Grass Prairie Preserve that vary with respect to land management practices (prescribed burning, grazing by cattle). Gastropod community composition was unique in the Prairie where mounds of grass litter form permanently moist cavities harbouring aquatic species, while dry-habitat species colonized the upper parts of these mounds. Gastropod communities in Prairie habitats were negatively affected by grazing and burning that occurred in the five years prior to our survey. Unburned Forest patches included both forest gastropod species and edge effect influenced open-habitat species and harboured the most diverse gastropod commu- nities. These unburned Forest patches potentially provide a species pool for post-burn prairie recolonization. The gastropod community of Wet Meadows was not affected by grazing and was composed mainly of aquatic species. In this gastropod survey five species were recorded from Manitoba for the first time. The rare Blade Vertigo (Vertigo milium) is also reported. Key words: Fire; grazing; freshwater snails; terrestrial snails and slugs; protected area; rare species; Manitoba Introduction Tall grass prairie once covered 68 million ha of North America before conversion to urban areas or cropland in the United States and Canada—less than 5% remains (Sampson and Knopf 1994). Tall grass prairie harbours a diversity of terrestrial and aquatic plants and animals within a mosaic of grassland, pot- hole-forming wetland/grassland systems, and shrubby/ wooded areas. Ecosystem services that include nu- trient cycling, water retention, aquifer recharge, the storage of atmospheric carbon, as well as enhanced water infiltration and improved runoff water qual- ity are all of great ecological and economic impor- tance (Glaser 2012). As a result of human activity, prairies are the most highly impacted of any of the continent’s terrestrial ecosystems. Current threats to the biodiversity and ecological functioning of the re- maining tall grass prairie include: habitat fragmen- tation, loss by conversion to cropland, incompatible grazing practices, undesirable habitat changes due to fire and fire exclusion, spread of invasive plant spe- cies, and stream degradation due to incompatible land management practices and soil erosion (Glaser 2012). The biodiversity of northern tall grass prairie has been poorly explored, especially that of soil related functional animal groups, such as terrestrial gastro- pods. Terrestrial gastropods are generally under-sur- veyed in most of Canada and are usually absent from management strategies for protected areas. Being a significant component of biodiversity among ground dwelling species, terrestrial gastropods are globally declining (Lydeard et al. 2004) and play a crucial role in ecological processes (Jordan and Black 2012) by aiding in decomposition, nutrient cycling and soil building processes, and by providing food and es- sential nutrients to wildlife. Also, terrestrial gastro- pod abundance and diversity can be used as ecolo- gical indicators at the litter-soil interface, such as for logging practice management in forests (British Columbia Ministry of Forests 2008). Previously, only a few terrestrial gastropod surveys have occurred in Manitoba, e.g., by the Manitoba Museum and by Nekola (2005), and none of these targetted the com- munity in the tall grass prairie. Humans have long used fire to influence North American ecosystems, including First Nations who A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 313 ©The authors. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 314 used fire to create large areas of grassland in the Great Plains regions (Pyne 1983; Botkin 1990). While First Nations used fire to promote a habitat mosaic and a resource diversity that provided greater stability to their lives, later European settlers used burns to create uniformity in ecosystems (Lewis 1985). Prescribed fire has become an important management tool for prairie and forest conservation in North America (Gottesfeld 1994; Williams 2000), and is used to limit the spread of invasive plants (Brooks and Lusk 2008), promote growth and reproduction in native prairie vegetation (Towne and Owensby 1984), and improve and expand habitat for grassland and parkland birds (e.g., Burkman 1993; Madden et al. 1999; Davis et al. 2000; Ludwick and Murphy 2006; Vierling and Lentile 2006; Buehler et a/. 2007; Grant et al. 2010; Klaus et al. 2010; Austin and Buhl 2013) and rare prairie plants (e.g., Becker 1989; Bleho et al. 2015). Some authors have expressed concern about the det- rimental impacts of prescribed burns on prairie that include providing optimal germinating conditions for invasive plant seedlings by opening the vegetation canopy (Ohrtman er a/. 2011), and negative direct and indirect effects on the abundance of small mammals (Kaufman et al. 1990), birds (Reinking 2005), arthro- pods (Swengel 1996; Harper et a/. 2000), and terres- trial gastropods (Nekola 2002; reviewed by Saestedt and Ramundo 1990; Knapp et al. 2009). In addition to structural modification by fire, tall grass prairie has also been intermittently grazed by large ungulates, 1.e., Bison (Bison bison, Knapp et al. 1999). Domestic Cattle (Bos taurus) are now the dominant grazers at most prairie sites. Grazing can enhance plant diversity by encouraging the growth of some prairie species (Damhoureyeh and Hartnett 1997, 2002). The effect on prairie fauna is also selec- tive; birds (Sliwinski 2012 as cited in Glaser 2012), arthropods (van Klink et al. 2015), and terrestrial gastropods (Baur et al. 2007) respond differently to grazing regimes, defined by stocking rate, grazing frequency, and livestock type. One of the largest remaining tall grass prairie complexes in Manitoba is protected by the Nature Conservancy of Canada (NCC) and partners as part of the Manitoba Tall Grass Prairie Preserve (MTGPP). As part of an effort to preserve tall grass prairie biodi- versity and the ecosystem services it provides, man- agers need to understand how management practices influence the gastropod community in the MTGPP. Currently, the NCC uses rotational prescribed burn- ing and grazing by cattle to maintain a spatial and structural mosaic of grassland, wetland, and forest within the tall grass prairie system. For managers of protected prairie habitat, such as the NCC, the ques- tion of which management strategy to apply remains THE CANADIAN FIELD-NATURALIST Vol. 133 an ongoing challenge. The goal is to identify the ap- propriate regime of prescribed burns and grazing re- quired to maintain a generally rich floral and faunal diversity without negatively affecting the diversity of functional groups, such as gastropods involved in lit- ter-soil processes, or those of conservation concern. To assess the impact of current management prac- tices on the composition of the gastropod community in the MTGPP a gastropod survey was undertaken at variously managed sites (burning or grazing within the past five years) within the preserve. Study Area The 4100 ha MTGPP is located in the rural mu- nicipality of Stuartburn, in southeastern Manitoba, Canada (Figure 1). The majority (>70%) of MTGPP lands are owned by the NCC; the Manitoba Habitat Heritage Corporation and Nature Manitoba own the remainder. Preserve lands are jointly man- aged through a Management Committee that in- cludes landowners as well as Manitoba Sustainable Development and Environment and Climate Change Canada. The preserve is comprised of dozens of dis- tinct management units, allowing temporal and spa- tial variation in management practices. The habitats comprise two grassland types (Prairie and Wet Meadow) and two woodland patches (Forest and Wet Forest). The two woodland types range from small groves to larger forest areas and provide mostly edge habitat, but also include humid microhabitats under logs and drier microhabitats on the surface of logs and on branches. The habitat types (classification based on Minnesota Department of Natural Resources 2005) differ in vegetation com- position and structure as well as in seasonal cycle of flooding as follows: Prairie (P) Tall grass prairie communities dominated by tall and mid-height graminoid species up to 1.50 m tall. Big Bluestem (Andropogon gerardi Vitman), Prairie Dropseed (Sporobolus heterolepis (A. Gray) A. Gray), Little Bluestem (Schizachyrium scoparium (Michaux) Nash), Yellow Indiangrass (Sorghastrum nutans (L.) Nash), and Plains Porcupine Grass (Hesperostipa spartea (Trinius) Barkworth) are most common. Not flooded but forming very humid micro- habitats of roots and decaying grass leaves in the up- per soil layer between mounds of grass. Wet Meadow (WM) Meadow dominated by graminoid species up to 0.50 m tall. Broad-leaved species such as Slim- stemmed Reedgrass (Calamagrostis stricta (Timm) Koeler), Prairie Cordgrass (Sporobolus michauxi- anus (Hitchcock) P.M. Peterson & Saarela), Sartwell’s Sedge (Carex sartwellii Dewey), and Woolly Sedge 2019 NICOLAI ET AL.: GASTROPODS IN TALL GRASS PRAIRIE 315 96°45'W Survey site Disturbed upland Prairie wet meadow Northern mesic prairie Northern wet prairie Northwestern dry-mesic-oak woodland Northwestern mesic-aspen-oak woodland Northwestern aspen woodland Northwestern wet aspen forest Northwestern wet-mesic aspen woodland 96°45'W 96°42'W 96°39'W 96°42'W 96°39'W FiGcure 1. Phytosociology classification in the Manitoba Tall Grass Prairie Preserve and gastropod survey sites: F = Forest, WE = Wet Forest, P = Prairie, WM = Wet Meadow. Two drainage sites have also been analyzed, D1/D2 and D3/D4, with sampling points on each side of the road. (Carex pellita Muhlenberg ex Willdenow) are typical, with Tussock Sedge (Carex stricta Lamarck) an oc- casional dominant. Habitat is subjected to moderate inundation by standing water following spring thaw and heavy rains, and to periodic drawdowns during the summer. Forest (F) Forest patches within the grassland that are not flooded and are dominated by trees and herbaceous species. Herbaceous plant cover commonly includes: Wild Lily-of-the-valley (Maianthemum canadense Desfontaines), Northern Bedstraw (Galium boreale 316 L.), Wild Sarsaparilla (Aralia nudicaulis L.), Ame- rican Vetch (Vicia americana Muhlenberg ex Will- denow), and Lindley’s Aster (Symphyotrichum cili- olatum (Lindley) A. Léve & D. Love). Bur Oak (Quercus macrocarpa Michaux) and Trembling Aspen (Populus tremuloides Michaux) are dominant tree species. Wet Forest (WF) Forest patches dominated by trees and herbaceous Species that are subjected to the same inundation re- gime as Wet Meadow sites. Herbaceous cover com- monly includes: Star-flowered False Solomon’s Seal (Maianthemum stellatum (L.) Link), Wild Strawberry (Fragaria virginiana Miller), Northern Bedstraw (G. boreale), Calico Aster (Symphyotrichum lateriflo- rum (L.) A. Love & D. Love), and Dwarf Raspberry (Rubus pubescens Rafinesque). Trembling Aspen (Populus tremuloides), Balsam Poplar (Populus bal- samifera L.), or Black Ash (Fraxinus nigra Marshall) are the most important tree species. Historical and recent fire and grazing manage- ment on MTGPP property is highly diverse; there is no specific information on the historical frequency of grazing or burning for this area. Long-term manage- ment plans include prescribed burns once every five years, typically in spring or fall. In the year prior to prescribed fire, properties are not grazed. However, the interval between fires can be variable due to occa- sional wildfires and seasonal weather conditions not conducive to the use of prescribed fire (Bleho ef al. 2015). A twice-over rotational grazing system is used at the MTGPP but is individually managed by cat- tle owners. Information on frequency and intensity of grazing and fire was not available. Timing of fire and grazing (Table 1) was based on best available in- formation. Sites within historically human-built drains are also part of the MTGPP ecosystem, and potentially could serve as a source for post-management recol- onization by gastropods. Drainage wells were there- fore also investigated for richness and abundance of aquatic gastropods that might be available to colonize wet and flooded grassland or forest habitat. Methods All sites (7 = 16) examined within the MTGPP had been managed either by burning or grazing (ex- clusively cattle) within the last five years (n = 10) or had received no active management over the past five years or more (n = 6). Although we were able to sam- ple recently managed (<5 years) and unmanaged (=5 years) sites for both woodland and grassland habitat types, we were unable to find any Wet Forest that had been subject to both grazing and burning in the pre- vious five years (Table 1). THE CANADIAN FIELD-NATURALIST Vol. 133 In September 2013, a visual search and hand col- lection of gastropods occurred in the litter and the up- permost soil layer using six 0.2 x 0.2 m plots per site (Figure 1). The plots were separated by a distance of at least 20 m on a random, non-linear transect to sam- ple different microhabitats within the same vegetation zone (= site). Additionally, four drainage-well sites of 10 x 10 m were searched during 30 min each for aquatic gastropod species to establish the full gastro- pod species list in the MTGPP. Gastropods were identified by A.N. and R.G.F. Vouchers of all species have been deposited in the Manitoba Museum (Catalogue numbers: MM65991 to MM65999 and MM66178 to MM66311). Live gas- tropods were stored as wet samples at the Biodiversity Institute of Ontario (BIO), Guelph, Ontario, and in- corporated into the BOLD database under field sam- ple numbers ANi039 to ANi056 and under the BIO collection numbers BIOUG09921-C08 to -CO9 and BIOUG09922-B10, -C02, -C05 to -C07, -C10 to -C12, -D01 to -D07, -E01, -E03 to -E12, and -FO02. Individu- als of each species were counted to determine abun- dance/m? for each site. Due to the diversity of management combinations, it was not possible to assemble a set of replicates, so we used an exploratory approach in our multivari- ate analysis. Species richness was compared among habitat types using an adjusted ¢-test (Welch) and one-way analysis of variance (ANOVA). The gastro- pod community composition was analysed with nonmetric multidimensional scaling (NMDS) with Euclidean distance similarity coefficient applied to TABLE 1. Management history of the sites in the Tall Grass Prairie Preserve, Manitoba, prior to the 2013 sampling year. Information about grazing was available since 1993 and about fire since 1992. F = Forest, WF = Wet Forest, P = Prairie, WM = Wet Meadow. Years since Years since Habitat Sites last grazing last fire Woodland Fl 5 2, F2 1 10 F3 <20 <21 F4 <20 <21 WFI1 <20 2 WEF2 1 <21 WF3 <20 <21 WF4 <20 <21 Grassland Pl 5 2 P2 o, <21 PS 5 <21 P4 <20 <2] WMI > 2 WM2 1 <21 WM3 <20 <21 WM4 <20 i 2019 abundance data (N/m’) based on the normalized minimal threshold density (Legendre and Legendre 2007, Ramette 2007). The stress coefficient indi- cates the badness-of-fit, this is the quality of the NMDS (S < 0.10: good). Mann-Whitney and Kruskal- Wallis tests were used on scores of the axes to ana- lyse gastropod community differences between hab- itat types. Hierarchical clustering was performed with the centroid method on a Euclidean distance matrix calculated on the abundance of gastropods (N/m?). Approximately unbiased P-values were com- puted by multiscale bootstrap resampling, m = 1000 (Shimodaira 2004). Spatial autocorrelation of com- munity composition was analysed with the Mantel test using Euclidean distance and m = 1000 permutations. Moran’s I was calculated for species richness on an inverse distance matrix. All analyses were conducted with the software R 2.8.0 (R Core Team 2008). Results The forest and grassland communities formed by terrestrial and aquatic species are distinguishable on the NMDS model (Figure 2) by scores on the first axis (Mann-Whitney, W= 55, n = 16, P = 0.01) and on the third axis (Mann-Whitney, W = 53, n = 16, P = 0.03), but not on the second axis (Mann-Whitney, W = 41, n= 16, P = 0.38). Species richness (Table 2) did not differ between forest and grassland communities (¢- test, t,, 9; = 0.88, P = 0.39). Likewise, species compo- sition (measured as scores on the three NMDS axes; Figure 2) and species richness (Table 2) were not sig- nificantly different among Forest, Wet Forest (forest communities), Prairie and Wet Meadow (grassland communities; axis 1: Kruskal-Wallis, y?; = 6.9, P = 0.07; axis 2: Kruskal-Wallis, y’?,; = 1.3, P = 0.73; axis 3: Kruskal-Wallis, 77; = 5.1, P = 0.16; species rich- ness: ANOVA, F; ;.= 0.27, P = 0.84). Nevertheless, nine of 23 gastropod species showed habitat prefer- ence based on presence in a single habitat type (Table 2). Among aquatic gastropods, six of nine species are characterized as vernal species (Clarke 1981), being m@ Wet Forest NICOLAI ET AL.: GASTROPODS IN TALL GRASS PRAIRIE ST generally restricted to periodically flooded terres- trial habitats (Table 2). Only two of the vernal spe- cies were absent from the drainage well sites (with permanent water). Some typically open-habitat spe- cies, such as Costate Vallonia (Vallonia costata (O.F. Miller, 1774)) and Trumpet Vallonia (Vallonia par- vula Sterki, 1893), were only observed at the forest edge. Glossy Pillar (Cochlicopa lubrica (O.F. Miller, 1774)), Small Spot (Punctum minutissimum (1. Lea, 1841)), and V. parvula were only found in the dry, un- flooded, Forest, while Tapered Vertigo (Vertigo ela- tior Sterki, 1894), a species preferring very wet habi- tats (Nekola and Coles 2010), only occurred in the Wet Forest. Blade Vertigo (Vertigo milium (Gould, 1840)), a wet grassland species (Nekola and Coles 2010), was only recorded in Prairie sites while Multirib Vallonia (Vallonia gracilicosta Reinhardt, 1883) occurred only in Wet Meadow. Marsh Hive (Euconulus cf. prati- cola (Reinhardt, 1883); = E. alderi (Gray, 1840), see Forsyth and Oldham 2016), also a wet grassland spe- cies (Forsyth 2004, 2005), occurred in both grassland habitats. The cluster analysis of the gastropod commu- nity composition (Figure 3) based on the distances in the NMDS model (Figure 2) showed three dis- tinctive clusters (cluster P3-P4, cluster WM2-WM4, and a cluster including the remaining sites) that were not explained by spatial autocorrelation (Mantel test, z =-0.03, P = 0.50). Moreover, species richness was not spatially autocorrelated (Moran test, /,,, = —0.16, Tsp = 0.07, SD = 0.11, P = 0.38). This result indicated that management practices may influence gastropod community composition in some habitats. While, two recently managed Prairie sites, Pl and P2, were not significantly different from most sites (Figures 3 and 4a), P3 and P4, left unmanaged for at least five years, had a unique community composition charac- terized by high abundance of aquatic species (Figures 3 and 4a). P3 and P4 had deep litter filled holes be- tween mounds of grass, whereas recently burned O Forest WM1 ® Wet Meadow © Prairie ard i “TWF ar F2 | La as v | ‘i ie WF29F 7 no >| 2 wM 6 | P4 WM? 3 WES. 8 -10| é wM4 52008 Miia = -15 b k=3 (7,9 4 2 -10 9, a a a a A YE a Ww -30 -25 -20 -15 -10 -5 0 5 10 g NMDS axis 1 Ss FiGure 2. NMDS plot of gastropod communities in different habitats occurring in the Tall Grass Prairie. S indicates the stress and k the total number of axes used in the analysis. F = Forest, WF = Wet Forest, P = Prairie, WM = Wet Meadow. 318 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Frequency and abundance of gastropods in different habitats in the Tall Grass Prairie Preserve, Manitoba (fre- quency / minimal-maximal abundance per m? in four sites per habitat). Species richness is indicated as mean + SE. Family Species Terrestrial gastropods Agriolimacidae Cochlicopidae Discidae Euconulidae Gastrodontidae Punctidae Pristilomatidae Succineidae Valloniidae Vertiginidae Vitrinidae Aquatic gastropods Lymnaeidae Planorbidae Physidae Deroceras laeve, Meadow Slug Cochlicopa lubrica, Glossy Pillar Discus whitneyi, Forest Disc Euconulus fulvus, Brown Hive Euconulus polygyratus, Fat Hive Euconulus cf. praticola, Marsh Hive Nesovitrea electrina, Amber Glass Striatura milium, Fine-ribbed Striate Zonitoides arboreus, Quick Gloss Punctum minutissimum, Small Spot Hawaiia minuscula, Minute Gem Mediappendix cf. vermeta, Suboval Ambersnail Novisuccinea ovalis, Oval Ambersnail Oxyloma sp., an ambersnail Vallonia costata, Costate Vallonia Vallonia gracilicosta, Multirib Vallonia Vallonia parvula, Trumpet Vallonia Vallonia pulchella, Lovely Vallonia Gastrocopta tappaniana, White Snaggletooth Vertigo elatior, Tapered Vertigo Vertigo milium, Blade Vertigo Vertigo ovata, Ovate Ambersnail Vitrina angelicae, Eastern Glass-snail Galba sp., a fossaria Stagnicola elodes, Marsh Pondsnail Gyraulus circumstriatus, Disc Gyro Gyraulus parvus, Ash Gyro Planorbella subcrenata, Rough Ramshorn Planorbella trivolvis, Marsh Ramshorn Promenetus umbilicatellus, Umbilicate Sprite Aplexa elongata, Lance Aplexa Physella gyrina, Tadpole Physa Species richness Prairie (P1, Table 1) had bare soil between small grass mounds over most of the habitat. A recently grazed Prairie (P2, Table 1) had smaller grass mounds and less litter than unmanaged Prairie, but the structure of the vegetation and the grass litter were not quan- tified. Although P1 and P2 had been managed within the past two years prior to our study, their gastro- pod community composition was similar to those of Forest and Wet Forest. In the Wet Meadow, the re- cently grazed (WM2) and unmanaged WM4 formed a cluster distinct from the remaining sites (Figures 2 and 3). WM2 and WM4, sites separated only by a small gravel road, had a higher abundance of aquatic Woodland Grassland Forest WetForest Prairie Net Dramoee eadow 4/3-5 4/3-5 1/3 3 3/3-5 2/3-8 1/8 2/3-8 3/5-I1 2/3-24 3/521 2/2-3 1/3 1/24 3/8-16 4/3-16 3/13-37 4/8-32 1/3 4/5-19 2/21-27—_-1/3 3/3-5 1/3 2/5-I1 1/13 RES 2/3-5 2 /3-11 2/5-16 1/3 2/5-I1 2/3-16 1/11 7138 1/16 3/3-5 1/3 F/3 1/3 (Bas 2/3-8 1/27 1/5 1/8 1/3 1/3 1/3 1/8 2/5-6 1/3 4/13-104 3/5-53 2/89 1/8 1/16 3/3-11 4/3-24 1/24 3/3-32 2/8-13 1/8 1/5 1/8 1/3 3/8-19 2/48-59 4/8-27 1/3 4/356 3/5-48 3/5-45 4/19-61 2/821 TOL1V3: F34135. 78210 “8 3205.. 604 11 species (Figure 5a) relative to other Wet Meadow sites (WM1 and WM3, Figure 5b). Discussion The prairie ecosystem is a patchy assemblage of grassland, groves, and small forests. In general, the different habitats are moist due to periodic flood- ing, especially in the Wet Forest and Wet Meadow. Species richness and gastropod community compo- sition are driven by climate parameters such as wa- ter balance at a large sub-continental scale (Horsak and Chytry 2014) and by soil moisture, temperature, and calcium-content at a local scale (Dvorakova and 2019 15 20 25 Height 10 FiGurRE 3. Dendrogram of gastropod community clusters in different habitats occurring in the Tall Grass Prairie. Approximately unbiased P-value computed by multi- scale bootstrap resampling (7 = 1000) are indicated on the branches. F = Forest, WF = Wet Forest, P = Prairie, WM = Wet Meadow. NICOLAI ET AL.: GASTROPODS IN TALL GRASS PRAIRIE 319 Horsak 2012; Hettenbergerova et al. 2013). Because of the small size of the forest patches, most of the for- ested habitat includes forest edges that are suitable for open land species. Therefore, community composi- tion in general is very similar for most grassland and woodland sites. However, the Prairie sites, when un- disturbed by human activity, host a very particular gastropod community, characterized by the presence of vernal species, such as Lance Aplexa (Aplexa elon- gata (Say, 1821)) and Umbilicate Sprite (Promenetus umbilicatellus (Cockerell, 1887)), and the presence of dry-habitat species, such as Lovely Vallonia (Vallonia pulchella (O.F. Miller, 1774)) and V. gracilicosta. Dead vegetation accumulates in prairie habitat over years to form mounds of grass litter. Within these mounds, cavities retain water permanently. This per- manent water availability allows aquatic species to colonize the cavities within mounds of grass litter, while upper parts of the mounds are exposed to dry- ing and are suitable for dry-habitat species. Burning MP1 OP2 a 120 100 ‘Se 80 © = 60 cD) oO OO — SSS oS e joe bo ey i Di Q = 0 WO? WW wr _® WW WO WM De W@W WwW w’ FF FPF GF oA 07 sh” gh 6? ° &) PO GO WM GMT Ve ‘a 6% as ET MK we 4+ © & of & Yr Ge Xe \} “ Terrestrial Aquatic Gastropod species mP3 OP4 b 1 100 7 80 ™~ ae 60 fs) Pa 40 S c 20 =) Q 60 c — 40 ec £ 20 fo} =< 0 oe ~” aS we & eo 28 x0 Os &, e” > Q Rs »” se se se Fgh ge oF Gh a & Cs & cs gh & - os a Ae ; cS w Q? $e rl Terrestrial Aquatic Gastropod species b gWM1 OWM3 120 _. 100 £ 80 e > 60 oO SA ec i oe (a ne} xt oO Terrestrial ) Rs ayy ° &° As Rs & & Ky. NG we? & oO Aquatic Gastropod species FicureE 5. Abundance of gastropod species in a. recently grazed Wet Meadow (WM2) and in a Wet Meadow where the last management was at least five years ago (WM4), and in b. recently burned Wet Meadow (WM1) and in a Wet Meadow where the last management was at least five years ago (WM3), in the Tall Grass Prairie Preserve in Manitoba. WM2 and WM4 form a distinct cluster in the NMDS model (see Figures 2 and 3). Pooled species richness for WM1 and WM3 is nine terrestrial and four aquatic gastropod species, and for WM2 and WM4 is six terrestrial and six aquatic gastropod species. See Table 2 for full species names. and grazing, as well as trampling by cattle, may re- duce the mounds of grass litter and impact the aquatic micro-habitat. For this reason, only undisturbed Prairie sites are unique, harbouring a gastropod com- munity composed of both aquatic and dry habitat spe- cies. However, our characterization of sites as “undis- turbed” or “unmanaged” only means that they have not be subject to grazing or burning in recent years. All sites sampled had a prior history of burning and grazing, with the rotation of these land management practices over time and space nonetheless responsi- ble for this unique faunal assemblage. Without fire at some point, these sites would all have transitioned to Forest or Wet Forest, and thus harbour a different gastropod community. Short term declines in gastro- pod diversity or abundance that result from manage- ment measures may be an essential part of ensuring the long term maintenance of both grassland and the gastropod species dependent on open habitats. Nekola (2002) indicates that frequent prescribed burns represent a significant threat to the diversity of grassland snail communities, directly and indi- rectly affecting snail survival. Snails in the upper Lit- ter layer die from exposure to high heat during burn- ing (Nekola 2002). Post-burn mortality of snails is also high due to desiccation, due to the loss of shelter and micro-habitats (Ray and Bergey 2015). Fire de- stroys that part of the soil habitat upon which most litter-soil organisms depend and 1s therefore the most important factor affecting survival (Bellido 1987). In prairie habitat this means the loss of mounds of grass litter forming moist cavities. At the recently burned Prairie sites soil was bare between the re-es- tablishing mounds of grass litter, but moist cavities were absent. Therefore, aquatic species were nega- tively affected by the loss of micro-habitat. Burning may be beneficial for forest/grassland species, such as V. milium (only observed in the burned Prairie site in our study), which can exploit open-burned habitat close to the forest edge. Similarly, in the 2019 Mediterranean region Santos et al. (2012) only found gastropods of the family Geomitridae: Montserrat Heath Snail (Xerocrassa montserratensis (Hidalgo, 1870)) and Striped Heath Snail (Xerocrassa penchi- nati (Bourguignat, 1868)), two endemic open-habi- tat species, in burned sites. The availability of cryp- tic refuges within these Mediterranean forest patches (Kiss and Magnin 2003, 2006) presumably facilitates the survival of open-habitat snail populations (Santos et al. 2012). Likewise, in the tall grass prairie system, the manner in which fire spreads through burn units varies depending on habitat and weather conditions. Skips, or ‘refugia’, within the burn extent are com- mon (Sveinson Pelc 2013). The resulting patchy con- sumption of litter layer and exposure of mineral soil allows recolonization from unburned areas. The re- sult is that most sites recently burned (<2 years) had gastropod composition similar to unmanaged sites, demonstrating rapid recolonization in this patchy ecosystem. Ray and Bergey (2015) showed that in favourable weather conditions snail communities in post-burn habitats that include leaf litter showed growth rate increases due to higher soil pH follow- ing fire. In Prairie habitat once reestablishment of the litter layer is underway, recolonization from adjacent sites such as Wet Forest patches or drainage sites (es- pecially by aquatic species) can be rapid. Grazing also contributes to the preservation of the prairie ecosystem mosaic by limiting the spread of woody species and the expansion of forest. In the Tall Grass Prairie Preserve grazing had a significant ef- fect on snail community composition only at Prairie sites where vegetation structure was destroyed. At other sites, grazing intensity (frequency, number of animals, length of grazing period, and their combina- tions) may be low enough to keep disturbance below a threshold and to maintain snail species composition. When formerly managed grassland was abandoned in Romania, open-habitat gastropod species decreased (Cremene eft al. 2005). However, grazing intensity negatively influenced the snail fauna in Swiss grass- lands, independent of livestock species (Boschi and Baur 2007). Different mechanisms involved in graz- ing may affect the snail community. The choice of food plants by livestock may impact seed disper- sal and therefore plant composition, affecting food sources and micro-habitat for snails. Also, trampling may affect snail survival directly, or indirectly by de- stroying micro-habitat (Fischer et al. 1996; Rook et al. 2004). In the Tall Grass Prairie Preserve the struc- ture of mounds of grass litter in Wet Meadow is less important than in Prairie sites. Aquatic species might take more advantage of long periods of flooding in the former. Also, drainages are usually wet and pro- vide a species pool for colonizing wet meadows af- NICOLAI ET AL.: GASTROPODS IN TALL GRASS PRAIRIE 321 ter periods of drought. Dry-habitat species were ab- sent from the Wet Meadow, however some terrestrial gastropods, such as Fat Hive (Euconulus polygyratus (Pilsbry, 1899)) and Amber Glass (Nesovitrea elect- rina (Gould, 1841)) are adapted to both moist and dry habitats and, were present, in Wet Meadow. Two Wet Meadow sites were distinct from all others due to a high abundance of aquatic species which may be the result of a particular flooding regime. Management recommendations for grasslands in general include low intensity burns that preserve the organic litter layer. Intervals between burns of >5 years (Kiss and Magnin 2006) and >15 years (Nekola 2002) have been recommended to allow for restoration of the gastropod community. However, it is not clear if an interval >15 years would apply in the MTGPP sys- tem, where fire rotation historically has ranged 3-6 years (Hamel et a/. 2006) and is currently 5—6 years. Unfortunately, the gastropod community composition prior to this management strategy is unknown. First Nation fire management was also frequent (<5 years; Lewis 1985). Our observations suggest that short burn intervals have low impact when habitat is patchy, and gastropods can easily recolonize from adjacent un- burned areas. In European grasslands, Boschi and Baur (2007) advise extensive grazing. Independent of livestock species, the number of livestock present and the duration of grazing has an impact on the gastropod community. Because there can be an interaction be- tween different management methods (Damhoureyeh and Hartnett 1997), it is difficult to predict the effect of the highly diverse fire-grazing management combina- tions on gastropod communities in the different habi- tats of the tall grass prairie system. Mounds and leaf litter seem to be important for populations of gastropods to recover after burns. Leaf litter supplementation may be a management option. The gastropod fauna of Manitoba is poorly known and there is little information on the terrestrial mol- luscs of the Canadian prairie ecosystem. This study increases our knowledge concerning the range of V. parvula, P. minutissimum, Fine-ribbed Striate (Striatura milium (Morse, 1859)), E. cf. praticola, and Suboval Ambersnail (Mediappendix cf. vermeta (Say, 1829)), all recorded for the first time in Manitoba dur- ing our study. Vertigo milium, reported previously in Canada only from a few sites in Ontario and one site in Manitoba (Nekola and Coles 2010) is ranked as Nationally Imperilled to Vulnerable (N2N3, CESCC 2016). Galba Schrank, 1803 sp. could not be identi- fied to species because of taxonomic issues, and un- common grassland species/subspecies within the genus, recorded previously in Alberta (Boag and Wishart 1982) are poorly known. 322 Acknowledgements We would like to thank the Nature Conservancy of Canada (NCC) for funding this project and Mhairi McFarlane from NCC for advice on the survey de- sign. Information on habitats and land management history was jointly developed and shared by Manitoba Tall Grass Prairie Preserve Management Committee member organizations (Manitoba Sustainable De- velopment, Nature Manitoba, Environment and Cli- mate Change Canada, Manitoba Habitat Heritage Corporation, and NCC). We acknowledge also Brent Sinclair for hosting the work on the collec- tion in his lab at the University of Western Ontario (UWO) as well as Jennifer Ho, undergraduate stu- dent at UWO, for help in sorting the collection. Many thanks also to Jeremy de Waard at the Biodiversity Institute of Ontario for including voucher specimens in the Barcode of Life Database (BOLD), Randall D. Mooi and Janis Klapecki at the Manitoba Museum for curating dry voucher specimens, and Valérie Briand at the Université Rennes 1, for literature search and formatting. This manuscript was vastly improved by the editing of Associate Editor Don McAlpine. Literature Cited Austin, J.E., and D.A. Buhl. 2013. Relating Yellow Rail (Coturnicops noveboracensis) occupancy to habitat and landscape features in the context of fire. Waterbirds 36: 199-213. https://doi.org/10.1675/063.036.0209 Baur, B., C. Cremene, G. Groza, A.A. Schileyko, A. Baur, and A. Erhardt. 2007. Intensified grazing affects en- demic plant and gastropod diversity in alpine grasslands of the Southern Carpathian mountains (Romania). Biologia 62: 438—445. https://doi.org/ 10.2478/s11756-007-0086-4 Becker, D.A. 1989. Five years of annual prairie burns. Pages 163-168 in Proceedings of the Eleventh North American Prairie Conference. Prairie Pioneers: Ecology, History and Culture. 7-11 August 1988, Lincoln, Nebraska. Edited by T.B. Bragg and J. Stubbendieck. University of Nebraska Printing, Lincoln, Nebraska, USA. Accessed 10 January 2020. http://digital.library.wisc.edu/1711.dl/ EcoNatRes.NAPC11. Bellido, A. 1987. Field experiment about direct effect of a heathland prescribed fire on microarthropod commu- nity. Revue d’Ecologie et de Biologie du Sol 24: 603-633. Bleho, B.I., N. Koper, C.L. Borkowsky, and C.D. Hamel. 2015. Effects of weather and land management on the Western Prairie Fringed-orchid (Platanthera praeclara) at the northern limit of its range in Manitoba, Canada. American Midland Naturalist 174: 191-203. https://doi. org/10.1674/0003-0031-174.2.191 Boag, D.A., and W.D. Wishart. 1982. Distribution and abundance of terrestrial gastropods on a winter range of bighorn sheep in southwestern Alberta. Canadian Jour- nal of Zoology 60: 2633-2640. https://do1.org/10.1139/z 82-338 Boschi, C., and B. Baur. 2007. The effect of horse, cat- tle and sheep grazing on the diversity and abundance of THE CANADIAN FIELD-NATURALIST Vol. 133 land snails in nutrient-poor calcareous grasslands. Basic and Applied Ecology 8: 55—65. https://doi.org/10.1016/j. baae.2006.02.003 Botkin, D.B. 1990. Discordant Harmonies: A New Ecology for the Twentieth Century. Oxford University Press, New York, New York, USA. Burkman, M.A. 1993. The use of fire to manage breeding habitat for yellow rails. M.Sc. thesis, Northern Michigan University, Marquette, Michigan, USA. British Columbia Ministry of Forests. 2008. Land snails and slugs as ecological indicators of logging practices: recommendations for adaptive management. Accessed 28 December 2019. http://www. for.gov.be.ca/hfd/library/ FIA/2008/FSP_Y083030a.pdf. Brooks, M., and M. Lusk. 2008. Fire Management and In- vasive Plants: a Handbook. United States Fish and Wild- life Service, Arlington, Virginia, USA. Buehler, D.A., A.M. Roth, R. Vallender, T.C. Will, J.L. Confer, R.A. Canterbury, S.B. Swarthout, K.V. Rosenberg, and L.P. Bulluck. 2007. Status and con- servation priorities of Golden-winged Warbler (Vermi- vora chrysoptera) in North America. Auk 124: 1439- 1445. https://doi.org/10.1093/auk/124.4.1439 CESCC (Canadian Endangered Species Conservation Council). 2016. Wild Species 2015: The General Status of Species in Canada. National General Status Working Group. Accessed 10 January 2020. http://www.registr elep-sararegistry.gc.ca/virtual_sara/files/reports/Wild %20Species%202015. pdf. Clarke, A.H. 1981. The Freshwater Molluscs of Canada. National Museum of Natural Sciences. National Mu- seum of Canada, Ottawa, Ontario, Canada. Cremene, C., G. Groza, L. Rakosy, A.A. Schileyko, A. Baur, A. Erhardt, and B. Baur. 2005. Alterations of steppe-like grasslands in Eastern Europe: a threat to regional biodiversity hotspots. Conservation Biology 19: 1606-1618. https://doi.org/10.1111/j.1523-1739.2005.00 084.x Damhoureyeh, S.A., and D.C. Hartnett. 1997. Effects of bison and cattle on growth, reproduction, and abundances of five Tall Grass Prairie forbs. American Journal of Botany 84: 1719-1728. https://doi.org/10.2307/2446471 Damhoureyeh, S.A., and D.C. Hartnett. 2002. Variation in grazing tolerance among three tallgrass prairie plant species. American Journal of Botany 89: 1634-1643. https://doi.org/10.3732/ajb.89.10.1634 Davis, M.A., D.W. Peterson, P.B. Reich, M. Crozier, T. Query, E. Mitchell, J. Huntington, and P. Bazakas. 2000. Restoring savanna using fire: impact of the breed- ing bird community. Restoration Ecology 8: 30-40. https://doi.org/10.1046/j.1526-100x.2000.80005.x Dvorakova, J., and M. Horsak. 2012. Variation of snail assemblages in hay meadows: disentangling the predic- tive power of abiotic environment and vegetation. Malacologia 55: 151-162. https://doi.org/10.4002/040. 055.0110 Fischer, S.F., P. Poschlod, and B. Beinlich. 1996. Ex- perimental studies on the dispersal of plants and ani- mals on sheep in calcareous grasslands. Journal of Applied Ecology 33: 1206-1222. https://doi.org/10.23 07/2404699 2019 Forsyth, R.G. 2004. Land snails of British Columbia. Royal BC Museum Handbook. Royal British Columbia Museum, Victoria, British Columbia, Canada. Forsyth, R.G. 2005. Terrestrial Gastropods of the Upper Fraser Basin of British Columbia. Living Landscapes, Royal British Columbia Museum, Victoria, British Columbia, Canada. Accessed 28 December 2019. https:// royalbcmuseum.bc.ca/exhibits/living-landscapes/upper fraserbasin/ufb_snails/UFB-Snails. pdf. Forsyth, R.G., and M.J. Oldham. 2016. Terrestrial mol- luscs from the Ontario Far North. Check List 12: 1881. https://doi.org/10.15560/12.3.1881 Glaser, A. 2012. America’s Grasslands Conference: status, threats, and opportunities. /n Proceedings of the Ist Bi- ennial Conference on the Conservation of America’s Grasslands. August 15—17, 2011, Sioux Falls, South Da- kota. National Wildlife Federation and South Dakota State University, Washington, DC and Brookings, South Dakota, USA. Gottesfeld, L.M.J. 1994. Aboriginal burning for vegeta- tive management in northwestern British Columbia. Human Ecology 22: 171-188. https://doi.org/10.1007/ BF02169038 Grant, T.A., E.M. Madden, T.L. Shaffer, and J.S. Dock- ens. 2010. Effects of prescribed fire on vegetation and passerine birds in northern mixed-grass prairie. Journal of Wildlife Management 74: 1841-1851. https://doi.org/ 10.2193/2010-006 Hamel, C.D., R. Reisz, G. Fortney, R. Jones, and D. Pie- truszewski. 2006. Conservation Area Plan for the Tall- grass Aspen Parkland. Nature Conservancy of Canada, Manitoba Region, Winnipeg, Manitoba, Canada / The Nature Conservancy, Karlstad Field Office, Karlstad, Minnesota, USA. Harper, M.G., C.H. Dietrich, R.L. Larimore, and P.A. Tessene. 2000. Effects of prescribed fire on prairie ar- thropods: an enclosure study. Natural Areas Journal 20: 325-335. Hettenbergerova, E., M. Horsak, R. Chandran, M. Hajek, D. Zeleny, and J. Dvorakova. 2013. Patterns of land snail assemblages along a fine-scale moisture gra- dient. Malacologia 56: 31—42. https://doi.org/10.4002/ 040.056.0227 Horsak, M., and M. Chytry. 2014. Unimodal latitudinal pattern of land-snail species richness across northern Eurasian lowlands. PLoS ONE 9: e104035. https://doi. org/10.1371/journal.pone.0104035 Jordan, S.F., and S.H. Black. 2012. Effects of forest land management on terrestrial mollusks: a literature re- view. USDA Forest Service, Region 6 & USDI Oregon/ Washington Bureau of Land Management, Portland, Oregon, USA. Accessed 10 January 2020. https://www. fs.fed.us/r6/sfpnw/issssp/documents2/cpt-ig-effects-to- mollusks-bibliography-2012-02.docx. Kaufman, D.W., E.J. Finck, and G.A. Kaufman. 1990. Small mammals and grassland fires. Pages 46-80 in Fire in North American Tall Grass Prairies. Edited by S.L. Collins and L.L. Wallace. University of Oklahoma Press, Norman, Oklahoma, USA. Kiss, L., and F. Magnin. 2003. The impact of fire on some Mediterranean land snail communities and patterns of NICOLAI ET AL.: GASTROPODS IN TALL GRASS PRAIRIE 323 post-fire recolonization. Journal of Molluscan Studies 69: 43-53. https://doi.org/10.1093/mollus/69.1.43 Kiss, L., and F. Magnin. 2006. High resilience of Me- diterranean land snail communities to wildfires. Bio- diversity and Conservation 15: 2925-2944. https://doi. org/10.1007/s10531-005-3430-4 Klaus, N.A., S.A. Rush, T.S. Keyes, J. Petrick, and R.J. Cooper. 2010. Short-term effects of fire on breeding birds in southern Appalachian upland forests. Wilson Journal of Ornithology 122: 518-531. https://doi.org/10. 1676/09-105.1 Knapp, A.K., J.M. Blair, J.M. Briggs, S.L. Collins, D.C. Hartnett, L.C. Johnson, and A.G. Towne. 1999. The keystone role of bison in North American tallgrass prai- rie: bison increase habitat heterogeneity and alter a broad array of plant, community, and ecosystem pro- cesses. BioScience 49: 39-50. https://doi.org/10.1525/ bisi.1999.49.1.39 Knapp, E.E., B.L. Estes, and C.E. Skinner. 2009. Ecological effects of prescribed fire season: a litera- ture review and synthesis for managers. USDA General Technical Report PSW-GTR-224. U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station, Albany, California, USA. Accessed 10 Janu- ary 2020. https://www.fs.fed.us/psw/publications/docu ments/psw_gtr224/, Legendre, P., and L. Legendre. 2007. Numerical Ecology. Elsevier, Amsterdam, The Netherlands. Lewis, H.T. 1985. Why Indians burned: specific versus general reasons. /n Proceedings of Symposium and Wil- derness Fire, Missoula, Montana, Nov 15-18, 1983. GTR- INT-182. Edited by J.E. Lotan, B.M. Kilgore, W.C. Fischer, and R.W. Mutch. USDA Forest Service, Inter- mountain Forest and Range Experiment Station, Ogden, Utah, USA. Ludwick, T.J., and R.K. Murphy. 2006. Fire history, pas- serine abundance, and habitat on a North Dakota drift plain prairie. Prairie Naturalist 38: 1-11. Lydeard, C., R.H. Cowie, W.F. Ponder, A.E. Bogan, P. Bouchet, S.A. Clark, K.S. Cummings, T.J. Frest, O. Gargominy, D.G. Herbert, R. Hershler, K.E. Perez, B. Roth, M. Seddon, E.E. Strong, and F.G. Thompson. 2004. The global decline of nonmarine mol- lusks. Bioscience 54: 321-330. https://doi.org/10.1641/ 0006-3568(2004)054[0321 :tgdonm]2.0.co;2 Madden, E.M., A.J. Hansen, and R.K. Murphy. 1999. Influence of prescribed fire history on habitat and abun- dance of passerine birds in northern mixed-grass prai- rie. Canadian Field-Naturalist 113: 627-640. Accessed 28 December 2019. https://biodiversitylibrary.org/page/ 34235497. Minnesota Department of Natural Resources. 2005. Field Guide to the Native Plant Communities of Min- nesota: The Prairie Parkland and Tallgrass Aspen Parklands Provinces. Ecological Land Classification Program, Minnesota County Biological Survey, and Natural Heritage and Nongame Research Program, Minnesota Department of Natural Resources, St. Paul, Minnesota, USA. Nekola, J.C. 2002. Effects of fire management on the rich- ness and abundance of central North American grass- 324 land land snail faunas. Animal Biodiversity and Con- servation 25: 53-66. Nekola, J.C. 2005. Geographic variation in richness and shell size of eastern North American land snail com- munities. Records of the Western Australian Museum, Supplement 68: 39-51. https://doi.org/10.18195/issn.0313- 122x.68.2005.039-051 Nekola, J.C., and B.F. Coles. 2010. Pupillid land snails of eastern North America. American Malacological Bulletin 28: 1-29. https://doi.org/10.4003/006.028.0221 Ohrtman, M.K., S.A. Clay, D.E. Clay, E.M. Mousel, and A.J. Smart. 2011. Preventing saltcedar (Jamarix spp.) seedling establishment in the Northern Prairie Pothole Region. Invasive Plant Science and Management 4: 427-436. https://do1.org/10.1614/ipsm-d-11-00012.1 Pyne, S.J. 1983. Indian fires: the fire practices of North American Indians transformed large areas from forest to grassland. Natural History 92: 6-11. R Core Team. 2008. R: a language and environment for statistical computing. R Foundation for Statistical Com- puting, Vienna. Ramette, A. 2007. Multivariate analyses in microbial ecol- ogy. FEMS Microbiology Ecology 62: 142-160. https:// doi.org/10.1111/j.1574-6941.2007.00375.x Ray, E.J., and E.A. Bergey. 2015. After the burn: factors affecting land snail survival in post-prescribed-burn woodlands. Journal of Molluscan Studies 81: 44—S0. https://doi.org/10.1093/mollus/eyu051 Reinking, D.L. 2005. Fire regimes and avian responses in the central tallgrass prairie. Studies in Avian Biology 30: 116-126. Rook, A.J., B. Dumont, J. Isselstein, K. Osoro, M.F. WallisDeVries, G. Parente, and J. Mills. 2004. Matching type of livestock to desired biodiversity out- comes in pastures—a review. Biological Conservation 119: 137-150. https://doi.org/10.1016/j.bio con.2003.11.010 Saestedt, T.R., and R.A. Ramundo. 1990. The influence of fire on belowground processes of tallgrass prairie. Pages 99-117 in Fire in North American Tall Grass Prairies. Edited by §.L. Collins and L.L. Wallace. University of THE CANADIAN FIELD-NATURALIST Vol. 133 Oklahoma Press, Norman, Oklahoma, USA. Sampson, F., and F. Knopf. 1994. Prairie conservation in North America. Bioscience 44: 418—421. Santos, X., V. Bros, and E. Ros. 2012. Contrasting re- sponses of two xerophilous land snails to fire and natu- ral reforestation. Contributions to Zoology 81: 167-180. https://doi.org/10.1163/18759866-08103004 Shimodaira, H. 2004. Approximately unbiased tests of re- gions using multistep-multiscale bootstrap resampling. Annals of Statistics 32: 2616-2641. https://doi.org/10. 1214/009053604000000823 Sveinson Pelc, J. 2013. Characterization of a wild fire event in the tall grass prairie region of Manitoba. Pages 32-— 33 in Proceedings of the Manitoba Tall Grass Prairie Preserve Research Symposium 2014. Edited by M. Pearn and C.D. Hamel. Nature Conservancy of Canada, Winnipeg, Manitoba, Canada. Swengel, A.B. 1996. Effects of fire and hay management on the abundance of prairie butterflies. Biological Con- servation 76: 73—85. https://doi.org/10.1016/0006-3207 (95)00085-2 Towne, G., and C. Owensby. 1984. Long-term effects of annual burning at different dates in ungrazed Kansas tallgrass prairie. Journal of Range Management 37: 392-397. https://doi.org/10.2307/3899622 van Klink, R., F. van der Plas, C.G.E. van Noordwijk, M.F. Wallis DeVries, and H. Olff. 2015. Effects of large herbivores on grassland arthropod diversity. Biological Review 90: 347-366. https://doi.org/10.1111/brv.12113 Vierling, K., and L. Lentile. 2006. Red-headed wood- pecker nest-site selection and reproduction in mixed ponderosa pine and aspen woodland following fire. Condor 108: 957-962. https://doi.org/10.1093/condor/ 108.4.957 Williams, G.W. 2000. Reintroducing Indian-type fire: im- plications for land managers. Fire Management Today 60: 40-48. Received 25 January 2019 Accepted 19 December 2019 The Canadian Field-Naturalist Batch spawning in five species of minnows (Cyprinidae) from Ontario, Canada NORMAN W-.S. QUINN 47 Old Troy Road, R.R. #3, Tweed, Ontario KOK 3J0 Canada; email: normq4@gmail.com Quinn, N.W.S. 2019. Batch spawning in five species of minnows (Cyprinidae) from Ontario, Canada. Canadian Field-Naturalist 133(4): 325-328. https://doi.org/10.22621/cfn.v133i4.1914 Abstract Batch spawning, the act of spawning more than once within a spawning season, is assessed in six species of minnows (Cyprinidae) from Ontario, Canada. The bimodal frequency distribution of egg size in mature specimens suggests that the following species are batch spawners: Blacknose Dace (Rhinichthys atratulus), Brassy Minnow (Hybognathus hankinsoni), Common Shiner (Luxilus cornutus), Creek Chub (Semotilus atromaculatus), and Hornyhead Chub (Nocomis biguttatus). However, there is no evidence that Northern Pearl Dace (Margariscus nachtriebi) is a batch spawner. Thus, we now have evidence that 11 of 39 cyprinid species in Ontario are batch spawners. Knowledge about the reproductive habits of these species should be integrated into the comprehensive standards for the protection of fish habitat in Ontario to ensure the sur- vival of populations. Key words: Cyprinidae; minnows; spawning; batch; Ontario Introduction Batch (or fractional) spawning is widespread among fishes (e.g., Conover 1985). The phenom- enon is defined as spawning more than once dur- ing a spawning season as opposed to spawning only once in a relatively short period, hereafter, referred to as “conventional” spawning (Conover 1985). Batch Spawning presents a problem to fisheries managers because it confounds or renders impossible any at- tempt to estimate total fecundity (e.g., Conover 1985). Batch spawning has been frequently reported in the minnows (Cyprinidae; e.g., Heins and Rabito 1986). The objective of this study was to report on the oc- currence of batch spawning in some Ontario cyprin- ids through the examination of ovaries of mature in- dividuals of six species: Blacknose Dace (Rhinichthys atratulus), Brassy Minnow (Hybognathus hankin- soni), Common Shiner (Luxilus cornutus), Creek Chub (Semotilus atromaculatus), Hornyhead Chub (No- comis biguttatus), and Northern Pearl Dace (Marga- riscus nachtriebi). Methods In 2013-2015, minnows were captured with stan- dard (40 x 20 cm) cylindrical wire traps set overnight from late April (ice out) to 30 June, a period when spawning of these fish is underway. Five of the six Species were caught in Clarke Creek (45°06'N, 77° 48'W) near Bancroft, Ontario. Hornyhead Chub was caught in an unnamed creek near Madoc, Ontario (44° 30'N, 77°39'W). Standard length and weight of fish were recorded on capture. Ovaries were removed and preserved in 10% buffered formalin. The gonadosomatic index (GSI) was calculated as ovary weight divided by to- tal weight (including ovaries). Cyprinids typically spawn with a GSI of about 10% (e.g., Abiden 1986). The approach used to determine mode of spawn- ing was based on frequency distribution of the size of eggs in ovaries. Batch spawners in or near spawn- ing condition should show a multimodal distribu- tion of egg sizes. Large, fully mature eggs should be observed in the presence of mid-sized eggs, the lat- ter representing the batch to be spawned at a later date. Conventional, one-batch spawners should show only mature eggs amid a mass of very small “recruit- ment” eggs (Conover 1985; Powles et al. 1992) to be spawned the following year. This approach has been used previously, including with cyprinids (Heins and Rabito 1986; Powles et al. 1992; Heins and Baker 1993; Wang et al. 2014); as Heins and Baker (1993: 15) state, two separate groups of developing eggs is “a profile typical of fish that produce multi- ple clutches”. Ovaries from specimens in or near spawning condition, that is, having mature eggs (as described below) were examined to determine the frequency distribution of egg sizes. The fixed ovaries were A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 325 ©The Ottawa Field-Naturalists’ Club 326 weighed to the nearest 0.01 g. A sample of the ovar- ian matrix was obtained by cutting out two small pieces, one from each ovary. Herrera and Fernandez- Delgardo (1994) and Al Saleh et al. (2012) found that the size of eggs is more or less independent of posi- tion in the ovaries of minnows. The samples were weighed (typically 0.05—0.15 g) and placed on a glass slide, covered with a drop of water, and the eggs were spread out with the flat of a scalpel. The sample was then examined under a microscope at 40x magnification and all eggs were counted and sorted into one of three size classes: 0.20—0.60 mm, 0.61-1.00 mm, and >1.00 mm. The slides had an underlying grid to help prevent double counting of eggs, and an ocular micrometer was used to meas- ure eggs when size class was not obvious. The over- all colour of eggs in each size class was noted. The size classes correspond to the three catego- ries in Powles et al. (1992) for the minnow Northern Redbelly Dace (Chrosomus eos): 1) “immature” (“recruitment” in Conover 1985), white-grey with no yolk; 2) “maturing” (or mid-sized), vitellogenic (accruing yolk) and yellow or orange; and 3) “ma- ture’, >1.00 mm and translucent, but with yellow hues. Eggs in the mature category were fully de- veloped (Conover 1985; Powles et al. 1992). No ma- ture eggs of any observed species were greater than 1.20 mm; thus, it is assumed that size at development stage of eggs of these species and that of C. eos eggs is comparable (Brassy Minnow, an exception, 1s dis- cussed below). The subsamples typically contained 150—400 eggs. An estimate of total number of eggs and number in the three size categories was made by multiplying the weight of the ovary divided by weight of subsample times eggs counted in the sub- sample. Mid-sized eggs in the presence of mature THE CANADIAN FIELD-NATURALIST Vol. 133 eggs were deemed evidence of batch spawning. To produce more precise frequency distributions for graphic illustration, eggs were counted and meas- ured a second time. The subsample was again placed under the microscope and the diameter of 100 eggs measured with an ocular micrometer. To avoid bias, eggs were measured in the order of appearance in the field of view while the slide traversed the field of view. Distorted and ovoid eggs were quite common, but only round eggs were measured. Results Most mature female Creek Chub, Common Shi- ner, Blacknose Dace, and Hornyhead Chub had hun- dreds of mid-sized eggs in the presence of mature eggs, supporting the hypothesis that they are batch spawners (Table 1). All Brassy Minnow specimens had relatively small eggs. The 12 Brassy Minnow fe- males (caught between 4 May and 24 June in all three years) had GSI >10% and hundreds of vitellogenic eggs, but none >1.00 mm. However, five females had bimodal frequency distributions of egg size (Figure 11). Thus, Brassy Minnow appears also to be a batch spawner. Northern Pearl Dace is the anomaly in this group; the four mature females had essentially all eggs in the mature category (Figure lo,p) with negli- gible immature or mid-sized eggs. With this limited evidence, Northern Pearl Dace appears to be a con- ventional spawner. Figure 1 shows selected frequency distributions of egg size (from the 100 measured eggs per speci- men). The histograms were selected as typical of pat- terns observed for each species. Note that most (ex- cept for Northern Pearl Dace) show mid-sized eggs in the presence of mature (>1.00 mm) eggs. TABLE 1. Gonadosomatic index (GSI) and egg-size distribution in mature females of six Ontario cyprinids. No. mature Mean standard females | ength, cm Creek Chub 29 11.05 (Semotilus atromaculatus) Common Shiner 22 8.40 (Luxilus cornutus) Blacknose Dace 28 7.46 (Rhinichthys atratulus) Brassy Minnow 12 7.74 (Hybognathus hankinsoni) Hornyhead Chub 8 9.52 (Nocomis biguttatus) Northern Pearl Dace 4 7.90 (Margariscus nachtriebi) Mean GSI Egg-size a No. (%) of fish SAE 0) ee with mid-sized eggs aR 9.78 29 (100) 2603/959/107 (100/36.8/4.1) 10.56 22 (100) 1587/427/480 (100/26.9/30.3) 12.95 25 (89) 1440/245/535 (100/17.0/37.2) 10.76 = 3294/0/1028 (100/0/31.2) 14.31 8 (100) 2560/723/850 (100/28.2/33.2) 15.87 0) 775/757/28 (100/97.7/3.6) *Means of total no. eggs/mature eggs/mid-sized eggs. Mature eggs >1 mm, mid-sized 0.6—1.0 mm. TSize categories of eggs of Brassy Minnow are an exception (see text for explanation). 2019 QUINN: BATCH SPAWNING IN ONTARIO CYPRINIDS 327 a b c d 40 40 cee 50 20 | | 20 | | | 30 | ny) 0 0 ft) 0.5 >1.0 0.5 >1.0 0.5 >1.0 0.5 >1.0 é f g h 40 40 40 40 ie 20 20 20 £ S 0 i) ! i) 0 oO 0.5 >1.0 0.5 >1.0 0.5 >1.0 0.5 >1.0 fa) : = i j k I BD (9 40 40 40 40 20 20 20 20 ny) 0 0 ) 0.5 >1.0 0.5 >1.0 0.5 >1.0 0.5 >1.0 m n oO p 40 20 100 100 ° | “alll | °| | *| | ny) 0 i) : i) 0.5 >1.0 0.5 >1.0 0.5 >10 0.5 >1.0 No. of eggs Figure 1. Selected egg-size distributions for six Ontario cyprinids. a—d: Creek Chub (Semotilus atromaculatus), e—g: Common Shiner (Luxilus cornutus), h-j: Blacknose Dace (Rhinichthys atratulus), k and |: Brassy Minnow (Hybognathus hankinsonii), m and n: Hornyhead Chub (Nocomis biguttatus), 0 and p: Northern Pearl Dace (Margariscus nachtriebi). Discussion Batch spawning is reported frequently in the Cyprinidae and from locations as disparate as Spain (Herrera and Fernandez-Delgrado 1994), Iraq (Al Saleh et al. 2012), and Malaysia (Abiden 1986). Con- ventional spawning is also occasionally reported (e.g., Wang ef al. 2014). This study adds five species to the six cyprinid species already documented as batch spawners in Ontario. These other species are: Black- nose Shiner (Notropis heterolepis; Roberts et al. 2006), Bluntnose Minnow (Pimephales notatus, Gale 1983), introduced Common Carp (Cyprinus carpio, Ivanov 1976), Fathead Minnow (Pimephales promelas, Gale and Buynak 1982), introduced Goldfish (Carasius auratus, Ivanov 1971), and Northern Redbelly Dace (Powles et al. 1992). Thus, 11 of the 39 Ontario cyprin- ids have been confirmed to be batch spawners. This study suggests that Northern Pearl Dace is a conventional spawner. The evidence for batch spawn- ing reported here is indirect because direct observa- tion in the field is difficult (Conover 1985). Ontario has developed comprehensive standards for the protection of fish habitat (e.g., Anonymous 2006). For example, timing restrictions force work in water away from periods when spawning or egg development may occur (Anonymous 2006). In sys- tems with complex fish communities, this can mean that work is restricted to a few weeks in late sum- mer. However, because of batch spawning, the repro- duction of cyprinids may be prolonged; some spe- cies, for example, Fathead Minnow, spawn more than 15 times in a season (Gale and Buynak 1982). Such a prolonged spawning period suggests that even late summer restrictions may be inadequate to fully pro- tect cyprinid populations. The evolution of batch spawning has been inter- preted according to three adaptive scenarios or hy- potheses. It may be a “bet hedging” life history pattern 328 (Morrongielo et al. 2012), whereby a variable post- hatch environment and consequent unpredictable mortality of young favour a reproductive effort that is spread out temporally, thus increasing the probability of survival of the progeny. Second, Schlosser (1998) and Matthews er al. (2001) suggest that fish in con- fined environments, such as streams, extend repro- duction to minimize intraspecific competition for the developing young. Third, Coburn (1986) argues that developmental and ecological factors limit egg size to a certain minimum. Thus, fish with small adult body size, having smaller ovaries, compensate for less out- put by laying multiple clutches. More basic research and data on cyprinid repro- ductive patterns are needed to verify these adaptive hypotheses. Acknowledgements I thank Erling Holm for assistance with identifica- tion of specimens. Perce Powles provided useful com- ments on an early draft of the manuscript. Fish were collected under the authority of the Ontario Ministry of Natural Resources (OMNR) permit #1079384, and I followed OMNR Class Animal Care Protocol for Fish as directed by that agency. Literature Cited Abiden, A.Z. 1986. The reproductive biology of a tropical cyprinid, Hampala macrolepidota, from Zoo Negra lake, Kuala Lumpur, Malaysia. Journal of Fish Biology 29: 381-391. https://doi.org/10.1111/j.1095-8649.1986.tb 04954.x Al Saleh, F., V. Hammond, A. Hussein, and R. Alhaz- za. 2012. On the growth and reproductive biology of asp, Aspius vorax, population from the middle reach- es of Euphrates River. Turkish Journal of Fisheries and Aquatic Sciences 12: 149-156. https://doi.org/10.4194/ 1303-2712-v12_1_17 Anonymous. 2006. MTO/DFO/MNR protocol for protect- ing fish habitat on provincial transportation undertak- ings. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Coburn, M.M. 1986. Egg diameter variation in eastern North American minnows (Pisces: Cyprinidae): correl- ation with vertebral number, habitat and spawning be- havior. Ohio Journal of Science 86: 110-120. Accessed 25 February 2020. https://kb.osu.edu/handle/1811/23141. Conover, D.O. 1985. Field and laboratory assessment of patterns in fecundity of a multiple spawning fish: the Atlantic Silverside Menidia menidia. Fishery Bulletin 83: 331-341. Accessed 25 February 2020. https://spo. nmfs.noaa.gov/sites/default/files/pdf-content/1985/833/ conover.pdf. Gale, W.F. 1983. Fecundity and spawning frequency ofcaged bluntnose minnows, fractional spawners. Transactions of the American Fisheries Society 112: 398—402. https://doi. org/10.1577/1548-8659(1983)112<398:fasfoc>2.0.co;2 Gale, W.F., and G. Buynak. 1982. Fecundity and spawning THE CANADIAN FIELD-NATURALIST Vol. 133 frequency of the fathead minnow—a fractional spawner. Transactions of the American Fisheries Society 111: 35— 40. https://doi.org/10.1577/1548-8659(1982)111<35:fasfo t>2.0.co;2 Heins, D.C., and J.A. Baker. 1993. Reproductive biology of the Brighteye Darter, Etheostoma lynceum (Teleostei: Percidaea), from the Homochitto River, Mississippi. Ichthyological Exploration of Freshwaters 4: 11—20. Heins, D.C., and F.G. Rabito, Jr. 1986. Spawning per- formance in North American minnows: direct evidence of multiple clutches in the genus Notropis. Journal of Fish Biology 28: 343-357. https://do1.org/10.1111/j.1095- 8649.1986.tb05171.x Herrera, M., and C. Fernandez-Delgardo. 1994. The age, growth, and reproduction of Chondrostoma polylepis willkommi in a seasonal stream in the Guadalquivir River basin (southern Spain). Journal of Fish Biology 44: 11-22. https://doi.org/10.1111/j.1095-8649.1994 tb01 581L.x Ivanov, Y.N. 1971. An analysis of the fecundity and inter- mittent spawning of lake Balkhash wild carp, Cyprinus carpio. Journal of Ichthyology 11: 666—673. Ivanov, Y.N. 1976. The formation of ultimate fecundity in intermittent spawning fish with reference to Southern One Finned Greenling, Pleurogrammus aronas, and the wild goldfish, Carassius auratus. Journal of Ichthyology 16: 56-63. Matthews, W.J., K.B. Gido, and E.D. Marsh-Matthews. 2001. Density-dependent overwinter survival and growth of Red Shiners from a southwestern river. Transactions of the American Fisheries Society 130: 478—488. https://doi. org/10.1577/1548-8659(2001)130<0478:ddosag>2.0.co;2 Morrongiello, J.R., N.R. Bond, D.A. Crook, and B.M. Wong. 2012. Spatial variation in egg size and egg num- ber reflects trade-offs and bet-hedging in a freshwater fish. Journal of Animal Ecology 81: 806—817. https://doi. org/10.1111/j.1365-2656.2012.01961.x Powles, P.M., S. Finucan, M. van Haaften, and R.A. Curry. 1992. Preliminary evidence for fractional spawn- ing by the Northern Redbelly Dace, Phoxinus eos. Canadian Field-Naturalist 106: 237-240. Accessed 9 March 2020. https:/Awww.biodiversitylibrary.org/page/ 34347457. Roberts, M.E., B.M. Burr, M.R. Whiles, and V. San- tucci. 2006. Reproductive ecology and food habits of the Blacknose Shiner, Notropis heterolepis, in north- ern Illinois. American Midland Naturalist 155: 70-83. https://doi.org/10.1674/0003-0031(2006)155[0070:reafh 0]2.0.co;2 Schlosser, I.J. 1998. Fish recruitment, dispersal, and tro- phic interactions in a heterogeneous lotic environment. Oecologia 113: 260-268. https://doi.org/10.1007/s0044 20050377 Wang, J., F. Liu, X. Zhang, W.-X. Cau, H.-Z. Liu, and X. Gao. 2014. Reproductive biology of Chinese minnow, Hemiculturella sauvagei, 1888 in the Chishui River, China. Journal of Applied Ichthyology 30: 314-321. https://doi.org/10.1111/jai.12353 Received 16 October 2017 Accepted 10 January 2020 The Canadian Field-Naturalist Note High reliance on a diet of Moose (Alces americanus) by Eastern Coyotes (Canis latrans var.) in Cape Breton Highlands National Park, Nova Scotia, Canada JASON W.B. PowER!?*, MICHAEL J. BOUDREAU’, ERICH M. Muntz’, and SOREN BoNDRUP-NIELSEN! 'Department of Biology, Acadia University, Wolfville, Nova Scotia B4P 2R6 Canada ?Current address: Nova Scotia Department of Lands and Forestry, Wildlife Division, 136 Exhibition Street, Kentville, Nova Scotia B4N 4E5 Canada 3Cape Breton Highlands National Park of Canada, P.O. Box 158, Cheticamp, Nova Scotia BOE 1H0 Canada “Corresponding author: Jason. WB.Power@novascotia.ca Power, J.W.B., M.J. Boudreau, E.M. Muntz, and S. Bondrup-Nielsen. 2019. High reliance on a diet of Moose (Alces amer- icanus) by Eastern Coyotes (Canis latrans var.) in Cape Breton Highlands National Park, Nova Scotia, Canada. Canadian Field-Naturalist 133(4): 329-331. https://doi.org/10.22621/cfn.v13314.2138 Abstract Eastern Coyote (Canis latrans var.) scats were collected along transects in Cape Breton Highlands National Park, Nova Scotia, Canada, from May 2012 to August 2013 to determine diet. Based on 294 scats, Moose (Alces americanus) remains made up the highest percentage by volume in scats during fall, winter, and spring. During the summer, Moose remains were found in over 30% of scats (18% by volume), although fruit and berries were more commonly found. No other study has documented such high annual use of Moose. As there was no evidence that the consumed Moose were killed by Coyotes, presumably Coyotes scavenged Moose that had died of natural causes. Key words: Eastern Coyote; Canis latrans var.;, diet, Moose; Alces americanus; Cape Breton Highlands National Park Eastern Coyotes (Canis latrans var.) were first recorded in Cape Breton Highlands National Park (CBHNP; Figure 1), Nova Scotia, Canada, in 1980 (E.M.M. pers. obs.). A high level of coyote—-human aggressive encounters, including a human fatality (E.M.M. pers. obs.), resulted in the park initiating a study of the ecology of Coyotes within its boundaries. One aspect of this study was to understand their diet. Coyotes typically exhibit a generalist diet (Young and Jackson 1951; Bekoff 1977; Prugh 2005; Lukasik and Alexander 2011) adjusting to seasonal avail- ability of prey and other food sources (Patterson ef al. 1998; Lukasik and Alexander 2011). Food selec- tion ranges from preying on small mammals, such as rodents and lagomorphs, to large ungulates, live- stock, or pets, as well as foraging for fruit, eating garbage, and scavenging (1.e., Bowyer ef al. 1983; Fedriani et al. 2001; Lukasik and Alexander 2011). Eastern Coyotes have been known to prey effectively on adult White-tailed Deer (Odocoileus virginianus, Parker 1986; Patterson and Messier 2000) and, more recently, they have been documented killing adult Moose (Alces americanus) in Ontario (Benson and Patterson 2013). Here, we report on an unusually high reliance on a diet of Moose by Eastern Coyotes year round in CBHNP. From May 2012 to August 2013, scats were col- lected every three weeks from 21 2-km-long transects randomly selected along established paths and trails throughout CBHNP (Figure 1). Percentage by volume for each prey remain was determined using the point- frame method (Chamrad and Box 1964) after scats were washed to retain hair and bones and other hard material and dried. A Kruskal-Wallis test (R Studio, version 0.98.490; R version 3.0.2 reports y?) was used to test for differences in prey remains among calen- dar seasons. In total, 294 Coyote scats were collected along 966 cumulative km of trail transects. Dietary analy- sis of these scats indicated that Moose, fruit/berries, and Snowshoe Hare (Lepus americanus) made up the highest percentage of volume by season (Table 1). B29 ©The Ottawa Field-Naturalists’ Club 330 Nova Scotia, Canada THE CANADIAN FIELD-NATURALIST Vol. 133 National Park oa tee tm, mee, wee () Transect ae km A Moose carcass —— Cabot Trail me Park trail Figure 1. Locations of scat transects and Moose (Alces americanus) carcasses found between May 2012 and August 2013 in Cape Breton Highlands National Park. Overall, Moose was the most common food by vol- ume found in Coyote scats (over 70% during spring and winter), followed by fruit/berries (fall 29%, sum- mer 56%) and Snowshoe Hare (winter 25%). Small mammals (23% fall), birds (9% fall), and deer (8% spring) were less common. While opportunistically back tracking Coyotes (57 tracks for a total of 109 km) in winter, we found 19 Moose carcasses, all female ranging from 1.5 to 10.5 years old (aged by analyzing tooth pattern of lower jaw; Figure 1). None of the carcasses was lo- cated on or near scat transects. These carcasses had been scavenged by Coyotes; there was no evidence that Coyotes had killed any of these Moose. These results indicate that Eastern Coyotes in CBHNP have a generalist diet with a high reliance on Moose. Studies in eastern Maine (Litvaitis and Harrison 1989), northwestern Wyoming (Dowd and Gese 2012), southeastern Quebec (Richer et al. 2002), western Maine (Major and Sherburne 1987), and New Brunswick (Dumond ef a/. 2001) have found Moose to make up a smaller proportion of the diet of Coyotes. Only in eastern Quebec did Boisjoly ef al. (2010) report a high frequency of 51% Moose in scats. Our study documents the highest percentage by vol- ume of Moose in scats of Eastern Coyotes in CBHNP during the winter (71%). At the time of this study, Moose were likely the TABLE 1. Analysis of prey remains identified from 294 scats of Eastern Coyote (Canis latrans var.) collected on trail tran- sects in Cape Breton Highlands National Park, Nova Scotia, from May 2012 through August 2013. % prey by volume (mean + SD) in each season* Dietary remains Fall Winter Spring Summer e P (n= 40) (n = 80) (n= 64) (n= 110) Moose (Alces americanus) 23.3 + 39.4 71.24 44.5 70.9 + 44.5 18.0 + 35.8 78.98 0.000 White-tailed Deer (Odocoileus — Loe D3 Le 27U — 13.99 0.003 virginianus) Snowshoe Hare (Lepus americanus) 16.2 + 34.6 24.8 + 42.2 18.1 + 37.5 11.8 + 30.4 302° 0.317 Bird 8.6+ 241 0.9458 16 125 1946.4 19.76 0.000 Small mammal P29 + 30.6 Load Let 1241 12.0 + 27.6 33.49 0.000 Fruit 29.1 +419 — — 56.3+43.1 147.20 0.000 *Fall = 22 September to 20 December, winter = 21 December to 19 March, spring = 20 March to 20 June, summer = 21 June to 21 September. 2019 most biomass-rich food source available to Coyotes, especially in the highlands. Moose density in the highlands of CBHNP is typically high, with well over 1000 individuals in the park (Bridgland et al. 2007), although no estimate of Moose density was available during our study period. Gray Wolves (Canis [upus) in western Quebec were observed feeding on sin- gle Moose carcasses for up to 23 days (Messier and Créte 1985); thus, a Moose carcass could likely sus- tain a Coyote pack for several weeks as a protein- and energy-rich food source. Furthermore, less energy is likely expended scavenging a Moose carcass dur- ing winter and spring months compared with hunting small mammals. Cyclical lows of the Snowshoe Hare population during this study (E.M.M. pers. obs.) may have contributed to the primary occurrence of Moose in Coyote scats. Coyotes in CBHNP may rely on Moose carcasses because of their apparent availabil- ity and the lack of other prey, such as Snowshoe Hare, a common food source of Coyotes in other parts of Nova Scotia (Patterson et al. 1998). Literature Cited Bekoff, M. 1977. Canis latrans. Mammalian Species 79: 1-9. https://doi.org/10.2307/3503817 Benson, J.F., and B.R. Patterson. 2013. Moose (Alces al- ces) predation by eastern coyotes (Canis latrans) and eastern coyote x eastern wolf (Canis latrans x Canis lycaon) hybrids. Canadian Journal of Zoology 91: 837— 841. https://doi.org/10.1139/cjz-2013-0160 Boisjoly, D., J.P. Ouellet, and R. Courtois. 2010. Coyote habitat selection and management implications for the Gaspésie caribou. Journal of Wildlife Management 74: 3-11. https://doi.org/10.2193/2008-149 Bowyer, R.T., S.A. McKenna, and M.E. Shea. 1983. Seasonal changes in coyote food habits as determined by fecal analysis. American Midland Naturalist 109: 266-273. https://doi.org/10.2307/2425406 Bridgland, J., T. Nette, C. Dennis, and D. Quann. 2007. Moose on Cape Breton Island, Nova Scotia: 20th cen- tury demographics and emerging issues in the 21st cen- tury. Alces 43: 111-121. Accessed 5 March 2020. https:// pdfs.semanticscholar.org/67fd/c73fbd5800fe5ab87f08c 76218352f8cf374. pdf. Chamrad, A.D., and T.W. Box. 1964. A point frame for sam- pling rumen contents. Journal of Wildlife Management 28: 473-477. https://doi.org/10.2307/3798199 Dowd, J.L.B., and E.M. Gese. 2012. Seasonal variation of Coyote diet in northwestern Wyoming: implications for dietary overlap with Canada Lynx? Northwest Science POWER ET AL.: DIET OF EASTERN COYOTE 331 86: 289-299. https://doi.org/10.3955/046.086.0405 Dumond, M., M.A. Villard, and E. Tremblay. 2001. Does coyote diet vary seasonally between a protected and an unprotected forest landscape? Ecoscience 8: 301-310. https://doi.org/10.1080/11956860.2001.11682657 Fedriani, J.M., T.K. Fuller, and R.M. Sauvajot. 2001. Does availability of anthropogenic food enhance dens- ities of omnivorous mammals? An example with coy- otes in southern California. Ecography 24: 325-331. https://doi.org/10.1111/).1600-0587.2001.tb00205.x Litvaitis, J.A., and D.J. Harrison. 1989. Bobcat—coyote niche relationships during a period of coyote population increase. Canadian Journal of Zoology 67: 1180-1188. https://doi.org/10.1139/z89-170 Lukasik, V.M., and S.M. Alexander. 2011. Human—coyote interactions in Calgary, Alberta. Human Dimensions of Wildlife 16: 114-127. https://doi.org/10.1080/10871209. 2011.544014 Major, J.T., and J.A. Sherburne. 1987. Interspecific rela- tionships of coyotes, bobcats, and red foxes in western Maine. Journal of Wildlife Management 51: 606-616. https://doi.org/10.2307/3801278 Messier, F., and M. Créte. 1985. Moose—wolf dynam- ics and the natural regulation of moose popula- tions. Oecologia 65: 503-512. https://doi.org/10.1007/ BF00379664 Parker, G.R. 1986. The seasonal diet of Coyotes, Canis latrans, in northern New Brunswick. Canadian Field- Naturalist 100: 74-77. Accessed 5 March 2020. https:// www. biodiversitylibrary.org/page/28072149. Patterson, B.R., L.K. Benjamin, and F. Messier. 1998. Prey switching and feeding habits of eastern coyotes in relation to snowshoe hare and white-tailed deer dens- ities. Canadian Journal of Zoology 76: 1885-1897. https://doi.org/10.1139/z98-135 Patterson, B.R., and F. Messier. 2000. Factors influencing killing rates of White-tailed Deer by coyotes in eastern Canada. Journal of Wildlife Management 64: 721-732. https://doi.org/10.2307/3802742 Prugh, L.R. 2005. Coyote prey selection and community stability during a decline in food supply. Oikos 110: 253— 264. https://doi.org/10.1111/j.0030-1299.2005.13478.x Richer, M.C., M. Créte, J.P. Ouellet, L.P. Rivest, and J. Huot. 2002. The low performance of forest versus rural coyotes in northeastern North America: inequality be- tween presence and availability of prey. Ecoscience 9: 44-54. https://doi.org/10.1080/11956860.2002.11682689 Young, S.P., and H.H.T. Jackson. 1951. The Clever Coyote. Stackpole Company, Harrisburg, Pennsylvania, USA. Received 31 January 2019 Accepted 18 December 2019 The Canadian Field-Naturalist A review of the historical and current status of American Beaver (Castor canadensis) on Prince Edward Island, Canada ROSEMARY CURLEY'”*, DAviD L. KEENLYSIDE®, HELEN E. KRISTMANSON’, and RANDALL L. DIBBLEE!> ‘Department of Agriculture and Forestry, P.O. Box 2000, Charlottetown, Prince Edward Island C1A 7N8 Canada ?Current address: 9 Harland View Drive, Stratford, Prince Edward Island C1B 1W2 Canada ’Prince Edward Island Museum and Heritage Foundation, 2 Kent Street, Charlottetown, Prince Edward Island C1A 1M6 Canada “Intergovernmental and Public Affairs, Aboriginal Affairs Secretariat, PO. Box 2000, Charlottetown, Prince Edward Island CIA 7N8 Canada >Current address: 672 St. Catherine’s Road, Bonshaw, Prince Edward Island COA 1CO Canada “Corresponding author: rcurleypei@eastlink.ca Curley, R., D.L. Keenlyside, H.E. Kristmanson, and R.L. Dibblee. 2019. A review of the historical and current status of American Beaver (Castor canadensis) on Prince Edward Island, Canada. Canadian Field-Naturalist 133(4): 332— 342. https://doi.org/10.22621/cfn.v13314.2145 Abstract Evidence supporting the native status of American Beaver (Castor canadensis) on Prince Edward Island (PEI) before European contact in 1534 has yet to be established; however, the postglacial and archaeological records have not previ- ously been reviewed in this context. We demonstrate the coincidence of a land bridge between the mainland and PEI and the occurrence of beavers in the region dating between 9500 and 5000 BP (before present, with present defined as 1950). We provide an archaeological record of 14 beaver incisors in six locations, deposited between 500 and 1650 AD and also show that beavers could swim to PEI. Based on this evidence, we conclude that beavers were native to the province. The current population, originating via reintroductions from New Brunswick, has populated much of the available habitat and engendered considerable controversy. Key words: American Beaver; Castor canadensis; status; dispersal; Prince Edward Island; history; archaeology Introduction The historical status of mammals of Prince Edward Island (PEI) during the early years of European contact and settlement has been exten- sively researched by Sobey (2007). While admitting that American Beaver (Castor canadensis) may have been extirpated from PEI before 1700, he believed that the evidence supporting native status for this Species in the province was remarkably weak. In a more recent review of mammal status in the region, the beaver has been described as extirpated from PEI and reintroduced (Forbes et a/. 2010), but with- out supporting details. With regard to the existing population, Cameron (1958) noted the reintroduction of beavers to PEI from Algonquin Park, Ontario, in 1908-1910 and Dibblee (1994) found contemporary records of importation by private individuals; how- ever, both efforts were unsuccessful because of un- regulated trapping. The current PEI population origi- nated via reintroductions from New Brunswick (NB) after 1940 (Cameron 1958; Dibblee 1994). It is perhaps the uncertainty about beaver status on PEI that led the Atlantic Salmon Federation to la- bel the beaver a non-native species and to call for its removal from several eastern PEI rivers as part of a conservation strategy for Atlantic Salmon (Salmo salar, Guignion 2009). Similarly, Cairns et al. (2010) suggested that the beaver’s effect on Atlantic Salmon associated with river damming could be classified as (negatively) anthropogenic, rather than natural. However, management decisions about beavers on PEI should be based on a comprehensive understand- ing of the historical and current status of the species. When assessing mammal colonization of islands, Mazza et al. (2013) suggest considering palaeonto- logical, climate and sea-level evidence, characteris- tics and behaviour of the species, the historical re- cord, and the primary source of information, the fossil record. To determine whether the beaver is an alien species on PEI or a native mammal that was ex- tirpated and reintroduced, we reassess the historical evidence from Sobey (2007) and others, as well as 332, ©The Ottawa Field-Naturalists’ Club 2019 the post-glaciation history and geography of the prov- ince, and archaeological materials. Finally, we pre- sent the current status of the beaver in the province. Methods We reviewed both historical and scientific litera- ture for references to beavers and beaver habitat on PEI and elsewhere. Our search included local his- tory documents, scientific literature, government re- ports, archaeological reports, and collections at the Canadian Museum of History and Parks Canada (Halifax). Sobey’s (2002, 2006a,b, 2007) wildlife his- tory research covered much of the historical account. Archaeological field research was undertaken by D.L.K. from 1980 to 2008 and by H.E.K from 2009 to 2018. Dating of beaver teeth from these archaeo- logical studies was based on the site characteristics, cultural associations, and, more specifically, radio- carbon dating of associated charcoal in shell mid- dens at South Lake and Greenwich. An incisor from MacMillan’s Point was radiocarbon dated through accelerator mass spectrometry at Beta Analytic Testing Laboratory (Miami, Florida, USA). Dr. Fran- ces Stewart, a peer-recognized leading zooarchaeol- ogist for eastern Canada with an extensive reference collection of skeletons, determined the species iden- tity of bones at George Island. The distance beavers may have swum or rafted to get to islands was calculated using Google Earth (Keyhole, Inc., Mountain View, California, USA). Where island hopping was possible, the longest open- water swim using islands was calculated, as well as the straight-line distance through water. Monitoring of beaver populations from 1972 to 2007 was conducted by R.L.D. while employed by the PEI government. Areas of beaver-influenced wet- lands were delineated and measured on aerial photos from 1990, 2000, and 2010 captured at a 1:17 500 scale (PEICLUI 2010). Beaver dams and the triangu- lar flooded areas behind them are readily recognized at this scale. Wetland sizes ranged from 0.1 to 240 ha. In 1990 and 2000, coverage of PEI included the use of infrared photography. Both active and inac- tive beaver dams were delineated on the 1990 pho- tographs. Each photo was overlain with a same-scale transparent map showing roads and streams. These lines were then transferred to the map and digitized to create the first vector-based PEI wetland inventory. In 2000, the analog film was then scanned and the resultant imagery used to create complete orthorec- tified imagery of PEI. Using the orthomaps and ex- isting digital 1990 inventory, the 2000 PEI wetland inventory, including beaver dams, was incorporated into a province-wide digital land use inventory. In 2010, digital imagery was acquired in both col- CURLEY ET AL.: BEAVERS ON PEI 333 our and colour-infrared with 40-cm resolution. Soft- copy photogrammetry was used to generate the PEI 2010 Land Use Inventory in which the same interpre- tation parameters were applied. Results Historical evidence of Castor canadensis Cameron (1958: 45) listed the beaver as a native mammal in PEI, taking as proof “the presence of bea- ver tooth marks on sticks found in peat bogs”, but pre- sented no further details. Sobey (2007) acknowledged no firsthand account of beavers in the PEI historical record since French settlement in 1721. There are two early French reports. In 1721, Denys de La Ronde stated that there were no beavers in Ie Saint-Jean (as it was then named; Sobey 2007). However, footnoted evidence reveals that Pere René-Charles Breslay, who lived in Ile Saint-Jean from 1721 to 1723, took eight beaver skins to France (Sobey 2006a). Beavers were included in an 1802 shipment of pelts and an 1808 list of pelt prices from PEI. It is notable that credi- ble mammal listers (e.g., Johnston 1822; MacGregor 1828) in the early 1800s failed to include the bea- ver for the island. Peter Sinott emigrated to PEI in 1821 and stated in 1876 that the beaver had been pre- sent when he was younger (Sobey 2006b), whereas, in the late 19th century, the opinion of a permanent resident, Sutherland (1861), and the visiting Rowan (1876) indicated an absence of beavers (Sobey 2007). A Summerside Journal article by Marks (1900) re- lated the words of an old gentleman that the last bea- ver he saw had been killed 40 years earlier, ca. 1860, and the 31 October 1916 Charlottetown Guardian reported that the late Professor Caven of Prince of Wales College, Charlottetown, had found traces of beaver dams on the Dunk River (Dibblee 1994). Natu- ralist Francis Bain (1890) reiterated that remains of beaver dams could still be seen. Glacial and post-glacial history, geography, and dispersal The current PE] mammalian fauna arrived after glaciers retreated about 11 000 BP (before present, with present defined as 1950; Shaw ef al. 2006). Although overland access to PEI was necessary for some species and facilitated by a land connection that was in place from 9500 to 5000 years BP across what is now Northumberland Strait (Kranck 1972; Shaw et al, 2002), it is unlikely that the now-flooded strait is a barrier to beaver dispersal (Table 1). On PEI, sev- eral specimens of River Otter (Lontra canadensis), previously regarded as extirpated, have been col- lected since 2016 (G. Gregory pers. comm. 20 June 2019) including a juvenile that was whelped on PEI, although at least some have swum or travelled on ice across 13+ km of marine waters, identical to distan- 334 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE I. Unassisted occupation of islands by American Beaver (Castor canadensis) and the shortest straight-line distances by water, measured using Google Earth. Island and jurisdiction Water body Nueva Island, Chile South of Beagle Channel Lennox Island, Chile South of Beagle Channel Admiralty Island, Alaska Stephens Channel Isle Royale, Michigan Lake Superior Grand Manan Island, NB_ Bay of Fundy Newfoundland, NL Cape Breton Island, NS Prince Edward Island George Island, PEI Strait of Belle Isle or other route Strait of Canso Northumberland Strait Malpeque Bay Distance to mainland or island (km) Salinity Reference 6.5—11.5* Marine Anderson et al. 2009 6 Marine Anderson et al. 2009 SES Marine MacDonald and Cook 1996 23 Fresh Mech 1966 11 Marine Ingersoll and Gorham 1978 17+ Marine Cameron 1958 1 Marine Cameron 1958 13 Marine Cameron 1958; this study 1 Marine This study Note: NL = Newfoundland and Labrador, NB = New Brunswick, NS = Nova Scotia, PEI = Prince Edward Island. *With possible island hopping, the longest open-water swim using islands is shown as well as the shortest possible distance by water. ces regularly swum by River Otters in marine wat- ers of Alaska (Blundell et a/. 2002). The beaver and the otter were regarded as equally effective dispers- ers in colonizing insular Newfoundland and Labrador (Dodds 1983), and possibly not via the narrow but tur- bulent Strait of Belle Isle (Cameron 1958). Beavers from Minnesota, USA, have dispersed to Isle Royale, Michigan, USA, at least twice across 23 km of freshwater in recent times (Mech 1966), far ex- ceeding the 13 km that a beaver would need to swim to PEI. American Beavers commonly occupy inshore islands in Canada (Naughton 2012), and a rejuvenated beaver population in Newfoundland arrived at sev- eral smaller offshore islands in the mid-1900s (Dodds 1983). R.C. has twice seen a beaver swimming along the shore in coastal areas of PEI, as well as a beaver dam constructed across a coastal salt marsh. Beavers are well adapted to an aquatic environment, and they have several features that also protect them in marine waters. They breathe only through the nose and can prevent accidental swallowing of water. When they are underwater, flaps close off their nose and ears and a membrane protects the eyes (Naughton 2012). They are also buoyant and predisposed to enter the water, characteristics that enable colonization of islands (Mazza et al. 2013). On Cape Breton Island, Nova Scotia (NS), beaver presence was recorded by 9500 years BP (Gorham ef al. 2007); thus, they were present in the Maritimes during the 4000-year period when the extensive land connection between PEI and NB and NS was in place (Kranck 1972; Shaw et al. 2002). Archaeological record of beavers on Prince Edward Island Sobey (2007) acknowledged the beaver inci- sor excavated at South Lake by Keenlyside (1982, 1983) but did not look for other records. The archae- ological record for the Maritimes has been evaluated (Murphy and Black 1996). Because of the great influ- ence of coastal erosion and relative rise in sea level, many possible sites of older coastal encampments of Indigenous peoples have long since disappeared un- der water. As well, pre-contact shell middens were systematically spread on the land by PEI farmers to counteract soil acidity and, thus, their contents were plundered and/or dispersed (Gesner 1846). There are 14 archaeological collections of beaver material from six sites, all with deposition dates af- ter the postglacial flooding of Northumberland Strait (Table 2). Incisors are the most easily identified bea- ver remains and, thus, are often noted immediately when found, as at the Sutherland site, Greenwich (CcCp-7; Keenlyside 2002). Faunal remains of beaver are currently known from four prominent archaeological sites on PEI (Figure 1). The MacDonald site (CcCm-12), located in South Lake, Kings County (Keenlyside 1982, 1983), consisted of two cultural components: a prob- able Acadian early-mid 18th century historical oc- cupation, and a second underlying late Maritime Woodland occupation dating to about 600-900 AD. Associated with the earlier indigenous occupation was a cut beaver incisor section that appears to have been used as a bit for a cutting or incising implement. No post-cranial elements were identified in the site faunal sample. The finds from the Sutherland site at Greenwich (CcCp-7) located on the north shore of St. Peters Bay, Kings County, now part of Prince Edward Island National Park (PEINP; Keenlyside 2002), in- cluded a modified incisor recovered from a test pit in a shell midden deposit, one of several found. The site revealed extensive habitation covering 2—3 ha and dates from 800 to 900 AD, a similar period as at the MacDonald site. The split incisors found at Rustico (Robinson) Island in PEINP were used as a knife by Indigenous CURLEY ET AL.: BEAVERS ON PEI 335 2019 [IOs UI Wd QE Jnoge poring ‘pray pomoyd ‘yd d1Y & Ul Y}O0} pouldyeg oJIUy & SB OSN JOF poyst -jod pure ‘punois ‘yds doop Wo Ep—pEe 9uoz ysnoyd ur ‘poyjoours pure yno puo “j1yd¢ (A]UO apIs QUO) IND pu poINnjoRIy Wd 6] :JNd WO QT ‘sUOTSNOUT o1ue1a9 Aq payep ‘uappru [JOYS YM yueq Sulposrg yueq uy daap wid g7—-¢€7 Uappruw [Joys WO1] s]JUSUUWOD, menneeceye) ‘siod UOSUBUT}SITY 6861 150) s¥a)=) 4 (0)9) OVAL uosnsJoy 99e][e@M =108-SAIIAL ‘686 PleuogT cO0C JIeMIIS “pu UeLIO/L ‘TOOT SPIsApUsAy cO0C JIeMaIS “pu UeLIO/, ‘TOOT SPIsATUsAy ‘pu uepLlopy ‘TOOT SPIsATUSAy TOOT APIsATUIIy Z86I SPISATUIy ‘pu ULTIOW “E861 ‘T8361 OpIsA[usayy syuowmnoop JOUdIOJOY ‘O08-IfCAL SYTZ ‘60TZ ‘L-d999 OWO 8Z6l ‘L-d999 OWO Z8r ‘ISP ‘L-dg99 OWO Isl ‘L-dg99 OWO eve C[-W DID OWO 90S C[-W DID OW) Joyono,, O€ + OCE OS + OLV OOT + O8ZI OOT + O8ZI OOIF 0871 OST ¥ OOTI O91 O81 O91 O81 CS ¥ 0S61/dd sea Jose poziyeulloN (OS9I-OLtI1) 06 = 09ST (O€SI-O€rI) OS = OSI (001-008) 00€ + OOIT (001-008) O0€ ¥ OOIT (001-008) O0€ ¥ OOIT (000I—00S) OSZ ¥ OSL (0001-008) 001 + 006 (0001-008) 001 + 006 (osued) dS + dV ‘pousodap sieod ayeurxoiddy [M AMd Wd W1d AW1d W1d W1d W1d 10199] [09 166l 886I-L86I 100c—000¢ 100c—000¢ S86l £66I-V66l O86l SO86I suoseos [e1oaas ayep UOT}D9TJOD Avg peoysaoa ‘WUIOd S_URT[TASe (1-1999) yled [euoneN [Ad “puv|s][ SUOSUIGOY (£-d999) yieg jeuonen Tdd “YsIMuseln “OUIS puvpoyINS (£-d999) yieg jeuonen Tdd “YosIMuseln “OUIS puvpIoyINS (£-d999) yieg Jeuonen Tdd “YosIMuselH “OUIS puvjJoyINS (L-d999) yled [euoneN [Ad “YOIMUdIH “OUIS puvpJoyINS (Z1-W999) aye’y yINos ‘ays preuoqoe~w (C1-W999) aye’T ynos ‘ays preuoqoe payday[oo uole00'T IOSIOUI IOMO'T JOSIOUT IOMO] JU pue roddn sug JOSIOUI Jo yUSWIseIy pure Josroul Jo jusWI sel} [OUIeUA JOSIOUI IOMO'T SJUSUIS IY JOSIOUI OMI, JOSIOU] JOSIOUI JO 9091 g JOSIOUI JO 3091 g oduies ‘LIOT—O861 SulInp puels] prempy cours UO BUI}IaT[OO OINSTUN,IOddoO puke SUOTAOXO [BOISOTOOBYOIC WIOIJ POUIeIGO PUL CV YS9I—OOS PoUsodap siOsIoUl (S7sUapHUDI OJSDD) JOARIG UCILIOULY JO SUIVUIOI UMOUY *7 ATAV 336 THE CANADIAN FIELD-NATURALIST Vol. 133 people (Leonard 1989; Wallace Ferguson 1989), but oc 2 _ is eee S23 es 2 not all teeth were modified. For instance, the com- DS Aas 6 5 oi Teg : : 2 |e z 28 S5 < plete incisor at a George Island (Pitawelkek) site, ® ag B58 Pe S where exploration continues, showed no modifica- : 3 225 22 |8 tions of any kind (Kristmanson 2007, 2009), nor did S & 2 8 ae a Ss. EB the MacMillan Point specimen. The latter had been S59 28 8 gS S Z| s found in an old fire pit in a plowed field about 6 km zi SS 6ER OES from the Rustico Island site (H.E.K. unpubl. data). Soonso& nase ; : 2 A rib fragment from an immature beaver, an un- ey S 5 dated phalanx, and incisor fragments were recovered ere S S =: from excavations at the Pitawelkek site on George 2 e|s Ss Es z Island. These are in general association with radio- 3 a |é = do | 2 carbon dates of ca. 700-800 AD through to the re- 9 5 z 5 5 = = cent historical period. For now, the age of the occu- a S g o a 5 pation has been cautiously extended back to at least < 0 AD based on the collection of diagnostic artifacts 5 Se Ue Be 5 Z at the Pitawelkek site and other locations on George a. | 22 é cans r 2=] <= 3 Island (Kristmanson 2019). The Malpeque site is a S |Bee 282 eel & cultivated field where it is believed that shell middens z Pome Ro ey. Bo Ss =f were spread on the land (Gesner 1846). * co s oi 3 = Current status of beaver So |S S Ss il ; : f : gH | 2 z = = The history of beaver reintroductions was in- SS |g eS £ ft vestigated by Dibblee (1994). Following the earli- E a|s 5 eb 2 8 est importations from Ontario in 1908 and from an s S es = 38 unconfirmed source in 1912, the population rose to < ss i Ve an estimated 500 beavers over several river systems. 20 = ie Z However, high fur prices in the 1920s and unregulated Su |< < < fF 2 =I trapping resulted in the animals’ disappearance. No gAg||] z z 2 © & beaver dams were detected on 1935 aerial photos. In i = = > ww &§ , ' Eo | 2 o o a vio& the late 1940s, a migratory birds protection officer, oO 4 7 % ¢ Sarl s 3 3 em A Spurgeon Jenkins, obtained beavers from NB biol- é a [> ~ = 2 5 ogist Bruce Wright, and introduced them into PEI. 3 g 8 3 Thus, Dibblee concluded that all beavers now present * 5 5 7 in PEI originated from NB. 3 |M ns ns g M § Beavers were initially introduced east of Char- Fe = 5 o lottetown, but by 1973 the population had expanded. Ti a Between 1973 and 1979, government personnel re- 4 es Me a = J a moved 32 beavers from eastern PEI where they were 3 2 S = = = 5 = considered to be a nuisance and released them into 2s He § watercourses of Prince County, where no beavers jo) me io) . f= O ep 2 ms were present. In January 1981, the first short open sea- a $ aS 8 son for trapping beavers took place in Prince County, 3 a 2 z 5) when 20 beaver “problems”, such as blocked culverts hall ee aa 3 x s ss & and flooded driveways indicated an expanded popu- eo [se 88 22 |e = 33 lation (Dibblee and Curley 1980). By 2000, the island 8 3 rm aa F a _ =. E £3& was well populated with beavers, although few were B28 2B A: aa ca2s & recorded in the hilly central portion of the island with x §s0 §s0 4s 35 Fe 32 its flashy streams (a flashy stream is one that rapidly = - Saas & collects flows from the steep slopes of its watershed ae 5 OVE’ ; 5 5 S = 5 1 Pars basin and produces flood peaks soon after the rain but S 2 2 B Y 3 g 2 5 the flow quickly subsides after the rainfall ends). This S o is ro is O88 5 v habitat is less suitable for beavers (Novak 1987), and z = | 2 QB a. 2 iM = os most are situated to the east and west on rivers with a Ala 5 =) Zar sm low gradients (Figure 1). 2019 . q “tinh, George Island e = ~_ dae? \ t Malpeque Queens. - County, - x pe CURLEY ET AL.: BEAVERS ON PEI Rustico Island 337 PEI Greenwich |. MacMillans, -’: :.. 7 Point Charlottetown- he e@ <,' e Port La Joye % Beaver incisor @ Beaver-influenced wetlands, 2010 = co i@ 12.5 25 37.5 50 kilometres Figure 1. Locations of intact or partial American Beaver (Castor canadensis) incisors and other geographic points of inter- est, Prince Edward Island, Canada, plus active and inactive beaver-influenced wetlands delineated in the Prince Edward Island Corporate Land Use Inventory (PEICLUI 2010). According to data maintained by the provincial fish and wildlife agency, from 1975 to 2015, an aver- age of 465 beavers (one per 11 km?) were trapped each year, varying from 91 in 1975 to 917 in 2011. Between 1972 and 2002, aerial surveys of index watersheds totalling 1363 km? of predominantly forested habitats were conducted by the province. Results indicate a peak of 276 active colonies in 1993 and 160 in 2002. The total wetland area of both active and inactive beaver flowages in PEI was calculated from aerial photography as 2233 ha in 1990, 3395 ha in 2000, and 5304 ha in 2010. Beavers dams are often perceived as detrimental to salmonid populations (Kemp ef al. 2012), and watershed enhancement groups express concerns re- garding Brook Trout (Salvelinus fontinalis) and At- lantic Salmon migration. Beaver numbers fluctuate inversely with fur prices and are a continuing issue for wildlife managers dealing with complaints. Many of the 24 community-based watershed groups have considered beavers in their long-term management plans, as encouraged by the PEI Beaver Management Policy (Anonymous 2011). In practice, plans often direct removal of beavers from the main stem of a river. A local watershed group continues to remove all beavers from northeastern rivers on PEI to support spawning of a unique population of Atlantic Salmon (Moore et al. 2014). Discussion Previous records of beavers In evaluating the historical evidence of PEI mam- mals, Sobey (2007) gave credence to first-hand ac- counts or records as verifying or disputing the pres- ence of various species. The acceptance of claims that beavers were not present in the province led him to explain away a considerable body of evidence that beavers may have been present. Two (Sutherland 1861 and Rowan 1876) of three mammal recorders who stated that beavers were not present produced their reports in the last half of the 19th century when the few beaver records may have been of new arriv- als from the mainland after a long period of absence. Rowan (1876), a travel writer who merely visited the province, also later stated in the same publication that beavers were extirpated from PEI. 338 These writings likely do not meet the standard of a first-hand account, nor, we assert, would the obser- vations by Denys de La Ronde, a naval officer, who spent only 13 months in PEI beginning in 1721, in- cluding time to travel to Louisbourg, Cape Breton Island (Sobey 2002). Although Denys de La Ronde visited all active PEI harbours, the French population in 1721 was perhaps 200 (Harvey 1926). Local know- ledge of wildlife would be cursory and cleared land scarce. Denys de La Ronde could not have spoken from personal knowledge of PEI beaver habitats, which would consist mainly of forested river sys- tems extending to the coast in a land mass exceeding 5000 km?. He may have obtained information from Mrkmag traders at Port la Joye, the seat of French government in Ile Saint-Jean, or from fishermen, but he did not acknowledge the source of his information. Harvey (1926) also calls into question his veracity as a reliable reporter. Novak (1987) reasoned, based on food availabil- ity, that beavers likely existed at lower densities in mature forests of the 1500s and the 1600s compared with the high beaver populations in the food-rich early successional riparian forests of today. Beavers prefer young saplings as food and only cut large trees further away from water when saplings are depleted (Gallant et al. 2004). PEI has short river systems and human travel in the pre-settlement mature for- ests dominated by American Beech (Fagus grandi- folia Ehrhart) was relatively easy, with some excep- tions (Sobey 2002). Thus, we speculate that beavers, present according to the archaeological record, were relatively accessible. They live in families of two adults and potentially three or four kits and two or three yearlings (mean group size in central Ontario is 7.5; Novak 1987), and their lodges are easily identi- fied and exploited. Fur trading began in the Maritimes in the mid- 1500s when Basque and French vessels began fish- ing for cod in the Gulf of St. Lawrence and exchang- ing goods with the Mrkmagq (Ray 1987; Cook 1993; Whitehead 1993). Nicolas Denys had an exclusive li- cense to enter into trade for fur and fish in the gulf dating from 1654, and PEI was included in his grant. A cod fisherman and now respected author, Denys (1672) recorded the presence of Basque ships in PEI waters and discussed how beaver pelts were obtained. The Mrkmaq scared beavers from their lodges in winter, clubbed or harpooned them, taking all within the colony, and also took beavers during ice-free sea- sons by draining their dams and attacking them with Spears and arrows. They met a strong economic de- mand for beaver pelts from Europeans, and extirpa- tion in PEI in the 1600s or earlier is a possibility, as noted by Sobey (2007). A reinterpretation of Sobey’s THE CANADIAN FIELD-NATURALIST Vol. 133 information discarding Denys de La Ronde’s opin- ion might also indicate that the beaver persisted into the 1800s. The decades of greatest beaver harvest in North America as a whole were 1700-1709 and 1790— 1799 (Novak 1987; Obbard et al. 1987). Archaeological record, glacial, and post-glacial history Sobey’s research (2007) does not take into ac- count the archaeological record for PEI. In 1995, ar- chaeological research on PEI constituted only 2% of published and unpublished primary studies in the re- gion (Murphy and Black 1996). Studies are needed from inland freshwater sites where beavers might naturally be found. Additional factors may also explain the lack of bones. According to religious custom assuring con- tinuation of the beaver, bones from beavers that were consumed by Mi’kmaq were not thrown into the fire or river, nor fed to dogs, although practices vary in detail and by location (Denys 1672; Wallis and Wallis 1955; Robinson and Heller 2017). Bone material and metal goods are generally not preserved in the acidic soils of PEI except in acid-neutralizing shell middens (Murphy and Black 1996). Indigenous peoples con- sumed beavers, and because beaver incisors were often used as cutting tools, their remains are found in the common areas of preservation, kitchen middens. Of the 14 records of beaver teeth presented here, not all have been dated, but dated specimens were de- posited from about 500 AD to as late as 1650 AD. The teeth could have been imported to PEI as tools (Sobey 2007; M. Betts pers. comm. 12 June 2013) but we have also shown that beavers swim or raft to islands, sometimes far offshore. The simplest explan- ation for the presence of beaver teeth at PEI archaeo- logical sites is that they are the remains of PEI bea- vers. As well, the rib bone of an immature beaver at a site that may be 2000 years old, suggests that beavers were breeding on PEI. Discounting this, one must find an explanation as to why beavers did not swim or raft to PEI, as they did to Newfoundland, or disperse to PEI when a land bridge was in place for 4000 years. With postglacial warming temperatures, vegeta- tion on the island changed rapidly from tundra (suit- able for beavers; Aleksiuk 1970; Jung et al. 2016; Tape et al. 2018) to forest, a spruce (Picea sp.)— nonarboreal birch (Betula sp.) association between 10000 and 8000 years BP, followed by pines (Pinus sp.; Anderson 1980). The presence of beaver on Cape Breton Island 9500 years BP (Gorham et al. 2007) aligns well with the maximum connection of the PEI land mass to the mainland 9000 years BP, a continu- ous land mass lasting until 5000 years BP (Shaw et al. 2002). Beavers are also efficient dispersers (Leege 1968; Hodgdon 1978; Sun et a/. 2000) and can swim 2019 long distances. We cannot identify any impediments to beavers populating the non-island in early post- glacial times or any dramatic ecosystem changes that would preclude beavers colonizing PEI. A land bridge and the presence of beavers in the region might suggest that they inhabited PEI soon after deglacia- tion. The ability of beavers to swim or raft to islands is convincing evidence that they inhabited PEI prior to 1534. Archaeological evidence indicates the pres- ence of beavers until at least 1650 AD. Current status of beaver The second-growth riparian forests of PEI provide suitable beaver habitat and the beaver has populated most of it. It is likely that human conflicts with beavers and their dams will persist as long as beavers flood transportation corridors and are viewed as negatively influencing the spawning success of salmonids. Conclusion It is quite credible that the beaver could have been extirpated from PEI in the roughly 200 years before French settlement in 1721. Extinction rates of mam- mals are orders of magnitude higher on islands than elsewhere and are often related to human predation in historical times (Loehle and Eschenbach 2012). Caribou (Rangifer tarandus), Canada Lynx (Lynx canadensis), North American Black Bear (Ursus americanus), River Otter, and American Marten (Martes americana) were all extirpated from PEI fol- lowing European settlement (Sobey 2007). Although Caribou in Nova Scotia were extirpated by 1921 (Benson and Dodds 1977), none were reported after 1765 in PEI. Human exploitation was also responsi- ble for the loss of Walrus (Odobenus rosmarus) from the Gulf of St. Lawrence beginning in the 1500s (McLeod et al. 2014), and Great Auk (Pinguinus im- pennis) became extinct in 1844, aided in part by their exploitation at Bird Rock in the Magdalen Islands, Quebec (Montevecchi and Kirk 1996). Cameron (1958) contended the beaver was “exter- minated” from PEI, and the data presented here sup- port its status as native, at least since 500 AD and possibly as early as 9500 years BP. Evidence that might allow determining the point of extirpation is less clear, but it is almost certain that a beaver popu- lation was no longer present in PEI after 1860. It may well have been the first mammal extirpated from PEI, before 1700. Although the current beaver population is known to be derived from animals introduced from NB, it is also possible that some individual beavers have reached PEI via natural dispersal from NS or NB and could account for the late 19th century re- cords from PEI. Future genetic studies may shed light on whether NS beavers have contributed to the cur- rent gene pool. In addition, and considering there are CURLEY ET AL.: BEAVERS ON PEI 539 no known endemic species in PEI because of its geo- logically recent land connection with the mainland, beavers sourced from NB are predicted to be simi- lar genetically to the original PEI population. It may be possible to test this using more archaeological re- mains of beavers as they become available. Additional radiocarbon dating of beaver incisors from middens may also reveal new information. Finally, because beaver-chewed sticks were seen as an indication that beavers were native mammals (Cameron 1958), mon- itoring bogs that are being mined for peat might yield older beaver records. Although the founders of the current population were introduced to support fur harvesting (Dibblee 1994), the population meets International Union for the Conservation of Nature guidelines as a reintro- duction, being “the intentional movement and re- lease of an organism inside its indigenous range from which it has disappeared” (IUCN/SSC 2013: 2). We suggest that the American Beavers now extant on PEI be regarded as a native population and that the prov- incial government apply the precautionary principle in the unlikely event that population decline threat- ens the species. The second-growth riparian forests of PEI provide suitable beaver habitat and the beaver has populated most of the island. It is likely that hu- man conflicts with beavers and their dams will occur as long as beavers flood transportation corridors and are viewed as negatively influencing spawning suc- cess of salmonids. Author Contributions Original Draft: R.C.; Writing — Review & Editing: R.C., D.L.K., H.E.K., and R.L.D.; Conceptualization: R.C:; Investigation: D.L.K., H.E.K., R.C., and R.L.D.; Methodology: R.L.D.; Formal Analysis: D.K. and R.L.D.; Funding Acquisition: H.E.K. and R.L.D. Acknowledgements Thanks to Scott Buchanan for encouragement to write on this topic and for pointing to historical refer- ences regarding exploitation of fur and fish in the Gulf of St. Lawrence. Thanks also to Donald McAlpine of the New Brunswick Museum for reviewing an early draft of this article. For use of unpublished data, we thank Garry Gregory, Prince Edward Island (PEI) Department of Environment, Water and Climate Change, Matthew Betts of the Canadian Museum of History, Wayne Jordan of Stratford, PEI, and Brigitta Wallace of Parks Canada, Halifax. Independent re- searcher, Frances Stewart, provided archaeological services for H.E.K. Kim Proulx, PEI Department of Environment, Water and Climate Change, prepared the map. Vicki Johnson, Charlottetown, PEI, pre- pared the tables. We are grateful to the anonymous 340 reviewers whose comments and suggestions led to improvements to the manuscript. Literature Cited Aleksiuk, M. 1970. The seasonal food regime of arctic beavers. Ecology 51: 264-270. https://doi.org/10.2307/ 1933662 Anderson, C.B., G.M. Pastur, M.V. Lencinas, P.K. Wallem, M.C. Moorman, and A.D. Rosemond. 2009. Do introduced North American beavers Castor cana- densis engineer differently in southern South Amer- ica? An overview with implications for restoration. Mammal Review 39: 33-52. https://doi.org/10.1111/).13 65-2907.2008.00136.x Anderson, T.W. 1980. Holocene vegetation and climatic history of Prince Edward Island, Canada. Canadian Journal of Earth Sciences 17: 1152-1165. https://do1.org/ 10.1139/e80-122 Anonymous. 2011. Beaver management policy. Depart- ment of Environment, Energy and Forestry, Prince Ed- ward Island. Accessed 28 January 2018. https://www. princeedwardisland.ca/sites/default/files/publications/ 2011_beaver_policy.pdf. Bain, F. 1890. The Natural History of Prince Edward Is- land. G.H. Haszard, Charlottetown, Prince Edward Is- land, Canada. Benson, D.A., and D.G. Dodds. 1977. The Deer of Nova Scotia. Nova Scotia Department of Lands and Forests, Halifax, Nova Scotia, Canada. Blundell, G.M., M. Ben-David, P. Groves, R.T. Bowyer, and E. Geffen. 2002. Characteristics of sex-biased dis- persal and gene flow in coastal river otters: implica- tions for natural recolonization of extirpated popula- tions. Molecular Ecology 11: 289-303. doi.org/10.1046/ j.0962-1083.2001.01440.x Cairns, D.K., D.L. Guignion, T. Dupuis, and R.E MacFarlane. 2010. Stocking history, biological char- acteristics, and status of Atlantic Salmon (Salmo salar) on Prince Edward Island. Canadian Scientific Advisory Secretariat Research Document 2010/104. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Cameron, A.W. 1958. Mammals of the Islands in the Gulf of St. Lawrence. Bulletin 154. National Museum of Can- ada, Ottawa, Ontario, Canada. Cook, R. 1993. The Voyages of Jacques Cartier with an Introduction by Ramsay Cook. University of Toronto Press, Toronto, Ontario, Canada. Denys, N. 1672. Description geographique et historique des costes de l’Amerique Septentrionale, avec |’ Histoire na- turelle du Pais. [Re-published 1908 as The Description and Natural History of the Coasts of North America (Acadia). Translated and edited by W.F Ganong. Champlain Society, Toronto, Ontario, Canada.] Dibblee, R. 1994. The beaver on Prince Edward Island, seeking a balance. Island Magazine 35: 18—22. Dibblee, R., and R. Curley. 1980. Population studies of beaver on Prince Edward Island. Presented to Atlantic Chapter, Canadian Society of Environmental Biologists, 27-28 October 1980, Amherst, Nova Scotia, Canada. Dodds, D. 1983. Terrestrial mammals. Pages 509-550 in Biogeography and Ecology of the Island of Newfound- THE CANADIAN FIELD-NATURALIST Vol. 133 land. Edited by G.R. South. Dr. W. Junk Publishers, The Hague, Netherlands. Forbes, G.J., D.F. McAlpine, and F.W. Scott. 2010. Mammals of the Atlantic Maritime Ecozone. Pages 693-718 in Assessment of Species Diversity in the At- lantic Maritime Ecozone. Edited by D.F. McAlpine and I.M. Smith. NRC Research Press, Ottawa, Ontario, Canada. Gallant, D., C.H. Bérubé, E. Tremblay, and L. Vasseur. 2004. An extensive study of the foraging ecology of beavers (Castor canadensis) in relation to habitat qual- ity. Canadian Journal of Zoology 82: 922-933. https:// doi.org/10.1139/z04-067 Gesner, A. 1846. Report of the Geological Survey of Prince Edward Island. Appendix 4 in Journal of the Legislative Council of Prince Edward Island, Fourth Session of the Fifteenth General Assembly. Queens Printer, Char- lottetown, Prince Edward Island, Canada. Gorham, E., C. Lehman, A. Dyke, J. Jannsens, and L. Dyke. 2007. Temporal and spatial aspects of peatland initiation following deglaciation in North America. Quaternary Science Reviews 26: 300-311. https://doi. org/10.1016/j.quascirev.2006.08.008 Guignion, D. 2009. A conservation strategy for Atlantic Salmon in Prince Edward Island. Atlantic Salmon Federation. Accessed 16 June 2019. http://www.salmon conservation.ca/wp-content/uploads/2018/11/PEI- Atlantic-Salmon-Strategy-Report-2009.pdf. Harvey, D.C. 1926. The French Régime in Prince Edward Island. Yale University Press, New Haven, Connecticut, USA. Accessed 17 March 2020. https://archive.org/ details/frenchrgimeinpOOharv/page/n5/, Hodgdon, H.E. 1978. Social dynamics and behavior within an unexploited beaver (Castor canadensis) population. Ph.D. thesis, University of Massachusetts, Amherst, Massachusetts, USA. Ingersoll, L.K., and S.W Gorham. 1978. A history of the mammals of Grand Manan. Grand Manan Historian 20: 31-54. IUCN/SSC (International Union for the Conservation of Nature/Species Survival Commission). 2013. Guide- lines for Reintroductions and Other Conservation Trans- locations. Version 1.0. IUCN, Gland, Switzerland. Ac- cessed 15 June 2019. https://portals.iucn.org/library/ efiles/documents/2013-009. pdf. Johnstone, W. 1822. A Series of Letters Descriptive of Prince Edward Island in the Gulph of St. Lawrence. J. Swan, Dumfries County, Scotland. Pages 86-161 in 1955 re-publication Journeys to the Island of St. John or Prince Edward Island 1775-1832. Edited by D.C. Harvey. MacMillan, Toronto, Ontario, Canada. Jung, T.S., J. Frandsen, D.C. Gordon, and D.H. Mossop. 2016. Colonization of the Beaufort Coastal Plain by Beaver (Castor canadensis). a response to shrubifica- tion of the tundra? Canadian Field-Naturalist 130: 332— 335. https://do1.org/10.22621/cfn.v13014.1927 Keenlyside, D.L. 1982. Prince Edward Island archaeo- logical research, 1980. Pages 62—99 in Archaeological Research in the Maritimes 1980. Reports in Archaeology 5. Edited by C.J. Turnbull. Council of Maritime Pre- miers, Fredericton, New Brunswick, Canada. 2019 CURLEY ET AL. Keenlyside, D.L. 1983. In search of the Island’s first people. Island Magazine 13: 3-7. Keenlyside, D.L. 2002. Report on archaeological re- search in Prince Edward Island, 2000. Progress report Sutherland site CcCp-7 analysis, St. Peters Bay, PEI. Manuscript on file with Aboriginal Affairs Secretariat, Office of the Provincial Archaeologist, Charlottetown, Prince Edward Island, Canada. Kemp, P.S., T.A. Worthington, T.E.L. Langford, A.R.J. Tree, and M.J. Gaywood. 2012. Qualitative and quan- titative effects of reintroduced beavers on stream fish. Fish and Fisheries 13: 158-181. https://doi.org/10.1111/). 1467-2979.2011.00421.x Kranck, K. 1972. Geomorphological development and post- Pleistocene sea level changes, Northumberland Strait, Maritime Provinces. Canadian Journal of Earth Sci- ences 9: 835-844. https://doi.org/10.1139/e72-067 Kristmanson, H. 2007. Archaeological tests at Pitawelkek [Hog Island], Malpeque Bay, PEI, October 2007. Un- published report, on file with Aboriginal Affairs Secre- tariat, Office of the Provincial Archaeologist, Charlotte- town, Prince Edward Island, Canada. Kristmanson, H. 2009. Malpeque Bay archaeological pro- ject, October 7 & 9, 2009. Unpublished report on file with Aboriginal Affairs Secretariat, Office of the Pro- vincial Archaeologist, Charlottetown, Prince Edward Is- land, Canada. Kristmanson, H. 2019. Pitawelkek: a 2000 year old archae- ological site in Malpeque Bay. Island Magazine 84: 2-14. Leege, T.A. 1968. Natural movements of beavers in south- eastern Idaho. Journal of Wildlife Management 32: 973— 976. https://do1.org/10.2307/3799579 Leonard, K. 1989. Faunal analysis of the Rustico Is- land shell midden (7F) CcCT-1, Prince Edward Is- land,Canada. Preliminary report for University of Tor- onto, Department of Anthropology, and Canadian Parks Service, Halifax, Nova Scotia, Canada. Loehle, C., and W. Eschenbach. 2012. Historical bird and terrestrial mammal extinction rates and causes. Di- versity and Distributions 18: 84—91. https://doi.org/10. 1111/).1472-4642.2011.00856.x MacDonald, S.O., and J.A. Cook. 1996. The land mam- mal fauna of southeast Alaska. Canadian Field-Naturalist 110: 571-598. Accessed 10 October 2019. https://bio diversitylibrary.org/page/34343481. MacGregor, J. 1828. Historical and Descriptive Sketches of the Maritime Colonies of British North America. Longman, Rees, Orme, Brown and Green, London, United Kingdom. [Reprinted 1968 by S.R. Publishers, Johnson Reprint Corporation, New York, New York, USA] Marks, J.R. 1900. No beaver in Prince Edward Island. Summerside Journal, 19 Dec.: 1. Mazza, P.P.A., S. Lovari, F. Masini, M. Masseti, and M. Rustioni. 2013. A multidisciplinary approach to the analysis of multifactorial land mammal coloniza- tion of islands. BioScience 63: 939-951. https://doi.org/ 10.1525/bi0.2013.63.12.7 McLeod, B.A., T.R. Frasier, and Z. Lucas. 2014. Assess- : BEAVERS ON PEI 341 ment of the extirpated Maritimes walrus using morpho- logical and ancient DNA analysis. PLoS ONE 9: e99569. https://doi.org/10.1371/journal.pone.0099569 Mech, L.D. 1966. The wolves of Isle Royale. Fauna series 7. National Park Service, United States Department of Interior, Washington, DC, USA. Accessed 14 June 2019. http://npshistory.com/series/fauna/7. pdf. Montevecchi, W.A., and D.A. Kirk. 1996. Great Auk (Pin- guinus impennis), Version 2.0. In The Birds of North America. Edited by A.F. Poole and FB. Gill. Cornell Lab of Ornithology, Ithaca, New York, USA. https://doi. org/10.2173/bna.260 Moore, J.-S., V. Bourret, M. Dionne, I. Bradbury, P. O’Reilly, M. Kent, G. Chaput, and L. Bernatchez. 2014. Conservation genomics of anadromous Atlantic salmon across its North American range: outlier loci identify the same patterns of population structure as neutral loci. Molecular Ecology 23: 5680-5697. https:// doi.org/10.1111/mec.12972 Morlan, R. n.d. Canadian archaeological radiocarbon data- base. Canadian Museum of Civilization, Gatineau, Que- bec, Canada. Murphy, B.M., and D.W. Black. 1996. Zooarchaeology in the Canadian Maritimes. Canadian Zooarchaeology 9: 2-20. Accessed 10 October 2019. https://journals.uvic. ca/index.php/zooarchaeology/article/view/5261. Naughton, D. 2012. Natural History of Canadian Mammals. Canadian Museum of Nature and University of Toronto Press, Toronto, Ontario, Canada. Novak, M. 1987. Beaver. Pages 282-312 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J.A. Baker, M.E. Obbard, and B. Malloch. Ontario Ministry of Natural Resources and On- tario Trappers’ Association, Toronto, Ontario, Canada. Obbard, M.E., J.G. Jones, R. Newman, A. Booth, A.J. Satterthwaite, and G. Linscombe. 1987. Furbearer harvests in North America, 1600-1984. Pages 1007— 1034 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J.A. Baker, M.E. Obbard, and B. Malloch. Ontario Ministry of Natural Resources and Ontario Trappers’ Association, Toronto, Ontario, Canada. PEICLUI (Prince Edward Island Corporate Land Use Inventory). 2010. Prince Edward Island Corporate Land Use Inventory. Resource Inventory and modelling, Forests, Fish and Wildlife Division, PEI Department Communities, Land and Environment, Charlottetown, Prince Edward Island, Canada. Ray, A.J. 1987. The fur trade in North America: an over- view from ahistorical geographic perspective. Pages 21— 30 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J.A. Baker, M.E. Obbard, and B. Malloch. Ontario Ministry of Natural Resources and Ontario Trappers’ Association, Toronto, Ontario, Canada. Robinson, B.S., and A.S. Heller. 2017. Maritime cul- ture patterns and animal symbolism in Eastern Maine. Journal of the North Atlantic 10(sp10): 90-104. https:// doi.org/10.3721/037.002.sp1009 Rowan, J.J. 1876. The Emigrant and Sportsman in Canada. 342 Edward Stanford, London, England. [Reprinted 1972, facsimile edition by Coles Publishing, Toronto, Ontario Canada]. Shaw, J., P. Gareau, and R.C. Courtney. 2002. Palaeo- geography of Atlantic Canada 13-0 kyr. Quaternary Science Reviews 21: 1861-1878. https://doi.org/10.1016/ S0277-3791(02)00004-5 Shaw, J., D.J.W. Piper, G.B.J. Fader, E.L. King, B.J. Todd, T. Bell, M.J. Batterson, and D.G.E. Liverman. 2006. A conceptual model of the deglaciation of Atlantic Canada. Quaternary Science Reviews 25: 2059-2081. https://doi.org/10.1016/j.quascirev.2006.03.002 Sobey, D.G. 2002. Early descriptions of the forests of Prince Edward Island. I. The French period (1534-1758). Department of Agriculture and Forestry, Charlotte- town, Prince Edward Island, Canada. Sobey, D.G. 2006a. Early descriptions of the forests of Prince Edward Island. Hl. The British and Post-Con- federation periods—1758—c. 1900. Part A: the analy- sis. Department of Environment, Energy and Forestry, Charlottetown, Prince Edward Island, Canada. Sobey, D.G. 2006b. Early descriptions of the forests of Prince Edward Island. II. The British and Post- Confederation periods—1758-—c. 1900. Part B: the ex- tracts. Department of Environment, Energy and For- estry, Charlottetown, Prince Edward Island, Canada. Sobey, D.G. 2007. An analysis of the historical records for the native mammalian fauna of Prince Edward Island. Canadian Field-Naturalist 121: 384-396. https://doi.org/ 10.22621/cfn.v12114.510 Stewart, F.L. 2002. Small faunal remains from the Suther- THE CANADIAN FIELD-NATURALIST Vol. 133 land site, Prince Edward Island, 2000 and 2001 test ex- cavations by David Keenlyside. Unpublished report. National Museum of Civilization, Ottawa, Ontario, Canada. Sun, L., D. Miiller-Schwarze, and B.A. Schulte. 2000. Dispersal pattern and effective population size of the beaver. Canadian Journal of Zoology 78: 393-398. https://doi.org/10.1139/z99-226 Sutherland, G. 1861. A Manual of the Geography and Natural and Civil History of Prince Edward Island. John Ross, Charlottetown, Prince Edward Island, Canada. https://doi.org/10.5962/bh1 title.49420 Tape, K.D., B.M. Jones, C.D. Arp, I. Nitze, and G. Grosse. 2018. Tundra be dammed: beaver colonization of the Arctic. Global Change Biology 24: 4478-4488. https://doi.org/10.1111/gcb.14332 Wallace Ferguson, B. 1989. Selective exploitation of shell- fish at Rustico Island, Prince Edward Island. Presented to the Meeting of Canadian Archaeological Association, 10-14 May, Fredericton, New Brunswick, Canada. Wallis, W.D., and R.S. Wallis. 1955. The Micmac Indians of Eastern Canada. University of Minnesota, St. Paul, Minnesota, USA. Whitehead, R.H. 1993. Nova Scotia: the protohistoric period 1500-1630. Curatorial report 75. Nova Scotia Museum, Halifax, Nova Scotia, Canada. Accessed 13 May 2015. https://ojs.library.dal.ca/NSM/article/view/ 4080/3735. Received 4 October 2018 Accepted 23 December 2019 The Canadian Field-Naturalist Sixty years of White-tailed Deer (Odocoileus virginianus) yarding in a Gray Wolf (Canis lupus)—deer system L. Davip Mecu'2)* and SHANNON M. BARBER-MEYER!? 'United States Geological Survey, Northern Prairie Wildlife Research Center, 8711 - 37th Street SE, Jamestown, North Dakota 548401-7317 USA "United States Geological Survey, The Raptor Center, 1920 Fitch Avenue, St. Paul, Minnesota 55108 USA 7United States Geological Survey, 1393 Highway 169, Ely, Minnesota 55731 USA “Corresponding author: david_mech@usgs.gov; mechx002@umn.edu Mech, L.D., and S.M. Barber-Meyer. 2019. Sixty years of White-tailed Deer (Odocoileus virginianus) yarding in a Gray Wolf (Canis lupus)—deer system. Canadian Field-Naturalist 133(4): 343-351. https://doi.org/10.22621/cfn.v13314.2136 Abstract This article synthesizes information from over a six-decade period of studies of White-tailed Deer (Odocoileus virginianus) use of a winter yard and subject to Gray Wolf (Canis /upus) predation in northeastern Minnesota. It also adds spring migra- tion data from 35 adult female deer and fawns studied there during 1998, 1999, 2001, 2014, and 2017. Twenty-nine of these deer migrated in spring a mean distance of 29 km (SE = 4), a maximum distance of 78 km, and at a mean bearing of 83° (SE = 12; range 21-348). These findings are similar to those from 49 deer (both sexes) from the same yard studied during 1974-1984, that migrated a mean distance of 25 km (SE = 1.8) and a mean bearing of 77° + 4 SE. Between the two periods, the wolf population fluctuated considerably, the winter range of deer in the area where these deer spent summer greatly diminished, and both derechos and fires disturbed the habitat. This study attests to the selective advantage of the migratory tradition of deer in this yard. Key words: Canis lupus, deer yard; migration; Odocoileus virginianus; predation; predator-prey relations; White-tailed Deer; wolf; yarding Introduction White-tailed Deer (Odocoileus virginianus) mi- grate between summer and winter ranges in many northern areas (Summarized by Nelson 1998). Two main drivers of these migrations have been proposed: (1) the need for optimal protection from adverse win- ter weather (Townsend and Smith 1933; Severinghaus and Cheatum 1956; Ozoga 1968) and (2) grouping to minimize predation risk (Nelson and Mech 1981, 1991; Messier and Barrette 1985). Most studies of migratory deer populations have been short term, describing migration distances, tim- ing, and triggers for seasonal movements. One excep- tion is an investigation of deer movements in south- eastern Quebec that also depicted the extent of two deer yards over three decades (Lesage ef al. 2000). Studies of deer migratory behaviour in areas where Gray Wolves (Canis /upus) are the primary predator of deer have been conducted for as long as 10 years (Forbes and Theberge 1995; Theberge and Theberge 2004), 15 years (Fieberg et al. 2008), and 28 years (Hoskinson and Mech 1976; Nelson and Mech 1981, 1987; Nelson 1995, 1998; Nelson et al. 2004). However, we know of no migratory White-tailed Deer herd subject to wolf predation that has been in- vestigated for more than three decades. As part of a long-term study of wolf ecology and population trend in northeastern Minnesota (Mech 2009), we have also researched White-tailed Deer there since 1964 (Mech and Frenzel 1971; Hoskinson and Mech 1976; Nelson and Mech 1981, 1987; Nelson 1998; Nelson et a/. 2004). During that time, the amount of winter range of the deer herd we studied diminished greatly (Mech and Karns 1977). Forty- five years later, some 3000 km? that deer previously used for decades during winter remained devoid of wintering deer (Nelson and Mech 2006), and most, and probably all, of it still remains devoid of winter- ing deer (Mech e¢ al. 2018). In addition, various habi- tat disturbances and other important changes detailed below have occurred in the wolf study area. The wolf study area (Figure 1) lies in northeast- ern Minnesota, USA at about 47.60°N to 48.7333°N and 90.8167°W to 91.8333°W excluding the north- west quarter of that region and includes much of the Garden Lake deer yard (GLY) along its western edge 343 ©This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 344 —-—. Ontario Wolf study 91°Wi area Superior National Forest Boundary Isabella Lake Co. 20 km ee awe L. Superior FicurE 1. The wolf study area with the Garden Lake Yard (GLY). Irregular grey and stippled areas represent the GLY as described by Mech and Karns (1977). Grey and stippled ovals represent areas listed as deer yards by Arnold et al. (1961). Stippled areas (both irregular and oval) are where deer have not overwintered since the early 1970s (Mech and Karns 1977; Nelson and Mech 2006; Mech et al. 2018). The darker bold oval just east of Ely is the GLY proper, where White-tailed Deer (Odocolieus virginianus) from previous studies mentioned in the Introduction and the present study were radio-collared. Inset map shows location of Superior National Forest (black) in Minnesota. near Ely, Minnesota, USA. The GLY is named for the area around Garden Lake and the adjacent area near the Winton Hydroelectric Power Plant where winter- ing deer concentrate the most under the most severe conditions and where deer were fed artificially in the early 1970s and probably for some time before that. Deer have continued to concentrate in the GLY dur- ing winter and to migrate to summer ranges in and through the wolf study area for over 60 years. We studied the migratory behaviour of deer in this yard from 1974 through 1984 (Hoskinson and Mech 1976; Nelson and Mech 1981, 1987, 1991; Nelson 1998) and again during 1998 through 2017. We document here the continued winter concentration of deer in that yard and their annual migrations despite those changes and despite a wolf population that depends on them for most of their diet (Barber-Meyer and Mech 2016). We also compare 1998-2017 demog- raphy and migratory status of the deer in that yard with results from 1974-1984 (Nelson and Mech 1981, 1987). The objective of this study is to demonstrate the extreme degree to which a migratory tradition in a given deer yard under natural conditions of wolf predation can persist, a record duration to our knowl- edge, and to compare the migratory behaviour over the period of this study. THE CANADIAN FIELD-NATURALIST Vol. 133 Study Area The extent of the GLY over the years has been de- scribed variously, no doubt because (1) deer popula- tions fluctuate greatly over the decades, and (2) deer use of winter range, and thus their migration move- ments, vary considerably by season, temperature, and snow conditions (Nelson 1995). As these conditions change, deer may move toward or away from win- ter yards, sometimes wintering for long periods only partly along their route to areas where they would concentrate more during the most extreme conditions (Nelson and Mech 1981). The Pohenegmook and Lac Temiscouata deer yards in southeastern Quebec, Canada provide a good example of such changes (see Figure 3 in Lesage et al. 2000). In 1953, the GLY was thought to encompass 128 ha, not including other yards west and east-northeast of the GLY (Erickson et al. 1961). Mech and Karns (1977) considered the GLY more inclusively, stretch- ing from about 35 km west-southwest of Ely to Ely, about 25 km east of Ely, and then northeast about 12 km, totalling about 72 km long, and centring on the Garden Lake area (Figure 1). In the mid-1970s the GLY was thought to extend about 16 km east-north- east (Hoskinson and Mech 1976) and later as holding <800 deer (Nelson and Mech 1987). East of Garden Lake, deer currently continue to winter along the area that Hoskinson and Mech (1976) described at times as far as some 18 km east of Garden Lake. Whether deer wintering elsewhere in the more ex- pansive GLY other than those from the capture area (Nelson and Mech 1981, 1987, this study) migrate in the same direction to summer ranges as those deer ra- dio tracked is unknown. The GLY lies along the western edge of our long- term wolf study area (Mech 2009) which covers about 2060 km? including the migration routes along which the wintering GLY deer travel to their sum- mer ranges (Figure 1). The wolf study area is situated well within the Minnesota wolf range (Fuller et al. 1992), and wolves have never been extirpated from the wolf study area. The area is replete with lakes and waterways, and American Beaver (Castor canaden- sis) and Moose (Alces americanus) are also availa- ble to wolves there (Mech and Karns 1977; Barber- Meyer and Mech 2016; Mech ef a/. 2018). Black Bear (Ursus americanus) 1s the only other major preda- tor of deer in the region (Kunkel and Mech 1994), al- though Coyote (Canis latrans), Fisher (Martes pen- nant), Bobcat (Lynx rufus), and Canada Lynx (Lynx canadensis) inhabit the area and could prey on fawns. General habitat, topography, and weather in the study area were described by Nelson and Mech (1981, 2006) and Heinselman (1996). 2019 In July 1999, a derecho windstorm leveled about 1600 km? of the forest through which some of the GLY deer migrate (National Oceanic and Atmospheric Administration 1999; Nelson and Mech 2006). Another derecho struck in 2016 that also affected the migration routes of these deer (Minnesota Department of Natural Resources 2016). In 2000 and 2007, fires burned 431 km”, just north- east beyond where radio-collared GLY deer migrate to but which could include summer ranges of other GLY deer (Fites et al. 2007). In 2011, the Pagami Creek fire burned 376 km? in which some GLY deer summered, or through which they migrated (Nelson and Mech 1987). Other habitat changes during the study included forest maturation, and alterations and variation in logging practices from clear cutting to to- tal protection. Weather conditions also changed considerably throughout the study. Snow depth, density, and per- sistence, especially during the past decade, differed from earlier in the study, including winter 2010— 2011 when snow depth was extremely low and win- ter 2013-2014 when snow was very deep and fluffy. White-tailed Deer have inhabited the region for many decades. Johnson (1922) considered deer com- mon from 1912 to 1915. In 1938, Olson (1938: 330) published a map showing deer present in every town- ship in the wolf study area. From 1948 to 1952, Stenlund (1955) documented wolf-killed deer in win- ter on most of the major lakes there. Erickson et al. (1961) stated that deer were abundant in the Northern Forest Zone, which included our wolf study area, for more than 40 years, and those authors listed 16 winter yarding areas they checked in or near our wolf study area between 1949 and 1958. Estimated deer densi- ties in the Northern Forest Zone (although not neces- sarily in our wolf study area) ranged from 5.9 to more than 7.8/km/? in the late 1930s (Erickson et al. 1961). By the mid-1970s, almost no deer spent winter in the northeastern third of the wolf study area, and wolves there lived primarily on Moose and probably beavers (Mech and Karns 1977). Deer that had win- tered there had succumbed to a combination of de- teriorating habitat (maturing forests), a long series of severe winters, and heavy wolf predation (Mech and Karns 1977). Deer have not been observed over- wintering there since, despite regular winter flights (Nelson and Mech 2006; Mech et a/. 2018). Deer num- bers along the southern and western edges of this area dropped to about 0.8 deer/km? (Floyd et al. 1979) and in 2011 pre-fawn densities averaged <2/km? (Lenarz and Grund 2011). To the east of the wolf study area, deer migrated during autumn to winter yards along the shore of Lake Superior (Nelson and Mech 1981) and reached yarding densities during 1968-1976 of MECH AND BARBER-MEYER: MIGRATING DEER 345 39 to 55/km? (Mech and Karns 1977). Deer from those yards moved at least 22 km northwest inland (Morse and Zorichak 1941; Nelson and Mech 1981). Deer that wintered in yards along the west side of the wolf study area, primarily in and around Garden Lake, 8.8 km east-northeast of Ely, migrated in spring southeastward to northeastward for up to 54 km ata mean bearing of 77° (Nelson and Mech 1987). Moose have also occupied the region for many decades. Johnson (1922) found Moose very com- mon in 1912-1915 but scarce in 1920. Olson (1938) estimated a Moose density of 1/6.4 km? based on his observations during 1920-1936 and his discussions with various wardens, trappers and other woods- men, but Stenlund (1955: 22) considered their num- bers “not high” during 1948-1952. An historical es- timate of Moose density from 1915 to 1970 over the entire northeastern Minnesota Moose range, which included our wolf study area, was 1/3.8 km? to 1/21.9 km? (Peek et al. 1976). From 1984 to 2016 in this Moose range, densities based on annual aerial counts were 1/1.7 km? to 1/5.5 km? (calculated from Moose- count data; Mech et a/. 2018). Moose numbers in the overall northeastern Minnesota Moose range peaked in 1989, 1996, and 2006, declined to less than half their 2006 level by about 2012, and then leveled off for several years (DelGiudice 2017; Mech et al. 2018). Wolves have inhabited the region throughout re- corded history (Olson 1938; Stenlund 1955; Mech and Frenzel 1971). Wolf numbers in the wolf study area varied from 23-32 in winter 2016-2017 (L.D.M. and S.M.B.-M. unpubl. data) to 97 in 2008-2009, a den- sity ranging from 11—16/1000 km? to 47/1000 km? during 1968-2017 (Mech 1973, 1986, 2009; Mech et al. 2018). During and after the major deer decline in the 1970s, wolf numbers there also declined con- siderably and did not reach former levels until about 2000 after recovering from a prolonged infection by canine parvovirus (Mech et al. 2008). A few years af- ter Moose numbers began declining in 2006 and deer numbers declined due to severe winters, the wolf population began dropping to its lowest level during the study, 23-32 animals (Barber-Meyer and Mech 2016; Mech et al. 2018). The primary migration routes and many of the summer ranges of the GLY deer we studied usually fell within the territories of two wolf packs, known as the Wood Lake and Ensign Lake Packs in earlier pub- lications (Mech 1973, 1986). Over the decades, the actual locations of these pack territories varied con- siderably, and other packs that used parts of the GLY, the deer migration routes, or the summer ranges of the GLY deer formed and disintegrated as well. At times, as many as four radioed packs, totalling up to 29 members during winter used the GLY (L.D.M., 346 S.M.B.-M., and M.E. Nelson unpubl. data). In addi- tion, wolf packs sometimes inhabited the GLY year around. One such pack that inhabited 39 km? includ- ing Garden Lake itself hosted the highest wolf density ever recorded anywhere, 182 wolves/1000 km? dur- ing winter, from 1 April 1998 through 30 March 1999 (Mech and Tracy 2004). Based on 39 years during which the Wood Lake Pack was radio-collared and 24 years in which the Ensign Lake Pack was radio-collared between 1973 and 2017, their winter pack sizes averaged 5.3 + 0.41 SE and 5.6 + 0.55 SE and ranged up to 11 and 12 members, respectively (L.D.M., S,.M.B.-M., and M.E. Nelson unpubl. data). The numbers of wolves in these packs did not follow the trajectory of the over- all wolf numbers in the wolf study area, but rather remained relatively constant from winter 1973-— 1974 through about 2006, although they declined af- ter that (Mech 1973, 1986, 2009; L.D.M.,S.M.B.-M., and M.E. Nelson unpubl. data). In any given year, the packs that used the area including the GLY deer sum- mer ranges and migration routes usually migrated to the Garden Lake area itself during autumn and back to the deer summer ranges in spring (Mech and Boitani 2003; L.D.M. and S.M.B.-M. unpubl. data) except when resident packs resided year around there. Methods Using Clover traps from 1998 to 2017, we live trapped, anesthetized, ear tagged, and radio col- lared deer within 1.4 km of the GLY (Mech and Barber-Meyer 2020). Three others were captured near Snowbank Lake, some 23 km east northeast of Garden Lake but still in the more expansive definition of the GLY discussed above. In the current study we excluded the three Snowbank Lake deer (included in a study by Nelson ef al. [2004]) because that area was not included in the Nelson and Mech (1987) area with which we compare our data. Our GLY captures were basically in the same area where deer (both sexes) from this yard were studied earlier (Hoskinson and Mech 1976; Nelson and Mech 1981, 1987). We ex- tracted an incisor from adults for aging by Matson’s Laboratory (Missoula, Montana, USA). We located the deer by aerial radio tracking or by global posi- tioning system (GPS) collar locations during June, July, and August until at least two consecutive loca- tions were in the same general area to determine their summer ranges (because generally once on summer range they remain in a relatively small area [Nelson and Mech 1999]) and again each winter when they returned to the winter yard (Nelson et al. 2004). We examined the approximate spring migration routes of deer collared with prototype Advanced Telemetry Systems (Isanti, Minnesota, USA) drop-off GPS THE CANADIAN FIELD-NATURALIST Vol. 133 radio collars (details in Merrill et a/. 1998), including some studied by Nelson ef al. (2004). We plotted individual deer summer locations and a summary location representing the centre of the winter deer capture locations on Google Earth Pro 7.1.7.2606 and measured the migration distances and directions via the Google Earth Tool function after converting UTMs of these locations to latitudes and longitudes via “Convert Geographic Units online” (http://www.rcn.montana.edu/resources/converter. aspx). Although fawns captured during the same year and at the same location as an adult female and mi- grating to the same summer range as the adult (or not migrating but remaining at the same summer range as the adult) might have been fawns of the adult, we still included the fawns as independent data. We used Statistix 9.0 (2008) to compare migratory sta- tus (including fawns) between our 1998-2001 and our 2014/2017 results using Fisher’s Exact Test, re- spectively, and also to those from a previous study in the same area (Nelson and Mech 1981, 1987). We compared age structures (excluding fawns) between 1998—2017 and those from the previous study (Nelson and Mech 1981, 1987) via the Mann-Whitney U-test in R version 3.5.1 (R Core Team 2018). We considered all differences significant at alpha = 0.05. Results We live-trapped and radio-collared 27 adult does and eight fawns during winters 1998-2001, 2014, and 2017 in or near the GLY and aerially radio-tracked them to their summer ranges (Table 1), including eight whose spring migrations were studied in de- tail by Nelson ef al. (2004). Apart from fawns, their mean age was 6.3 (SE = 0.8) years (Figure 2). All of the 19 deer we radio-collared in 1998-2001, includ- ing fawns, migrated to summer ranges, but six (in- cluding two fawns) of the 16 that we followed in 2014 and 2017 remained during summer within 3 km of their winter capture point, a significant difference be- tween these two periods (Fisher’s Exact = proportion TABLE 1. Female White-tailed Deer (Odocoileus virgin- ianus) or fawns of either sex radio-collared (first capture only) in Garden Lake Yard, Ely, Minnesota, USA, 1998-— 2017 and radio-tracked to their summer ranges. Six deer did not migrate. n Age (year)* Aa (# fawns) Mean Range 1998 8 (2) 5.6 3-11 1999 5 (0) 0) 1-13 2001 6 (1) a7 1-13 2014 4 (1) 63 5-8 2017 12 (4) 71 paeile! 1998-2017 35(8) 6.3 1-13 *27 adults and yearlings; excludes two adults of unknown age. 2019 4 3 2 TELE a] ht tl 12 3 4 5 6 7 8 9 10 11 12 13 Years Number FIGURE 2. Age structure of adult and yearling female White- tailed Deer (Odocolieus virginianus) live-trapped (first cap- ture only), in or near the Garden Lake Yard, Minnesota, 1998-2017, radio-collared, and followed to summer range. difference 0.375, P =0.005). The mean age of the four adult non-migrating deer was 7.3 and that of the 21 non-fawn migrators was 6.1. The age structures of the groups did not differ (W = 33, P = 0.53). The 35 adults and fawns migrated in spring a mean distance of 29 km (SE = 4), a maximum dis- tance of 78 km, and at a mean bearing of 83° (SE = 12; range = 21-348) excluding the six non-migrators (Table 2; Figure 3). Although the deer during differ- ent years of the study varied in the distances and di- rections to which they migrated, most of the annual mean migration distances were 21-36 km, and most of the annual mean migration bearings were 58—90° (Table 2). The 114° mean bearing for five deer in 1999 was heavily influenced by one deer whose migra- tion bearing was 348°. Excluding that deer, the mean bearing was 55° (SE = 14). Notably, two other deer captured in the same general location as deer that mi- grated east-northeastward migrated in markedly dif- MECH AND BARBER-MEYER: MIGRATING DEER 347 ferent directions southwest, and south. Excluding all three deviant deer, and the non-migrators, the mean summer migration bearing was 65° (SE = 4; n = 26), the basic direction that the GLY extended. The mean migration distance of this sample was 29 km (SE = 4; 4-78 km). Discussion The sample of 35 does and fawns we studied from 1998 through 2017 generally was similar to that of the does and fawns studied from 1974-1984 in the same area (Nelson and Mech 1981, 1987). We compared these two periods (19 and 10 years long) because those were the periods for which we had comparable data. There was no significant difference in the radio- collared doe:fawn ratios (37:19 versus 39:28) between the early and later capture samples (Fisher’s Exact = proportion difference 0.079, P = 0.46). The mean age of adult does of the earlier sample was 5.0 years and that of the later sample was 6.3 years. The age struc- tures of the groups did not differ (W = 359, P = 0.24). The 1998-2017 sample of does and fawns that we fol- lowed through spring migration migrated similarly in mean distance (25 km + 1.8 SE) to those from 1974— 1984, but not maximum (78 km this study versus 54 km, measured from Nelson and Mech [1987: Figure 2.2]). They were also similar in the general directions they migrated (77° +4 SE; Nelson and Mech 1987). Of the 49 GLY deer (18 males: 31 females) whose spring migrations were studied from 1974 to 1984, 42 migrated (Nelson and Mech 1987), and with our 1998-2017 sample of 35 does and fawns, all except six migrated, a non-significant difference between pro- portions of migrators during the two periods (Fisher’s Exact = proportion difference 0.029, P = 0.77). The demography and migration we studied in the sample of deer wintering in the GLY differed little TABLE 2. Migration distance and direction of White-tailed Deer (Odocoileus virginianus) that were radio-collared during 1998 through 2017 and followed to their summer ranges. Fawns possibly of collared does were included separately. Summer migration Year No. of No. Distance (km) Direction (°) Remarks deer migrating SE Maximum ca SE Range 1998 8 8 36+6 62 64+ 8 26-97 1999 5 5 3144 45 114+ 60 21-348 2001 6 6 26+ 10 58 78 +17 39-153 2014 4 1 Dery 8 Includes three non-migrators 2014 4 1 8 8 58 — Excludes three non-migrators 2017 12 9 92 it 78 Includes three non-migrators 2017 12 9 2627 78 90 + 18 36-226 Excludes three non-migrators 1998-2001 19 19 3144 62 81416 21-348 2014—2017 16 10 16255 78 Includes six non-migrators 2014—2017 16 10 26:57 78 87 + 16 36-226 Excludes six non-migrators 1998-2017 35 oS) 24 +3 78 Includes six non-migrators 1998-2017 35 29 29+4 78 83+ 12 21-348 Excludes six non-migrators FicureE 3. Distances and directions of spring migrations of 29 adult female and fawn White-tailed Deer (Odocolieus virginianus) radio-collared in the Garden Lake Yard during five winters between 1998 and 2017 (Table 1). Six of the ori- ginal sample of 35 did not migrate. from those studied there during 1974-1984. During the interim, several important environmental changes took place, as discussed in the Introduction. Throughout this period and despite the chang- ing deer, Moose, and wolf populations, as well as the widespread habitat upsets (e.g., derechos, forest fires, snowpack differences, changes in forestry practices), the majority of GLY deer continued to migrate each winter to the GLY the way they have for decades. Furthermore, we cannot extrapolate our findings to other migrating ungulate-wolf systems and would ex- pect each deer yarding situation to be different be- cause each local yarding ecology will be different. Nelson (1995, 1998) and Nelson et al. (2004) pro- vided details of the earlier migrations. The wolves that inhabited the major portions of the GLY deer summer and winter ranges maintained their numbers through about 2006. After Moose began to decline in 2006, the number of these wolves decreased, but packs continued to migrate each year for which we had data, presumably in response to the deer migration (L.D.M. and S.M.B.-M. unpubl. data), similar to wolf packs in Algonquin Park, Ontario, Canada (Forbes and Theberge 1995; Theberge and Theberge 2004). During summer, the major age class of deer that local wolves kill are fawns (Nelson and Mech 1986; Barber-Meyer and Mech 2016), although the availa- bility of beavers and Moose might buffer that preda- tion (Mech and Karns 1977; Barber-Meyer and Mech 2016). Evidence from other parts of the wolf study area suggests that individual fawns are visited by wolves on average in summer about 5.5 times/100 days (Demma and Mech 2009) to daily (Mech er al. 2015), although the rate of fawn predation is unknown. Regardless, even though fawns comprise a high percentage of the THE CANADIAN FIELD-NATURALIST Vol. 133 diet of wolves in summer (Barber-Meyer and Mech 2016), enough fawns have survived in the summer ranges of the GLY deer each year to sustain the mi- grating deer population over the decades. GLY migrating deer spend 31-356 hours during migration and adhere closely to a straight line dur- ing the trip (Nelson et a/. 2004). While migrating, deer are much more vulnerable to wolf predation than at any other time as adults (Nelson and Mech 1991), so the persistence of GLY deer either aban- doning summer range or favouring winter range or both during winter must have some strong adaptive value. Reducing vulnerability to wolf predation dur- ing winter when deer are in poor nutritional condition (DelGiudice et a/. 1992) and hindered by snow condi- tions (Mech et a/. 1971) was the explanation Nelson and Mech (1981) gave for deer in this area migrat- ing to areas of high deer density, 1.e., the GLY, listing several advantages to yarding. This benefit was one of the points Nelson and Mech (1981) proposed as an anti-predator effect of yarding. We further note that Poszig and Theberge (2000) did find that non-yard- ing deer in their study were “highly vulnerable” when migrating wolves returned to their territory. Kolenosky (1972) had already shown that wolves tended to kill deer along the edges, rather than the centre of the deer yards he studied, and further sup- port for the antipredator explanation for deer migra- tion and yarding has since been found in other stud- ies. In northwestern Minnesota, wolves also tended to kill deer along the edges of yarding areas rather than in the densest areas (Fritts and Mech 1981) as did Coyotes in Quebec (Messier and Barrette 1985). On the other hand, Poszig and Theberge (2000) found evidence in Ontario that tended to dispute the hypothesized antipredator advantages of deer yard- ing. The only benefit of yarding they proposed would be an enhanced trail network through the snow that might give deer in high densities more of an advan- tage in escaping wolves. Henderson et al. (2018) emphasized the role of density-dependent competition for home ranges in winter that forced deer to space out during summer to obtain adequate nutrition. The spacing out of migrat- ing deer to their summer ranges, where their fawns are born, provides far more habitat per deer to ob- tain nourishment, with summer being the season of annual replenishment (Silver et al. 1969; Moen 1978; DelGiudice et al. 1992). However, it also brings sev- eral other survival benefits related to wolf predation: (1) familiar escape terrain and habitat; (2) an area with a proven history of survival characteristics; and (3) separation from other fawns that would attract 2019 predators. Fawns are most vulnerable during late spring and early summer (Kunkel and Mech 1994; Carstensen et al. 2009), so widely spaced fawns re- duce the chance that any individual fawn would be detected by predators, thus increasing survivability (although reducing potential benefits of group vigi- lance and defense). None of these benefits of return to summer range or migration to winter range (Nelson and Mech 1981) conflict with the Henderson et al. (2018) findings, for in complex ecosystems both foraging and pre- dation risk are factors between which animals must find trade-offs that enhance their survival (Lima and Dill 1990). Within the context of these trade-offs, our study demonstrates that, in an area where wolf preda- tion 1s the major natural mortality for adult deer, long deer migrations between winter and summer ranges and yarding in winter produces strong enough sur- vival value for the behaviour to have persisted for over six decades and many generations. Author Contributions Writing — Original Draft: L.D.M.; Writing — Re- view & Editing: L.D.M. and S.B.-M.; Conceptuali- zation: L.D.M.; Investigation: S.B.-M.; Methodology: L.D.M.; Formal Analysis: L.D.M. and S.B.-M.; Fund- ing Acquisition: L.D.M. Acknowledgements This study was supported by the United States Geological Survey (USGS) with the cooperation of the Superior National Forest. We thank numerous volunteer wildlife technicians for assisting with the deer captures; several United States Forest Service pilots for safe flying; and Dr. M.E. Nelson (USGS re- tired) for collecting the 1998—2001 data and for cri- tiquing an early draft of the manuscript. Any use of trade, firm or product names is for descriptive pur- poses only and does not imply endorsement by the United States Government. Literature Cited Barber-Meyer, S.M., and L.D. Mech. 2016. White-tailed deer subsidize gray wolves during a moose decline: a case of apparent competition? Canadian Field-Naturalist 130: 308-314. https://doi.org/10.22621/cfn.v13014.1924 Carstensen, M., G.D. DelGiudice, B.A. Sampson, and D.W. Kuehn. 2009. Survival, birth characteristics, and cause-specific mortality of white-tailed deer neonates. Journal of Wildlife Management 73: 175-183. https:// doi.org/10.2193/2006-107 DelGiudice, G.D. 2017. 2017 aerial moose survey. Minne- sota Department of Natural Resources, St. Paul, Minne- sota, USA. DelGiudice, G.D., L.D. Mech, K.E. Kunkel, E.M. Gese, and U.S. Seal. 1992. Seasonal patterns of weight, hema- tology, and serum characteristics of free-ranging fe- MECH AND BARBER-MEYER: MIGRATING DEER 349 male white-tailed deer in Minnesota. Canadian Journal of Zoology 70: 974-983. https://doi.org/10.1139/z92-139 Demma, D.J., and L.D. Mech. 2009. Wolf use of sum- mer territory in northeastern Minnesota. Journal of Wildlife Management 72: 380-384. https://doi.org/10. 2193/2008-114 Erickson, A.B., V.E. Gunvalson, M.H. Stenlund, D.W. Burcalow, and L.H. Blankenship. 1961. The white- tailed deer of Minnesota. Minnesota Department of Conservation, Project W-11-R. Technical Bulletin, No. 5, St. Paul, Minnesota, USA. Fieberg, J., D.W. Kuehn, and G.D. DelGiudice. 2008. Understanding variation in autumn migration of north- ern white-tailed deer by long-term study. Journal of Mammalogy 89: 1529-1539. https://doi.org/10.1644/07- mamm-a-277.1 Fites, J.A., A. Reiner, M. Campbell, and Z. Taylor. 2007. Fire behavior and effects, suppression, and fuel treatments on the ham lake and cavity lake fires. Accessed 25 January 2020. https://www.fs.fed.us/adaptivemanagement/reports/ fbat/Ham Lake 07_22_08 pdf. Floyd, T.J., L.D. Mech, and M.E. Nelson. 1979. An im- proved method of censusing deer in deciduous-conifer- ous forests. Journal of Wildlife Management 43: 258— 261. https://doi.org/10.2307/3800668 Forbes, G.J., and J.B. Theberge. 1995. Influences of a migratory deer herd on wolf movements and mortal- ity in and near Algonquin Park, Ontario. Pages 503-513 in Ecology and Conservation of Wolves in a Changing World. Edited by L.D. Carbyn, S.H. Fritts, and D.R. Seip. Canadian Circumpolar Institute, Edmonton, Al- berta, Canada. Fritts, S.H., and L.D. Mech. 1981. Dynamics, movements, and feeding ecology of a newly protected wolf popula- tion in northwestern Minnesota. Wildlife Monographs 80: 1-79. Fuller, T.K., W.E. Berg, G.L. Radde, M.S. Lenarz, and G.B. Joselyn. 1992. A history and current estimate of wolf distribution and numbers in Minnesota. Wildlife Society Bulletin 20: 42-55. Heinselman, M. 1996. The Boundary Waters Wilderness Ecosystem. University of Minnesota Press, Minneapo- lis, Minnesota, USA. Henderson, C.R., Jr., M.S. Mitchell, W.I. Myers, P.M. Lukacs, and G.P. Nelson. 2018. Attributes of seasonal home range influence choice of migratory strategy in white-tailed deer. Journal of Mammalogy 99: 89-96. https://doi.org/10.1093/jmammal/gyx148 Hoskinson, R.L., and L.D. Mech. 1976. White-tailed deer migration and its role in wolf predation. Journal of Wildlife Management 40: 429-441. https://doi.org/10. 2307/3799944 Johnson, C.E. 1922. Notes on the mammals of northern Lake County, Minnesota. Journal of Mammalogy 3: 33-— 39. https://doi.org/10.2307/1373450 Kolenosky, G.B. 1972. Wolf predation on wintering deer in east-central Ontario. Journal of Wildlife Management 36: 357-369. https://doi.org/10.2307/3799065 Kunkel, K.E., and L.D. Mech. 1994. Wolf and bear pred- ation on white-tailed deer fawns. Canadian Journal of Zoology 72: 1557-1565. https://do1.org/10.1139/z94-207 350 Lenarz, M., and M. Grund. 2011. Deer modeling 2011. Forest Wildlife Populations and Research Group and Farmland Wildlife Research Group, Minnesota Depart- ment of Natural Resources, St. Paul, Minnesota, USA. Lesage, L., M. Crete, J. Huot, A. Dumont, and J. Ouellet. 2000. Seasonal home range size and philopatry in two northern white-tailed deer populations. Canadian Journal of Zoology 78: 1930-1940. https://doi.org/10. 1139/z00-117 Lima, S.L., and L.M. Dill. 1990. Behavioral decisions made under the risk of predation: a review and prospec- tus. Canadian Journal of Zoology. 68:619—640. https:// doi.org/10.1139/z90-092 Mech, L.D. 1973. Wolf numbers in the Superior National Forest of Minnesota. USDA Forest Service Research Paper NC-97, St. Paul, Minnesota, USA. Mech, L.D. 1986. Wolf numbers and population trend in the Superior National Forest, 1967-1985. USDA Forest Service Research Paper NC-270, St. Paul, Minnesota, USA. Mech, L.D. 2009. Long-term research on wolves in the Superior National Forest. Pages 15-34 in Recovery of Gray Wolves in the Great Lakes Region of the United States: An Endangered Species Success Story. Edited by A.P. Wydeven, E.J. Heske, and T.R Van Deelen. Springer, New York, New York, USA. Mech, L.D., and S. Barber-Meyer. 2020. Garden Lake Deer Yard (Lake Co., MN) Migration Data, 1998-2017: U.S. Geological Survey data release. https://doi.org/10. 5066/f72b8xct Mech, L.D., and L. Boitani. 2003. Wolf social ecology. Pages 1-34 in Wolves: Behavior, Ecology, and Conser- vation. Edited by L.D. Mech and L. Boitani. University of Chicago Press, Chicago, Illinois, USA. Mech, L.D., J. Fieberg, and S.M. Barber-Meyer. 2018. An historical overview and update of wolf-moose inter- actions in northeastern Minnesota. Wildlife Society Bulletin 42: 40—47. https://doi.org/10.1002/wsb.844 Mech, L.D., and L.D. Frenzel, Jr. 1971. An analysis of the age, sex, and condition of deer killed by wolves in north- eastern Minnesota. Pages 35-51 in Ecological Studies of the Timber Wolf in Northeastern Minnesota. Edited by L.D. Mech and L.D. Frenzel, Jr. USDA Forest Service Research Paper NC-52, St. Paul, Minnesota, USA. Mech, L.D., L.D. Frenzel, Jr.. and P.D. Karns. 1971. The effect of snow conditions on the ability of wolves to capture deer. Pages 51-59 in Ecological Studies of the Timber Wolf in Northeastern Minnesota. Edited by L.D. Mech and L.D. Frenzel, Jr. USDA Forest Service Research Paper NC-52. North Central Forest Experi- mental Station, St. Paul, Minnesota, USA. Mech, L.D., S.M. Goyal, W.J. Paul, and W.E. Newton. 2008. Demographic effects of canine parvovirus on a free-ranging wolf population over 30 years. Journal of Wildlife Disease 44: 824-836. https://doi.org/10.7589/ 0090-3558-44.4.824 Mech, L.D., and P.D. Karns. 1977. Role of the wolf in a deer decline in the Superior National Forest. USDA Forest Service Research Report NC-148, St. Paul, Min- nesota, USA. Mech, L.D., A. Morris, and S. Barber-Meyer. 2015. THE CANADIAN FIELD-NATURALIST Vol. 133 White-tailed deer (Odocoileus virginianus) fawn risk from wolves (Canis lupus) during summer. Canadian Field-Naturalist 129: 368-373. https://doi.org/10.22621/ cfn.v12914.1758 Mech, L.D., and S. Tracy. 2004. Record high wolf, Canis lupus, pack density. Canadian Field-Naturalist 118: 127— 129. https://doi.org/10.22621/cfn.v118i1.896 Merrill, S.B., L.G. Adams, M.E. Nelson, and L.D. Mech. 1998. Testing releasable GPS collars on wolves and white-tailed deer. Wildlife Society Bulletin 26: 830-835. Messier, F., and C. Barrette. 1985. The efficiency of yard- ing behaviour by white-tailed deer as an antipredator strategy. Canadian Journal of Zoology 63: 785-789. https://doi.org/10.1139/z85-115 Minnesota Department of Natural Resources. 2016. Severe storms strike northland again: July 20-21, 2016. Accessed 25 January 2020. http://www.dnr.state.mn.us/ climate/journal/160720_21_severe.html. Moen, A.N. 1978. Seasonal changes in heart rates, activ- ity, metabolism, and forage intake of white-tailed deer. Journal of Wildlife Management 42: 715-738. https:// doi.org/10.2307/3800763 Morse, M.A., and J.L. Zorichak. 1941. Quarterly Pro- gress Report Wildlife Research Project 11-R, Vol. 1. Minnesota Department of Conservation, St. Paul, Min- nesota, USA. National Oceanic and Atmospheric Administration. 1999. July 4-5, 1999 derecho. Accessed 25 January 2020. http://www.spc.noaa.gov/misc/AbtDerechos/case pages/jul4-51999 page. htm. Nelson, M.E. 1995. Winter range arrival and departure of white-tailed deer in northeastern Minnesota. Canadian Journal of Zoology 73: 1069-1076. https://doi.org/10.11 39/z95-127 Nelson, M.E. 1998. Development of migratory behav- ior in northern white-tailed deer. Canadian Journal of Zoology 76: 426—432. https://doi.org/10.1139/z97-207 Nelson, M.E., and L.D. Mech. 1981. Deer social organiza- tion and wolf depredation in northeastern Minnesota. Wildlife Monographs 77: 1-53. Nelson, M.E., and L.D. Mech. 1986. Mortality of white- tailed deer in northeastern Minnesota. Journal of Wild- life Management 50: 691-698. https://doi.org/10.2307/ 3800983 Nelson, M.E., and L.D. Mech. 1987. Demes within a northeastern Minnesota deer population. Pages 27— 40 in Mammalian Dispersal Patterns. Edited by B.D. Chepko-Sade and Z. Halpin. University of Chicago Press, Chicago, Illinois. USA. Nelson, M.E., and L.D. Mech. 1991. Wolf predation risk associated with white-tailed deer movements. Canadian Journal of Zoology 69: 2696-2699. https://doi.org/10. 1139/z91-379 Nelson, M.E., and L.D. Mech. 1999. Twenty-year home range dynamics of a white-tailed deer matriline. Cana- dian Journal of Zoology 77: 1128-1135. https://doi.org/ 10.1139/z99-085 Nelson, M.E., and L.D. Mech. 2006. A 3-decade dearth of deer (Odocoileus virginianus) in a wolf (Canis lupus)- dominated ecosystems. American Midland Naturalist 2019 155: 373-382. https://doi.org/10.1674/00 03-0031(2006) 155[373:addodo]2.0.co;2 Nelson, M.E., L.D. Mech, and P.F. Frame. 2004. Tracking of white-tailed deer migration by global positioning sys- tem. Journal of Mammalogy 85: 505-510. https://doi. org/10.1644/BOS-120 Olson, S.F. 1938. A study in predatory relationship with particular reference to the wolf. Scientific Monthly 46: 323-336. Ozoga, J.J. 1968. Variations in microclimate in a con- ifer swamp deeryard in northern Michigan. Journal of Wildlife Management 32: 574-585. https://doi.org/10. 2307/3798938 Peek, J.M., D.L. Urich, and R.J. Mackiem. 1976. Moose habitat selection and relationships to forest management in north-eastern Minnesota. Wildlife Monographs 48: 1-65. Poszig, D., and J.B. Theberge. 2000. Gray wolf, Canis lupus lycaon, responses to shifts of white-tailed deer, Odocoileus virginianus, adjacent to Algonquin Provin- cial Park, Ontario. Canadian Field-Naturalist 114: 62— 71. Accessed 10 March 2020. https://www.biodiversity library.org/page/34236558. R Core Team. 2018. R: a language and environment for statistical computing. R Foundation for Statistical MECH AND BARBER-MEYER: MIGRATING DEER 351 Computing, Vienna, Austria. Severinghaus, C.W., and E.L. Cheatum. 1956. Life and times of the white-tailed deer. Pages 57-186 in The Deer of North America. Edited by W.P. Taylor. Stackpole Books, Harrisburg, Pennsylvania, USA. Silver, H., N.F. Colovos, J.B. Holter, and H.H. Hayes. 1969. Fasting metabolism of white-tailed deer. Journal of Wildlife Management 33: 490—498. https://doi.org/ 10.2307/3799370 Statistix 9. 2008. Analytical software. Tallahassee, Flor- ida, USA. Stenlund, M.H. 1955. A field study of the timber wolf (Canis /upus) on the Superior National Forest. Minnesota De- partment of Conservation, Technical Bulletin No. 2, Minneapolis, Minnesota, USA. Theberge, J.B., and M.T. Theberge. 2004. The wolves of Algonquin Park: a 12-year ecological study. Publication Series No. 56, University of Waterloo, Waterloo, On- tario, Canada. Townsend, M.T., and M.W. Smith. 1933. The white-tailed deer of the Adirondacks. Roosevelt Wildlife Bulletin 6: 161-325. Received 10 September 2018 Accepted 23 January 2020 The Canadian Field-Naturalist Nest site characteristics of cavity-nesting birds on a small island, in Haida Gwaii, British Columbia, Canada NEIL G. PILGRIM", JOANNA L. SmitH!?, KeErrH Moore, and ANTHONY J. GASTON! 'Laskeek Bay Conservation Society, P.O. Box 867, Queen Charlotte City, British Columbia VOT 1S0 Canada *Nature United, 366 Adelaide Street East, Suite 331, Toronto, Ontario MSA 3X9 Canada “Corresponding author: biologist@laskeekbay.org Pilgrim, N.G., J.L. Smith, K. Moore, and A.J. Gaston. 2019. Nest site characteristics of cavity-nesting birds on a small island, in Haida Gwaii, British Columbia, Canada. Canadian Field-Naturalist 133(4): 352-363. https://doi.org/10.22621/cfn. v13314.2277 Abstract Many studies of cavity-nesting birds in North America are conducted in large continental forests and much less is known about them in island ecosystems. We describe a 29-year study of tree species, nest site characteristics, and fledge dates of cavity-nesting birds ona small island in Haida Gwaii, British Columbia (BC). Seven cavity-nesting bird species were docu- mented on East Limestone Island and 463 nests were found in 173 different trees. Nest trees were significantly taller and had a greater diameter than a random sample of snags. Tree height did not differ among bird species but diameter at breast height was larger for trees used by Brown Creeper (Certhia americana) than for other species. Cavity-nesters selected tree decay classes 2—7 (all dead/near dead [snags]), with 85% in decay class 4 (35%) or 5 (50%), similar to the random snag sam- ple (class 4, 32%; class 5, 42%). Cavity height ranged from 2.6 to 44.9 m and for all species, except Brown Creeper, the mean nest height was >60% of the mean tree height. Nest heights were generally greater than observed elsewhere in BC. Nest cavity orientation was random except for Red-breasted Sapsuckers (Sphyrapicus ruber), for which only 13% of the cavity entrances faced southeast. Median fledging dates ranged from 7 June (Chestnut-backed Chickadee [Poecile rufescens]) to 28 June (Northern Flicker [Colaptes auratus]). Estimated median dates of clutch completion were similar for all species. Our results show that large snags provide habitat for a high diversity of cavity-nesting birds on Haida Gwaii. Key words: Wildlife trees; cavity-nesters; excavators; nest site; timing of breeding Introduction Dead and dying trees are essential for creating high quality nest sites for cavity-nesting birds (Li and Martin 1991) and primary excavators (those spe- cies that normally excavate new nest sites each year) are essential to many secondary species in providing the necessary conditions for them to nest or find shel- ter (Aitken and Martin 2007). Many factors can con- tribute to nest-site quality including tree height, nest height, nest-hole orientation, and the state of tree de- cay (McClelland and Frissell 1975; Inouye 1976). The selection of a nest tree and characteristics of nest sites are known to contribute to the reproductive success of cavity-nesters by affording protection for the breeder and their offspring from predators and improved microclimate in the nest cavity (Von Haartman 1957; Wesolowski 2002; Maziarz and Wesolowski 2013). Cavity-nesting birds can be divided into three groups related to how they acquire their cavity: (1) primary cavity-nesters excavate their own holes in live or dead trees and typically excavate a new hole each year, (2) secondary cavity-nesters use holes ex- cavated by other species (usually primary cavity nest- ers), use a naturally occurring hole and may re-use nests, and (3) weak cavity-nesters either make their own hole in a heavily decaying tree, nest in a cay- ity excavated by another species, or expand a natur- ally occurring hole. A bark nester, Brown Creeper (Certhia americana), has also been included in this paper, though it mainly nests under loose bark (Davis 1978). Nest site characteristics vary among and within bird species by geographic location and forest type (Scott et al. 1977; Newton 1994). Characteristics of cavity-nests most often reported include nest tree species, height, diameter, state of decay (or decay class), height of the nest site above the ground, and the cardinal direction of the cavity entrance. Most studies examined these characteristics for contin- ental forests (e.g., Carlson et al. 1998; Martin et al. 2004; Vaillancourt et a/. 2008), usually in relation to forest management guidelines in order to maintain stand structure to support cavity-nesters (e.g., Steeger and Dulisse 2002). Few studies have examined these A contribution towards the cost of this publication has been provided by the Thomas Manning Memorial Fund of the Ottawa Field-Naturalists’ Club. 352 ©The authors. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 characteristics in small island ecosystems. These may differ from continental ecosystems in having fewer suitable nest sites due to the limited forest area available and/or having more pronounced edge to in- terior effects, thus increasing risks from predation. The purpose of our study was to identify nest site characteristics of cavities as well as timing of breed- ing for all regularly-occurring cavity-nesting species on a small island in Haida Gwaii. Nest site charac- teristics were measured within the island’s mature forest ecosystem by researchers and citizen scien- tists of the Laskeek Bay Conservation Society (http:// www.laskeekbay.org), a local non-profit organization founded in 1990, with a well-established annual field program. We examined characteristics of nest trees selected by cavity-nesters and compared them with a random selection of available trees. We also compare the results from our island study to other ecosystems and discuss the likely selection pressures governing nest site choice in this ecosystem. Study Area Data were collected on East Limestone Island, Haida Gwaii, British Columbia (BC), Canada (52.90747°N, 131.613°W), a 48 ha island located in Laskeek Bay in the K’unna Gwaay heritage site/conservancy. It is ad- jacent to the southeast tip of Louise Island (27 200 ha) and is separated from it by only 400 m at the closest point. The island is mostly flat, or gently sloping, with the highest point of elevation being 65 m on the south ridge. Elevation gradients are most prominent along the east and west coasts where multiple coves lead to the sea via steep slopes. The northern coast of the is- land is the site of a large cove that encompasses most of that coast. East Limestone Island is in the Coastal Western Hemlock Zone, wet Hypermaritime subzone, a BC biogeoclimatic category characterized by cool win- ters and mild, cool, wet summers with periodic dry warm spells (Banner et a/. 2014). Strong winds are common and form an important climatic feature. Rainfall can exceed 1000 mm annually. The forest is primarily dominated by mature Sitka Spruce (Picea sitchensis (Bongard) Carriere), Western Hemlock (Tsuga heterophylla (Rafinesque) Sargent), and Western Red Cedar (Thuja plicata Donn ex D. Don). Red Alder (A/nus rubra Bongard), Pacific Crabapple (Malus fusca (Rafinesque) C.K. Schneider), Sitka Alder (Alnus alnobetula ssp. sinuata (Regal) Raus), and Scouler’s Willow (Salix scouleriana Barratt ex Hooker) are present along the shoreline and in a few places within the interior forest. The under- storey is sparse due to shade from the mature trees and intense browsing by the invasive Black-tailed Deer (Odocoileus hemionus, Stockton et al. 2005). PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 3953 Although shrubs are sparse, they occur through- out the island and include Vaccinium species (Red Huckleberry [Vaccinium parvifolium Smith], Oval- leaved Blueberry [Vaccinium ovalifolium Smith]), Salal (Gaultheria shallon Pursh), and Red Elderberry (Sambucus racemosa L.). The forest has not been commercially logged and most trees have been estimated to be more than 100 years old (K.M. pers. obs.). Like most temperate coastal old-growth systems, wind is a major factor for disturbance on the island, with windthrow the most common reason for gap creation and tree mortality (Pojar and MacKinnon 1994), in part due to shallow soils and high edge-interior effects. In 2010, a major windstorm hit Laskeek Bay and ~50% of the forest on East Limestone Island was blown down, resulting in high mortality for mature Western Hemlock and Sitka Spruce. Methods Nest location and monitoring Between 1991 and 2018, staff and volunteers of the Laskeek Bay Conservation Society searched for and recorded cavity-nests on East Limestone Island. Observations were made of the tree characteris- tics, the nest cavities, and the species that occupied them. This comprised the “wildlife tree monitoring program”, a citizen science effort involving numer- ous staff and volunteers each year from 1990 to 2018. Observations were made throughout May and June in all years and up to 9 July in all but five years (1990— 1992, 2002, 2003, and 2011). From the beginning of the monitoring program, trees containing active nests were tagged with unique numbers and mapped. In 1990, observations were incidental to other work. The next year a systematic methodology to detect oc- cupied breeding sites was designed and occurred an- nually using a written protocol. From 1991 to 1995, nests were located by listening for begging chicks during the nestling period. From 1996 onwards, all trees used at least once during the previous five years were included in that year’s sample of nest trees and observed three times for 30 min in late April or May during the nest building, egg laying, and incubation phases of breeding. The observations were made, generally, within a few days of each other by one or two observers with binoculars situated at least 15 m from the nest tree. If no activity was observed after these three visits the tree was considered inactive for that season. If activity was observed, the tree was considered active and checked for 30 min every three days during June for evidence of breeding activity (e.g., adults feeding nestlings or chicks calling). Once chicks were heard calling, nests were checked every two days for 30 min (weather permitting) to deter- 354 mine when chick calling ceased, assumed to be a sign that the nestlings had fledged. Up to three times per season (late-May to mid-July) a survey of the entire 45 ha island was conducted to locate any new nest sites; the island was divided into four quadrants and four to six observers would spend several hours mov- ing slowly throughout them, watching and listening for cavity nesting birds. Once active nests were con- firmed and chicks were being fed, all the remaining wildlife trees that had been surveyed earlier in the season were visited again and monitored for 10 min to confirm vacancy—ensuring that no active nests had been missed. This protocol was thought to have a very high chance of success for the primary cav- ity nesters, as all have young that call loudly from the nest site and in every year, four to six observers were present on the island throughout the nesting sea- son. However, our inventory was not likely to be com- plete for the other species, especially Brown Creeper, which has rather quiet young. All new nest trees were numbered, added to the monitored nest inventory, nest site characteristics measured and recorded, and location mapped. At the end of each season, any nest tree that had been inactive for five seasons was re- moved from the “active” inventory. Fledging date was assigned to the average of the last date when chicks were seen or heard and the first date with no sound or visuals. Sightings of fledg- lings out of the nest were also used as an indication of fledging date. For species with 10 or more rec- ords of active nest sites on the island, the dates of the onset of incubation were estimated by taking the es- timated date of fledging from the field surveys and subtracting incubation and fledging periods provided Descrip- | Live/healthy; no| Live/unhealthy;| Dead; recently tion | decay or structural| internal decay or | dead, needles or growth deformities} fine twigs present. or other structural damage (including stem damage, Dead; no needles/twigs; 50% of branches lost; only larger limbs remain; often loose bark. dead or broken tops); dying tree. THE CANADIAN FIELD-NATURALIST Dead; most branches/bark absent; some internal decay. Vol. 133 by the relevant species accounts in the Birds of North America (https://birdsna.org/Species-Account/bna/). Durations of incubation and fledging periods applied are given in Appendix 1. Multiple characteristics were noted for each active nest tree: bird species, tree species, total tree height (m), percent cover bark (main stem), tree classifica- tion (including number of bracket fungi; see Guy and Manning 1995), nest cavity entrance height, tree diameter at breast height (dbh), and nest cav- ity orientation. These characteristics were recorded when a hole was first discovered and subsequently if any changes occurred (e.g., tree height). In this paper, we use the BC Tree Classification System (Guy and Manning 1995) to determine the current level of decay of each tree when first used. The BC Tree Classification System has nine categories, ran- ging from 1—live/healthy to 9—debris (Figure 1). The term snag refers to a standing dead or dying tree. British Columbia’s Tree Classification class 2 is live/unhealthy and, in this paper, will be referred to as a snag. All the characteristics listed in BC Tree Classification System (Figure 1) were used to deter- mine what decay class a snag was considered to be. If characteristics of different decay classes were found in one snag, the snag was classified according to the maximum number of characteristics. Random sample of available nest trees In July 2004, an island-wide survey was carried out to obtain a random sample of all possible trees available for cavity-nesters in decay class 2 or higher. We selected random trees at 50 m intervals along the two main trails on the island. At each interval, we took a 90° bearing, perpendicular to the trail, and DEAD FALLEN approx. 2/3 original height : Dead; very litle | Dead; extensive internal decay; outer | Debris; downed branches or bark; | shell may be hard: lateral roots usually | trees or stumps. sapwood/heart- | completely decomposed; hollow or wood may be nearly hallow shells. sloughing from upper bole; decay more advanced: lateral roots of larger frees usually softening. approx. 1/2 original height approx. 1/3 original height Figure 1. British Columbia’s Tree Classification System (Guy and Manning 1995). 2019 laid out a 20 m transect, measuring all dead/near dead (decay class 2—8) trees that fell within ~5 m of either side of the transect. The same characteristics were re- corded for these trees that were recorded for the oc- cupied nest trees. Statistical analysis Five cavity-nesting species for which sample sizes were more than five were used in statistical compari- sons: Red-breasted Sapsucker (Sphyrapicus ruber), Hairy Woodpecker (Dryobates villosus), Northern Flicker (Colaptes auratus), Chestnut-backed Chicka- dee (Poecile rufescens), and Brown Creeper. The first three species are primary cavity-nesters and the fourth and fifth are, respectively, a weak excavator and a bark nester. For the analysis of tree characteris- tics used by cavity-nesters—tree height, tree species, dbh, and state of decay—we used the characteristics as described the first time that a tree was found in use by each bird species, regardless of how many years a nest tree was active. When analyzing individual nest height and orientation, all nests across all years were analyzed. Most statistical analysis were conducted using PAST3 (Hammer ef al. 2001) for Mac OSX: analy- sis of variance (ANOVA) with Tukey’s pairwise tests were used to compare tree height, nest height, and dbh among the cavity-nesting species. A two-sample t-test was used to compare the trees used by cavity- nesting species to a random sample of trees of similar decay class for tree height and nest height. Statistical program R version 3.4.3 (R Core Team 2017) was PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 35D used to conduct a Rayleigh test of uniformity to com- pare nest hole orientations among species. Means are given + | SD. Some data were not recorded for some nests, so that sample sizes are not the same for all analyses. Results During our study, the island supported seven cav- ity nesting birds: three primary cavity nesters: Red- breasted Sapsucker, Hairy Woodpecker, and Northern Flicker; two weak excavators: Chestnut-backed Chickadee and Red-breasted Nuthatch (Sitta can- adensis), a bark nester: Brown Creeper; and a second- ary cavity nester: Northern Saw-whet Owl (Aegolius acadicus). A total of 463 nests were found in 173 dif- ferent trees: Red-breasted Sapsucker (n = 344), Hairy Woodpecker (33), Northern Flicker (9), Chestnut- backed Chickadee (47), Red-breasted Nuthatch ( 9), Brown Creeper (19); and Northern Saw-whet Owl (2). The main excavator on the island was overwhelm- ingly Red-breasted Sapsucker, which occupied 74% of the cavity nests found. Tree characteristics We located and tagged 173 trees used by cavity- nesting birds between 1990 and 2018 (Table 1). Most of the cavity-bearing trees were Sitka Spruce (60%) or Western Hemlock (32%) with a small percentage of Red Alder (3%) and Western Red Cedar (1%), and a few of unknown identity, either because the spe- cies were missing in data records or the decay class did not allow species determination (4%; Table 2). TABLE 1. Mean, SD, minimum and maximum tree heights, and tree diameters of bark nesting (Brown Creeper [Certhia americana]) and cavity-nesting birds and random sample of snags on East Limestone Island, Haida Gwaii, from 1990 to 2018. n Mean Red-breasted Sapsucker (Sphyrapicus ruber) ie ae Hairy Woodpecker (Dryobates villosus) a4 207 Northern Flicker (Colaptes auratus) 7 et Chestnut-backed Chickadee (Poecile rufescens) oe — Red-breasted Nuthatch 5 415 (Sitta canadensis) Northern Saw-whet Owl 5 16 (Aegolius acadicus) Brown Creeper (Certhia americana) 16 a All cavity-bearing trees* 173 21.7 Random selection of snags 100 12.6 Tree height (m) Tree diameter (dbh; cm) SD Range Mean SD Range 10.4 7.2-52.8 104 40 40-260 97 3.8-40.8 93 32 50-200 6.0 14.1-32.8 93 4] 46-170 12.8 5.1-46.6 119 57 31-240 7.5 15.4—33.9 104 32 68-154 3.6 10.0-15.1 96 49 61-130 13.7 7.2-58.5 133 54 54-260 |e 3.8-58.5 104 43 31-260 1.1 1.3-63.3 62 46 11-229 *Total number of nest trees used throughout the study. These trees were used more than once by various bird species. 356 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 2. Percentages of tree species used by various cavity-nesting birds on East Limestone Island, Haida Gwaii. Sitka Spruce a ace Riedie Western Red Species (Picea (Touga (hinnsrubeey Cedar Unknown sitchensis) heterophylla) (Thuja plicata) Red-breasted Sapsucker (” = 130) 56.9 36.9 15 00 46 (Sphyrapicus ruber) Hairy Woodpecker (n = 26) (Drypebiies willosuss 65.4 34.6 0.0 0.0 3.9 Northern Flicker (” = 8) 75.0 125 125 00 00 (Colaptes auratus) Chestnut-backed Chickadee (n = 29) 793 172 35 0.0 0.0 (Poecile rufescens) Red-breasted Nuthatch (n = 5) 60.0 40.0 0.0 00 0.0 (Sitta canadensis) ; Brown Creeper (” = 16) (Certhia americana) yon ie oe oS Northern Saw-whet Owl (n = 2) 100.0 0.0 0.0 0.0 0.0 (Aegolius acadicus) All cavity-bearing trees (n = 173) 59.5 31.8 29 12 4.6 Random selection of snags (7 = 100) 64.0 31.0 4.0 1.0 0.0 The percentage distribution of cavity trees was simi- lar to the distribution of a random selection of snags throughout the island (Sitka Spruce = 64%, Western Hemlock = 31%, Red Alder = 4%, Western Red Cedar = 1%). Among cavity-nesting species with five or more active nest trees, use of Sitka Spruce ranged from 57 to 79%, Western Hemlock from 13 to 40%, Red Alder from 2 to 13%, and Western Red Cedar from | to 6%. There was no evidence of inter-species differences in nesting tree preference (Table 2). The nest trees of Red-breasted Sapsucker, Hairy Woodpecker, Chestnut-backed Chickadee, and Brown Creeper were significantly taller and larger in diam- eter than a random sample of snags on the island (Table 3); Northern Flicker nest trees were taller but not significantly larger in diameter. Nest tree height did not differ significantly among the bird species (ANOVA F595 = 0.27, P = 0.93), but diameter was significantly different among species (Fy, = 2.44, P = 0.04), with Brown Creeper using trees with sig- nificantly larger diameter than Hairy Woodpecker (Tukey’s pairwise: P < 0.05). Height and diameter were positively correlated for both the active and the ran- domly selected snags (active: 774, = 0.37, P < 0.001; random r’, = 0.31, P < 0.01). Cavity-nesters used trees in decay classes 2 through 7 and showed a strong preference for decay classes 4 and 5 (Table 4, Figure 2); 50% of all active nest trees were in snags of decay class 5 and 35% were class 4. Trees in classes 4 (32%) and 5 (42%) were also the most common in the randomly selected snag sam- ple, but the proportion of snags in decay class 2 and 3 TABLE 3. Analyses comparing mean tree heights and diameters of nest trees to a random sample of snags (n = 100), East Limestone Island, Haida Gwaii. Tree height (m) Tree diameter (dbh; cm) Effect size n t P t P d* Red-breasted Sapsucker 126 70 <0.01 71 <0.01 2.94 (Sphyrapicus ruber) Hairy Woodpecker ePieyeihutaswvillonns) 27 3.4 < 0.01 3.2 < 0.01 2.88 Northern Flicker (Compieetaranis) 8 2.9 0.01 1.8 0.07 4.41 Chestnut-backed Chickadee 9 39 <0.01 54 <0.01 232 (Poecile rufescens) Brown Creeper 16 31 <0.01 5.5 <0.01 2.17 (Certhia americana) *Cohen’s 2019 TABLE 4. Decay classes of nest trees used by cavity-nest- ing birds and a random sample of snags on East Limestone Island, Haida Gwaii. Decay class Species n Mean SD_ Range Red-breasted Sapsucker 104 46 07°~«2-6 (Sphyrapicus ruber) Hairy Woodpecker = (Dryobates villosus) aca ged fre Northern Flicker (Colaptes auratus) Pie ae oe ee Chestnut-backed Chickadee 7 347 «(0773-6 (Poecile rufescens) Red-breasted Nuthatch (Sitta canadensis) i= ee Brown Creeper (Certhia americana) a ol eet Northern Saw-whet Owl > 45 #07 4-5 (Aegolius acadicus) All cavity-bearing trees 163 47 O8 2-7 Random selection of snags 100 45 09 3-7 in the random sample significantly exceeded the pro- portion among used trees (14% versus 3%, respect- ively; contingency test, y?, = 14.8, P < 0.001). Hence, it appears that primary cavity excavators preferred trees in a more advanced state of decay than those in the random sample. a Red-breasted Sapsucker 70 60 50 Chestnut-backed Chickadee Number of trees on PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 357 Cavity characteristics Nest cavity heights ranged from 2.6 to 44.9 m from the base of the tree (Table 5). The Northern Flicker and Chestnut-backed Chickadee nests were, on average, the highest of the cavity-nesting spe- cies at 19.0 and 18.0 m, respectively. The lowest nests were the Northern Saw-whet Owl, but only two nests were found during the study period. For all but Brown Creeper, the mean nest height was more than 60% of the mean tree height (Table 5); Brown Creeper mean nest height of 9.0 + 4.2 m was significantly lower than those of Red-breasted Sapsuckers, Hairy Woodpeckers, Northern Flickers, and Chestnut- backed Chickadees (Table 5). Entrance orientation was not statistically signifi- cant for most species (P > 0.05; Table 6) with the ex- ception of Red-breasted Sapsucker, for which fewer cavity openings than expected faced southeast (91°— 180°; 13% of nests, P = 0.01); however, sample sizes for other species were much smaller. Timing of breeding Breeding of cavity-nesting species ranged from 21 May to 9 July (Figure 3). For Red-breasted Sapsuckers, the most common cavity-nesting spe- cies on East Limestone Island, the annual median fledging dates spanned a 16-day period from 10 June. Chestnut-backed Chickadees were usually the first to fledge, with a median date of 7 June (Table 7). Northern Flicker had the latest median fledging Cc Hairy Woodpecker 70 60 50 d Brown Creeper Snag class FiGurE 2. Distribution of snag classes used by different species of cavity-nesters on East Limestone Island, British Columbia, Canada (only species with n > 10). 358 THE CANADIAN FIELD-NATURALIST Vol. 133 TABLE 5. Nest heights of cavity-nesting species on East Limestone Islands, Haida Gwaii compared to provincial data (Campbell et al. 1990, 1997). Nest height (m) Birds of BC % Species n ie Tree height Min—max Mean SD Range & (>50% range)* Red: bi eas ed Sapsiicker 91173" 17 3.8-44.9 76.2 1. 8-24 (3-9) (Sphyrapicus ruber) Hairy Woodpecker * (Drjebates villosis) 26 16.8 8.2 5.4-32.8 81.2 0.9-38 (2-6) Northern Flicker * (Coleptescatmatus) i 19.0 6.2 13.0-31.8 96.4 0-27 (<3) Chestnut-backed Chickadee % 18.0" 13 26-413 81] 0-26 (2-6) (Poecile rufescens) Red-breasted Nuthatch (Silat canadensis) 5 13.4 0.8 12.7-14.5 62.3 0.5—20 (3-6) Northern Saw-whet Owl > 93 0.4 90-96 73.8 2 513.5 (Aegolius acadicus) ria Brown Ciee Der 12 9.01 4.2 4.0-16.0 40.2 0.2-15 (2-6) (Certhia americana) “Differs significantly from + at P < 0.05, Duncan Multiple Range Test. ‘Range within which greater than 50% of nests occurred. TABLE 6. Number of cavity entrances facing northeast (NE; 1°—90°), southeast (SE; 91°-180°), southwest (SW; 181°—270°), and northwest (NW; 271°—0°) for four cavity-nesting species and P-values from a Rayleigh’s test for uniformity for their nest cavity entrance orientation (P < 0.05 for a Rayleigh’s test indicates clustering). Species NE Red-breasted Sapsucker 70 (Sphyrapicus ruber) Hairy Woodpecker 6 (Dryobates villosus) Chestnut-backed Chickadee 2 (Poecile rufescens) Brown Creeper 2 (Certhia americana) date of 28 June. Median dates for the start of incu- bation were estimated to fall between 6—9 May for the four species with the largest sample sizes (Table 7). No evidence of second broods was found for any species, but four fledging dates for Chestnut-backed Chickadees fell after 21 June, two weeks after the long-term median in early June suggesting that some chickadees either laid very late or replaced earlier failed broods. Discussion Tree species On East Limestone Island, cavity-nesting species primarily used spruce and hemlock trees for nest- ing and these were used in proportion to the avail- able snags on the island; very few nests were in Red Alder and only Brown Creeper was found in Western Red Cedar. This was not surprising as most alders on the island were young, small diameter trees that were SE SW NW the 30 Se) 64 0.0077 3 4 3 0.5854 1 4 2 0.5721 3 4 1 0.7109 not very tall or in a state of decay. In other parts of BC, deciduous trees are used by cavity-nesters, for example, Martin and Eadie (1999) and Martin et al. (2004) found 95% of cavities in the Cariboo-Chilcotin region of central interior BC were in Trembling Aspen (Populus tremuloides Michaux). The major- ity of these were created by Red-naped Sapsucker (Sphyrapicus nuchalis), Hairy Woodpecker, and Northern Flicker—a very similar primary excavator community to that on East Limestone Island. Tree height and cavity height Of the cavity-nesters with more than five active nest trees during the study period, the mean heights of nest trees were significantly higher than a random selection of snags, strongly suggesting that height is an important factor for the location of nest cavities on this island. In addition, all bird species except Brown Creeper created or used nest cavities in the top half of the tree. Nests were also generally much higher 2019 PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 359 a Red-breasted Sapsucker Cc Hairy Woodpecker 80 10 60 : ~ 40 a 4 = % AS 0 Bt Dees he, eae ee NP So ae Ne vO” s ne ate re wade <= SF We WM a OW OP SP NS oh SO ws” = we \ <” S LY we s we Ss S we oe s ~ Ss we Ss ny S N > . O b Chestnut-backed Chickadee d Brown Creeper c Cy 5 (15 ; A 5) < 10 3 55 2 Z, 1 0 0 i % BD BD Md mB .% BD wh % A BD AD AD 0 HD WD as ae Se oy oe oy” ay ~ > oo aw oe Ra ae Ro wv a ~ > a 8 SVP SSS POV SY SA OS NS SUS Sl NOS ww es Sw fre rr es SPR NE Date of observation FiGurE 3. Observed fledging dates for cavity-nesting species on East Limestone Island, Haida Gwail, British Columbia, Canada (1990-2018). TABLE 7. Estimated median incubation and median and extreme fledging dates for cavity-nesters on East Limestone Island, Haida Gwaii, 1990-2018. Species Estimated median start of incubation Red-breasted Sapsucker (Sphyrapicus ruber) 219 7 May Hairy Woodpecker (Dryobates villosus) 24 6 May Northern Flicker 5 ia! (Colaptes auratus) Chestnut-backed Chickadee (Poecile rufescens) 34 6 May Red-breasted Nuthatch 3 s (Sitta canadensis) Brown Creeper 10 9 May (Certhia americana) than those reported as ‘typical’ (>50% of nests) by Campbell et al. (1990, 1997), with mean nest heights on East Limestone Island more than twice the max- imum of the typical range elsewhere for all species except Red-breasted Sapsucker and Brown Creeper. The high nest sites on East Limestone Island could be a function of predation risk, with higher nests hav- ing lower risk (Kilham 1971; Nilsson 1984). The main potential nest predator of cavity-nesting birds was Red Squirrel (Jamiasciurus hudsonicus), while adults might have been susceptible to predation by Red-tailed Hawk (Buteo jamaicensis) and Sharp- Median date of Earliest fledging Latest fledging chick fledging date date 17 Jun 1 Jun 13 Jul 10 Jun 29 May 30 Jun 28 Jun 3 Jun 5 Jul 7 Jun 21 May 1 Jul 12 Jun 31 May 16 Jun 10 Jun 28 May 28 Jun shinned Hawk (Accipiter striatus), both of which occur on the island. Red Squirrel was introduced to Haida Gwaii in 1950 (Golumbia ef a/. 2008) and re- corded on East Limestone Island by 1983. The spe- cies 1S an active predator on songbird nests in the area (Martin and Joron 2003). It was the only po- tential predator seen entering nest cavities on East Limestone Island (A.J.G. pers. obs.). The density of squirrels on the island fluctuates significantly among years (Martin et al. 2008) and is high in comparison with nearby larger islands that have other mammal- ian predators (e.g., Pine Marten [Martes americana], 360 Black Bear [Ursus americanus]). One possible ex- planation for nest heights is that squirrels avoid tall trees denuded of leaves and branches to avoid avian predators, such as Red-tailed Hawk (visitors to East Limestone Island), or the resident Common Raven (Corvus corax). Furthermore, a nest near the top of a snag could result in less rainwater running into the cavity hole, compared with a cavity further down the tree (Conner 1975). Brown Creeper nests lower than other species and build cryptic nests behind bark or rotten wood. Unlike other cavity-nesters, Brown Creeper nest- lings do not call loudly from the nest when the par- ents are absent. Brown Creeper may depend on these cryptic habits to avoid detection and minimize pred- ation. As a predator of small mammals and birds (Rasmussen ef al. 2008), Northern Saw-whet Owl may be sufficiently intimidating to deter squirrels from entering their nests, which might explain why both the two owl nests found were much lower (9.0 m and 9.6 m) than the average for other species. Only three nests of the Haida Gwaii subspecies of Saw- whet Owl (Aegolius acadicus brooksii) had been found by 2008 (Rasmussen et al. 2008), one of which was on East Limestone Island. Data are too limited to know whether low nest sites are characteristic of this subspecies. However, elsewhere in BC the spe- cies uses holes at similar heights to those found on East Limestone Island (Campbell et a/. 1990). Tree diameter Red-breasted Sapsucker, Hairy Woodpecker, Chest- nut-backed Chickadee, and Brown Creeper all used trees with significantly greater mean dbh than that of randomly selected snags (Table 3), a finding also made by Martin et a/. (2004) in interior BC and by Raphael and White (1984) in the Sierra Nevada. Brown Creeper selected significantly larger tree diameters than those used by Hairy Woodpecker. Height and dbh are correlated so we cannot distinguish which has the greater influence of nest site choice. While height may confer protection from predation and bet- ter drainage, greater girth may allow for deeper nests or better thermal protection (O'Connor 1978; Van Balen 1984). In addition, a larger cavity size could increase space for nestlings, reducing competition among them when being fed (Slagsvold 1989). As a bark nesting species, Brown Creeper (Davis 1978) has different selection criteria from the other species. It tends to select trees with large sections of loose bark to nest underneath, perhaps more fre- quently available on larger diameter trees. The spe- cies also prefers large diameter trees for foraging (Poulin et al. 2008) and choosing their nest site close to their food source could be advantageous. THE CANADIAN FIELD-NATURALIST Vol. 133 Decay Trees used for cavities on East Limestone Island were mostly in an advanced state of decay, with mean decay classes ranging from 4.5—5.0 (Figure 2). These trees would have decayed heartwood with relatively hard sapwood. Such trees may be more suitable as nest trees due to the decayed heartwood being soft enough for easy excavation, with an outer shell of relatively strong sapwood surrounding and pro- tecting the nest cavity (Kilham 1971; Conner et al. 1976; Miller and Miller 1980). It is worth noting that the value of differing decay states of different species of trees is not adequately represented by the BC Tree Classification System (cf, Guy and Manning 1995). Trees may have a similar appearance but be harder or softer depending on their location. The location could be subject to different, perhaps stronger winds, or different climatic conditions, all of which would give the tree a different appearance, hence a differ- ent decay class. Cavity orientation Orientation was measured to understand nest site selection relative to microclimate. The orientation of Red-breasted Sapsucker cavity entrances was not ran- dom, perhaps because they attempt to regulate nest microclimate by orienting their nest entrances away from the prevalent southeast winds, which bring the heaviest rainfall to the island. In addition, the top- ography of the island allows for winds from this dir- ection to be funneled into the interior of the island, strengthening its effect and perhaps strengthening the effect of cavity orientation. The apparent lack of preferred cavity orientation among other species may be a result of small sample size. Additional research is needed for Saw-whet Owl, as well as Northern Flicker and Red-breasted Nuthatch to shed further light on the nest site preferences of these species. Timing of breeding All of our nesting dates fell within the ranges indi- cated by Campbell et al. (1990, 1997) for individual species. However, Campbell et al. (1990, 1997) indi- cated a longer season (early May to end of July) for all species found on East Limestone Island. It ap- pears that breeding on East Limestone Island var- ies little among species, with all initiating incubation in the first half of May, and most nesting completed by the end of June. One exception was the case of Red-breasted Sapsucker in 1999, when median fledg- ing was six days later than in the next latest year. Breeding of open nesting species was later and less successful in 1999 because of low temperatures as- sociated with a strong La Nifia event (Gaston et al. 2005) and this may also have caused the late breeding of the sapsuckers. 2019 Conclusion This 29-year study has provided insights into the significant characteristics of nest sites created or used by cavity-nesting birds on a small island in Haida Gwail. The results of this work suggest that a rich di- versity and healthy populations of cavity-nesting spe- cies can be supported on small islands with intact mature forests. The predominance of Red-breasted Sapsucker, a primary excavator, over other hole-nest- ing species, suggests that suitable holes are probably abundant for secondary species, such as chickadees and nuthatches, both of which used old sapsucker holes on occasion. On the mainland, cavity nests are found in a greater variety of trees, often in live de- ciduous trees at much lower heights. In future, when surveys are conducted on small islands it is important that attention is paid to the upper parts of large snags to ensure that cavity nests are not overlooked. Our re- sults support the proposal that the protection of large old snags within northwest coastal forest ecosystems is essential to providing a healthy community of cav- ity-nesting birds (Cockle et al. 2011). Author Contributions Writing — Conceptualization & Field Work Design: A.G., K.M., J.S.; Field Work Oversight: K.M., J.S., N.P.; Data Analysis & Original Draft: N.P. & AG; Writing — Review & Editing: all authors. Acknowledgements We thank Colin French and Andrea Lawrence for initiating this project and all the board members, science advisors, volunteers, and employees with Laskeek Bay Conservation Society who contributed to it through the past 29 years, as well as all those who have supported the continuing efforts of the soci- ety. We would also like to thank the Haida Nation and the Province of British Columbia for their support of Laskeek Bay Conservation Society. Literature Cited Aitken, K.E.H., and K. Martin. 2007. The importance of excavators in hole-nesting communities: availabil- ity and use of natural tree holes in old mixed forests of western Canada. Journal of Ornithology 148 (Suppl. 2): S425-S434. https://doi.org/10.1007/s10336-007-0166-9 Banner, A., W.H. MacKenzie, J. Pojar, A. MacKinnon, S.C. Saunders, and H. Klassen. 2014. A field guide to ecosystem classifications and identification for Haida Gwaii. Land Management Handbook 68. BC Forests, Lands and Natural Resource Operations, Victoria, Bri- tish Columbia, Canada. Campbell, R.W., N.K. Dawe, I. McTaggart-Cowan, J.M. Cooper, G.W. Kaiser, and M.C.E. McNall. 1990. The Birds of British Columbia, Volume 2. University of British Columbia Press, Vancouver, British Columbia, Canada. PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 361 Campbell, R.W., N.K. Dawe, I. McTaggart-Cowan, J.M. Cooper, G.W. Kaiser, M.C.E. McNall, and G.E.J. Smith. 1997. The Birds of British Columbia, Volume 3. University of British Columbia Press, Vancouver, British Columbia, Canada. Carlson, A., U. Sandstrém, and K. Olsson. 1998. Avail- ability and use of natural tree holes by cavity-nesting birds in a Swedish deciduous forest. Ardea 86: 109-119. Cockle, K.L., K. Martin, and T. Wesolowski. 2011. Woodpeckers, decay, and the future of cavity-nest- ing vertebrate communities worldwide. Frontiers in Ecology and the Environment 9: 377-382. https://doi. org/10.1890/110013 Conner, R.N. 1975. Orientation of entrances to woodpecker nest cavities. Auk 92: 371-374. https://doi.org/10.23 07/4084566 Conner, R.N., O.K. Miller, Jr., and C.S. Adkisson. 1976. Woodpecker dependence on trees infected by fungal heart rots. Wilson Bulletin 88: 575-581. Davis, C.M. 1978. A nesting study of the Brown Creeper. Living Bird 17: 237-263. Gaston, A.J., J.-L. Martin, and S. Allombert. 2005. Sea surface temperatures mediated by the El Nifio-Southern Oscillation affect birds breeding in temperate coastal rain forests. Avian Conservation and Ecology 1: 4. Accessed 12 March 2019. http://www.ace-eco.org/voll/iss 1/art4/. Golumbia, T.E., L. Bland, K. Moore, and P. Bartier. 2008. History and current status of introduced ver- tebrates on Haida Gwaii. Pages 8-31 in Lessons from the Islands: Introduced Species and What They Tell Us about How Ecosystems Work. Proceedings from the 2002 Research Group on Introduced Species Symposium. Canadian Wildlife Service, Environment Canada, Ottawa, Ontario, Canada. Guy, S., and T. Manning. 1995. Wildlife/danger tree as- sessor’s course workbook. Canada-British Columbia Partnership Agreement on Forest Resource Develop- ment. Ministry of Forests, Victoria, British Columbia, Canada. Hammer, O., D.A.T. Harper, and P.D. Ryan. 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4: 4 pp. Inouye, D.W. 1976. Nonrandom orientation of entrance holes to woodpecker nests in aspen trees. Condor 78: 101-102. https://doi.org/10.2307/1366924 Kilham, L. 1971. Reproductive behavior of yellow-bellied sapsuckers I. Preference for nesting in Fomes-infected aspens and nest hole interrelations with flying squirrels, raccoons, and other animals. Wilson Bulletin 83: 159-171. Li, P., and T.E. Martin. 1991. Nest-site selection and nest- ing success of cavity-nesting birds in high elevation for- est drainages. Auk 108: 405-418. Martin, K.L., K.E. Aitken, and K.L. Wiebe. 2004. Nest sites and nest webs for cavity-nesting communities in interior British Columbia, Canada: nest characteristics and niche partitioning. Condor 106: 5-19. https://doi. org/10.1093/condor/106.1.5 Martin, J.L., S. Allombert, and A.J. Gaston. 2008. The effects of deer and squirrels on forest birds: community structure, population density, and reproduction. Pages 362 93-99 in Lessons from the Islands: Introduced Species and What They Tell Us about How Ecosystems Work. Proceedings from the Research Group on Introduced Species 2002 Symposium. Canadian Wildlife Service, Environment Canada, Ottawa, Ontario, Canada. Martin, K.L., and J.M. Eadie. 1999. Nest webs: a com- munity-wide approach to the management and conser- vation of cavity-nesting forest birds. Forest Ecology and Management 115: 243-257. https://doi.org/10.1016/ s0378-1127(98)00403-4 Martin, J.L., and M. Joron. 2003. Nest predation in for- est birds: influence of predator type and predator’s habitat quality. Oikos 102: 641-653. https://doi.org/10. 1034/j.1600-0706.2003.12040.x Maziarz, M., and T. Wesolowski. 2013. Microclimate of tree cavities used by Great Tits (Parus major) in a pri- meval forest. Avian Biology Research 6: 47—56. https:// doi.org/10.3184/175815513x13611994806259 McClelland, B.R., and S.S. Frissell. 1975. Identifying forest snags useful for hole-nesting birds. Journal of Forestry 73: 414—417. Miller, E., and D.R. Miller. 1980. Snag use by birds. Pages 337-356 in Management of western forests and grass- lands for nongame birds. Workshop proceedings, Salt Lake City, Utah, 11-14 February 1980. Technical co- ordinator R.M. DeGraff. US Forest Service General Technical Report. INT-86, Ogden, Utah, USA. Newton, I. 1994. The role of nest sites in limiting the num- bers of hole-nesting birds: a review. Biological Con- servation 70: 265-276. https://doi.org/10.1016/0006-32 07(94)90172-4 Nilsson, S.G. 1984. The evolution of nest-site selection among hole-nesting birds: the importance of nest preda- tion and competition. Ornis Scandinavica 15: 167-175. https://doi.org/10.2307/3675958 O’Connor, R.J. 1978. Nest-box insulation and the timing of laying in the Wytham woods population of great tits Parus major. {bis 120: 534-537. https://doi.org/10.1111/ J.1474-919X.1978 tb06822.x Pojar, J., and A. MacKinnon. 1994. Plants of the Pacific Northwest Coast. Lone Pine Publishing, Vancouver, British Columbia, Canada. Poulin, J.F., M.A. Villard, M. Edman, P.J. Goulet, and A.M. Eriksson. 2008. Thresholds in nesting habi- tat requirements of an old forest specialist, the Brown Creeper (Certhia americana), as conservation targets. Biological Conservation 141: 1129-1137. https://doi.org/ 10.1016/j.biocon.2008.02.012 R Core Team. 2017. R: a language and environment for sta- THE CANADIAN FIELD-NATURALIST Vol. 133 tistical computing. R Foundation for Statistical Com- puting, Vienna, Austria. Raphael, M.G., and M. White. 1984. Use of snags by cavity-nesting birds in the Sierra Nevada. Wildlife Monographs 86: 3—66. Rasmussen, J.L., S.G. Sealy, and R.J. Cannings. 2008. Northern Saw-whet Owl (4egolius acadicus), The Birds of North America Version 2.0. Cornell Lab of Ornithology, Ithaca, New York, USA. Accessed 9 March 2019. https://birdsna.org/Species-Account/bna/ species/nswowl. Scott, V.E., K.E. Evans, D.R. Patton, and C.P. Stone. 1977. Cavity-nesting birds of North American for- ests (No. 511). Forest Service, US Department of Agriculture, Washington, DC, USA. Slagsvold, T. 1989. On the evolution of clutch size and nest size in passerine birds. Oecologia 79: 300-305. https:// doi.org/10.1007/BF00384308 Steeger, C., and J. Dulisse. 2002. Characteristics and dy- namics of cavity-nest trees in southern British Columbia. Pages 275-289 in Proceedings of the symposium on the ecology and management of dead wood in west- ern forests. Edited by W.F. Laudenslayer, P.J. Shea, C.P. Witherspoon, and T.E. Lisk. USDA Forest Service, General Technical Report, PSW-GTR-181, Albany, California, USA. Stockton, S.A., S. Allombert, A.J. Gaston, and J.L. Martin. 2005. A natural experiment on the effects of high deer densities on the native flora of coastal temper- ate rain forests. Biological Conservation 126: 118-128. https://doi.org/10.1016/j.biocon.2005.06.006 Vaillancourt, M.A., P. Drapeau, S. Gauthier, and M. Robert. 2008. Availability of standing trees for large cavity-nesting birds in the eastern boreal forest of Québec, Canada. Forest Ecology and Management 255: 2272-2285. https://doi.org/10.1016/j.foreco.2007.12.036 Van Balen, J.H. 1984. The relationship between nest-box size, occupation and breeding parameters of the Great Tit Parus major and some other hole-nesting species. Ardea 72: 163-175. Von Haartman, L. 1957. Adaptation in hole-nesting birds. Evolution 11: 339-347. https://do1.org/10.1111/).1558-56 46.1957.tb02902.x Wesolowski, T. 2002. Anti-predator adaptations in nest- ing marsh tits Parus palustris: the role of nest-site se- curity. Ibis 144: 593-601. https://doi.org/10.1046/).14 74-919x.2002.00087.x Received 30 April 2019 Accepted 24 December 2019 2019 PILGRIM ET AL.: CAVITY-NESTING ON A SMALL ISLAND 363 APPENDIX 1. Incubation and fledging periods used in estimating dates of clutch completion for species with 10 or more records. Species Incubation period (days) Hairy Woodpecker B (Dryobates villosus) Red-breasted Sapsucker 14 (Sphyrapicus ruber) Chestnut-backed Chickadee B (Poecile rufescens) Brown Creeper 15 (Certhia americana) Literature Cited Jackson, J.A., H.R. Ouellet, and B.J. Jackson. 2018. Hairy Woodpecker (Dryobates villosus), Version 1.1. In The Birds of North America. Edited by P.G. Rodewald. Cornell Lab of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.haiwoo.01.1 Dahlsten, D.L., L.A. Brennan, D.A. McCallum, and S.L. Gaunt. 2002. Chestnut-backed Chickadee (Poecile ru- fescens), Version 2.0. In The Birds of North America. Edited by A.F. Poole and F.B. Gill. Cornell Lab of Or- nithology, Ithaca, New York, USA. https://doi.org/10. 2173/bna.689 Fledging period (days) Reference 29 Jackson et al. 2018 27 Walters et al. 2014 20 Dahlsten et al. 2002 17 Poulin et al. 2013 Poulin, J.F., E. D’Astous, M. Villard, S.J. Hejl, K.R. Newlon, M.E. McFadzen, J.S. Young, and C.K. Gha- lambor. 2013. Brown Creeper (Certhia americana), Version 2.0. Jn The Birds of North America. Edited by A.F. Poole. Cornell Lab of Ornithology, Ithaca, New York, USA. https://doi.org/10.2173/bna.669 Walters, E.L., E.H. Miller, and P.E. Lowther. 2014. Red- breasted Sapsucker (Sphyrapicus ruber), Version 2.0. In The Birds of North America. Edited by A.F. Poole. Cornell Lab of Ornithology, Ithaca, New York, USA. Accessed 9 March 2019. https://birdsna.org/Species- Account/bna/species/rebsap. The Canadian Field-Naturalist Rooting depth and below ground biomass in a freshwater coastal marsh invaded by European Reed (Phragmites australis) compared with remnant uninvaded sites at Long Point, Ontario CALVIN Ler’, SARAH J. YUCKIN!, and REBECCA C. ROONEY!“ 'Department of Biology, University of Waterloo, 200 University Avenue West, Waterloo, Ontario N2L 3G1 Canada “Corresponding author: rrooney@uwaterloo.ca Lei, C., S.J. Yukin, and R.C. Rooney. 2019. Rooting depth and below ground biomass in a freshwater coastal marsh invaded by European Reed (Phragmites australis) compared with remnant uninvaded sites at Long Point, Ontario. Canadian Field-Naturalist 133(4): 364-371. https://do1.org/10.22621/cfn.v133i4.2281 Abstract Invasive European Reed (Phragmites australis subsp. australis) outcompetes native vegetation, reducing floristic diversity and habitat value for wildlife. Research in coastal salt marshes has indicated that P. australis invasion may be facilitated by its relatively deep rooting depth, but in freshwater marshes the growth pattern of below ground tissues in relation to water depth is uncertain. To determine if P. australis is rooting more deeply than resident wetland plant species in a freshwater coastal marsh on Lake Erie, Ontario, we measured the vertical distribution of below ground biomass in P. australis invaded marsh sites and compared it to the below ground biomass distribution in nearby sites not yet invaded by P. australis. These invaded and uninvaded sites were paired by water depth, which is known to influence resource allocation and rooting depth. Below ground biomass in invaded sites was greater than in uninvaded sites (t,. = 3.528, P = 0.001), but rooting depth (..e., the depth at which 90% of total below ground biomass is accounted for) was comparable (¢,; = 0.992, P = 0.330). Using water depth and site type, general linear models could predict below ground biomass (F,;; = 9.115, P < 0.001) but not root- ing depth (F;,;;= 1.175, P = 0.316). Rooting depth is likely affected by other factors such as substrate type and the depth of the organic soil horizon. Key words: Below ground biomass; coastal marsh; Common Reed; ecosystem effects; invasive species; Lake Erie; rhizomes; roots; wetland Introduction European Reed (Phragmites australis (Cavanilles) Trinius ex Steudel subsp. australis) is considered highly invasive in North America (Saltonstall 2002) and has profound negative effects on both coastal and inland wetlands and shores. Researchers have reported that P. australis replaces native vegetation (Able et al. 2003; Tulbure and Johnston 2010), low- ers plant biodiversity (Keller 2000), and disrupts wet- land integrity and ecological function (Windham and Ehrenfeld 2003; Rothman and Bouchard 2007; Tulbure and Johnston 2010; Duke ef al. 2015). Phragmites australis invasion may also lead to sed- iment accretion, terrain flattening, and a reduction in water-filled depressions due to the accumulation of leaf litter and rhizome biomass (Able et a/. 2003). These invasion-driven changes in wetland habitat have consequences such as the loss of toad breeding habitat (Greenberg and Green 2013), reduced abun- dance of at-risk birds (Robichaud and Rooney 2017), and fewer suitable nesting areas and poor microhab- itats for turtle eggs (Bolton and Brooks 2010; Cook 2016). Consequently, P. australis was named the worst invasive plant species in Canada (Catling and Mitrow 2005, 2011). In the Great Lakes region, P. australis has replaced thousands of hectares of freshwater coastal wetlands. Around Lake Erie alone, invasion estimates range from 2553 ha within the coastal wetlands (Carson et al. 2018) to 8233 ha within a 10 km buffer around the American portion of Lake Erie (Bourgeau-Chavez et al. 2013). At Long Point on Lake Erie, P. australis in- vasion is predicted to continue expanding rapidly un- til 2022 (Jung et al. 2017), and the wetland commu- nities most commonly replaced are cattail, meadow marsh, sedge and grass hummocks, and other mixed emergent communities (Wilcox ef al. 2003). The invasion success of P. australis is due to ad- vantageous morphological features and its ability to modify its environment. For example, P. austra- lis stems can grow up to five metres tall, intercepting light and shading competitors (Hirtreiter and Potts 364 ©The authors. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 2012). With its large seed heads, P. australis can pro- duce hundreds of wind-dispersed seeds (Tulbure and Johnston 2010), which is an important strategy for cre- ating new individuals (Albert et al. 2015). However, local expansion mainly occurs by vegetative growth (Albert et al. 2015), using stolon fragments and rhi- zomes (Mal and Narine 2004; Tulbure and Johnston 2010). Rhizomes are also important storage organs that enable P. australis to send up spring ramets in advance of resident species and to manage nitrogen limitation (Granéli et al. 1992). Below ground, P. australis engineers its habitat to optimize its competitive advantage over native spe- cies (Minchinton ef a/. 2006). For example, a study on P. australis roots reported that hypodermal layers around roots and rhizomes protect against toxic or- ganic compounds and anoxia (Armstrong and Arm- strong 1988). Aerenchyma channels, which send at- mospheric oxygen from emergent plant tissues to plant parts in anoxic soils, also allow P. australis to sustain deep rooting depths (Armstrong and Arm- strong 1988). For example, studies from a marine coastal marsh gave estimates of P. australis roots growing from 30 cm water depth) or by meadow taxa, including graminoids, sedges, and forbs (<<40 cm water depth). Sites in the 30-40 cm depth range were either meadow marsh or Zypha spp. marsh, as the two communities stratify by depth and rarely mix. Sites were dispersed across the Crown 366 Marsh and Long Point Provincial Park management units, spaced between 100 and 2000 m apart. This area 1S representative of wet meadow and emergent lacustrine marsh in Lake Erie, with substrate rang- ing from organic in shallower depths to pure sand in deeper locations. Core collection Fieldwork was conducted in May 2017. Using a 2.54 cm diameter soil gouge auger, soil cores (0.3- 0.75 m deep) were sampled from sites invaded by P. australis and paired uninvaded marsh sites. It was not possible to obtain cores of uniform length due to differences in the thickness of the organic hori- zon and difficulties penetrating the underlying sand substrate. In total 29 pairs of cores were collected. The cores were then sub-sectioned into 10 cm long segments and frozen until they could be processed. For comparison, Moore et al. (2012) who also exam- ined belowground biomass trends in marsh invaded by European P. australis, collected 100 cm long cores from 10 tidal marshes along New Hampshire’s Atlantic coast using the same diameter gouge auger and sub-sectioned them into 5 cm long segments. Core processing Core segments were thawed for about 24 h and then washed over two nested sieves: a coarser (1.7 mm) sieve over a finer (425 um) sieve. All live rhi- zomes and all root tissues were retrieved and dried at 80°C to a constant weight (minimum 48 h). Dead roots may have been included in our weights as we did not find it possible to reliably differentiate live and dead roots. The dried below ground tissues were then weighed on a Mettler Toledo analytical balance (MS204S, Columbus, Ohio, USA) with a 0.0001 g ac- curacy. For comparison, Moore ef al. (2012) picked live roots and rhizomes from trays partially filled with water and then oven dried to a constant weight at 65°C for a minimum of 48 h. Data analysis For the purposes of this study, rooting depth was defined as the depth (cm) at which 90% of the cu- mulative below ground biomass was accounted for. Below ground biomass was defined as the total root and rhizome mass per unit area (g/m), recognizing that the core depths varied with the thickness of the organic horizon. To test whether below ground bio- mass was greater in P. australis invaded sites com- pared to uninvaded sites, we used a paired-samples, one-tailed 7-test. To test whether rooting depth was greater in P. australis invaded sites than uninvaded sites, we used another paired-samples, one-tailed t-test. Lastly, to test whether water depth is a signif- icant predictor of below ground biomass and rooting depth, we used general linear models (GLM) with THE CANADIAN FIELD-NATURALIST Vol. 133 a least squares estimation framework to model var- iation in below ground biomass and rooting depth based on water depth, site type (P. australis invaded or uninvaded), and their interaction. Models are thus represented by the general form: V=BW+ BT+B Tx W+B +, where W is water depth, 7 is site type, and ¢€ is error. If the interaction terms were not significant, the model would be re-run to only include the main factors: wa- ter depth and site type. In all cases, we used an alpha value of 0.05 and Type III sums of squares. Analyses were completed using IBM SPSS Statistics 20. Results Paired-samples t-tests for below ground biomass and rooting depth Phragmites australis invaded marsh had greater below ground biomass than uninvaded marsh habitat, when meadow and Zypha spp. marsh are considered jointly (paired-samples /-test, t,, = 3.528, P = 0.001; Figure la). Although the difference between Typha spp. dominated cattail marsh and P. australis invaded marsh is negligible, it revealed that the difference is primarily between P. australis invaded and meadow marsh sites (Figure la). There is no significant difference in rooting depth between P. australis invaded marsh and uninvaded marsh (paired-samples f-test, t,, = 0.992, P = 0.330; Figure 1b). This appears evident in both meadow marsh and cattail marsh components of the unin- vaded sites (Figure 1b). The down-core distribution of below ground bio- mass suggests that core depths were sufficient to cap- ture the bulk of total below ground tissues (Figure 2). This was true for invaded (Figure 2a) and uninvaded (Figure 2b) sites, across a range of water depth in- tervals between 13.7 and 55.7 cm. Below ground bio- mass was detected to a maximum of 80 cm soil depth yet peaked within the top 30 cm of the soil profile, re- gardless of site type (Figure 2). General linear models for below ground biomass and rooting depth For below ground biomass, the interaction term was not significant (Table Sla, Figure S1), so we re- moved it and re-ran the GLM as below ground biomass =B,W+ B,T +B, + €. This model provided a reasonable fit (adjusted r? = 0.222; GLM, F,;; = 9.115, P < 0.001; details in Table S1b). Likewise, for rooting depth, the interaction term was not significant (Table S2a), so we removed it and re-ran the GLM as rooting depth = B,W+ B,T + By + €. However, this model proved to be a poor predictor 2019 _—~ 8000 a £ a % 6000 & L Oo ne] =| 4000 o) = 5b 3 2 2000 oY a) 3 o Ee 0 it ee et Invaded Uninvaded Uninvaded Uninvaded meadow cattail Marsh vegetation type Rooting depth (cm) Invaded Uninvaded Uninvaded Uninvaded meadow cattail Marsh vegetation type FicurE 1. Total below ground biomass and rooting depth in European Reed (Phragmites australis) invaded and un- invaded marsh. Boxplots depicting a. total below ground biomass (g/m’) and b. rooting depth (cm), contrasting P. australis invaded marsh (dark grey; n = 29) and uninvaded marsh (white; 7 = 29) sites. Note that uninvaded marsh is divided into shallower depth meadow marsh (light grey; n = 15) and deeper water cattail (7ypha sp.) marsh (grey; n = 14) communities. of rooting depth (adjusted r? = 0.006; GLM, F;5; = 1.175, P =0.316; details in Table S2b). Discussion Our research objectives were to determine if P. australis invaded marsh produced more below ground biomass, and deeper rooting depths than un- invaded marsh in a freshwater coastal marsh, as has been observed in marine coastal marshes (e.g., Ravit et al. 2006; Moore et al. 2012). Controlling for wa- ter depth, we observed that P. australis invaded marsh had more below ground biomass than unin- vaded marsh, however, rooting depths did not differ significantly between P. australis invaded and unin- vaded marsh sites. Like site type, water depth was a significant predictor of below ground biomass (g/m?) but not of rooting depth. Interestingly, although the LEI ET AL.. BELOW GROUND CHANGES OF EUROPEAN REED 367 largest difference in below ground biomass was evi- dent between P. australis invaded sites and meadow marsh sites, which were restricted to shallower wa- ter depths, we detected no significant interaction be- tween water depth and site type when predicting ei- ther below ground biomass or rooting depth. Greater below ground biomass may _pro- vide P. australis a competitive advantage allow- ing it to usurp soil resources (van Wijk et al. 2003) and facilitate dispersion by vegetative reproduc- tion (Saltonstall 2002; Tulbure and Johnston 2010; Albert et al. 2015). The current literature reports below ground biomass values for P. australis in the range of 886 g/m” (Rothman and Bouchard 2007) to 1368 g/m? (Windham 2001); for cattail marsh in the range of 742 g/m? (Rothman and Bouchard 2007) to 2461 g/m? (Ouellet-Plamondon ef al. 2004); and for meadow species, such as Saltmeadow Cordgrass (Sporobolus pumilus (Roth) P.M. Peterson & Saarela) and Bluejoint Reedgrass (Calamagrostis canaden- sis (Michaux) Palisot de Beauvois), in the range of 256 g/m? (Ouellet-Plamondon et al. 2004) to 757 g/m? (Windham 2001). Our measures of below ground bi- omass show the same pattern in relative magnitude among the three communities but are noticeably higher than other published values: averaging 3137 g/ m? for P. australis, 2372 g/m? for cattail marsh, and 1146 g/m? for meadow marsh. Our measurements may be high due to particularly dense growth, favour- able edaphic conditions in intact freshwater coastal marsh, or because we were unable to differentiate live tissues from recently dead tissues. When uninvaded marsh was separated into cat- tail and meadow marsh communities, we noted higher average below ground biomass in uninvaded cattail marsh, clearly overlapping with the below ground bio- mass typical of P. australis. This indicates that the ef- fects of invasion on below ground biomass is likely more evident where P. australis replaces meadow marsh than where it invades cattail marsh. Yet, despite this difference in mean below ground biomass be- tween cattail and meadow marsh, we fit a single slope relating the below ground biomass of uninvaded sites to water depth collectively. Future research should ex- plicitly test for the role of resident vegetation commu- nity type on limiting the magnitude of P. australis in- vasion effects on invaded ecosystems. Importantly, though invasion by P. australis in freshwater coastal marsh may increase overall be- low ground biomass, the concern that P. australis below ground tissues might penetrate more deeply than resident species and thus alter nutrient and metal fluxes in freshwater marshes is unfounded. Contrary to previous studies (e.g., Ravit et al. 2006; Moore et al. 2012), we observed no difference in rooting depth 368 THE CANADIAN FIELD-NATURALIST Vol. 133 Water depth: <25 cm Invaded A Uninvaded b =2 0-10 0-10 —_ = CN n=2 = 10-20 = 10-20 2 7 oa) n=2 Ss 20-30 20-30 |_—_—_—_— o 539-40 fr 2 30-40 8 7 40-50 40-50 0 500 1000 1500 2000 2500 0 500 1000 1500 2000 2500 Average below ground biomass (g/m’) | Average below ground biomass (g/m) Water depth: 25—35 cm Invaded Uninvaded 0 200 400 600 800 1000 1200 0 200 400 600 800 Average below ground biomass (g/m’) Average below ground biomass (g/m’) Water depth: 35—45 cm Invaded e Uninvaded f 0-10 10-20 20-30 = 30-40 S 40-50 350-60 60-70 70-80 0 200 400 600 800 100012001400 0 200 400 600 800 100012001400 Average below ground biomass (g/m’) Average below ground biomass (g/m’) Water depth: >45 cm Invaded Uninvaded pl __ a a es h 0-10 — ae — 1) 0-10 — 210-20 {fi |e ass 379-30 S 10-20 | | o = <= n=3 3.3040 = 20-30 | A 40-50 I - ‘a e n=2 50-60 | 30-40 ii 0 1000 2000 3000 4000 5000 0 100 200 300 400 500 600 Average below ground biomass (g/m ) Average below ground biomass (g/m’) FiGurRE 2. Down-core distribution of below ground biomass at different water depths. Down-core distribution of below ground biomass, contrasting European Reed (Phragmites australis) invaded sites (a, c, e, g) and uninvaded sites (b, d, f, h) at different water depth intervals: <25 cm (a, b), between 25-35 cm (c, d), between 35—45 cm (e, f), and >45 cm (g, h). The 7 above each bar indicates the number of cores in which living below ground tissues were detected at the indicated water and soil depth, in the indicated site type. Error bars are SE. 2019 among the invaded and uninvaded sites. Moore et al. (2012) surmised that in marine coastal marsh, P. aus- tralis may produce deeper roots to access freshwater pockets. If this were so, it might explain why P. aus- tralis was not rooting more deeply than resident spe- cies in our freshwater coastal marsh. Alternatively, these published studies may differ from ours in the frequency and amplitude of water depth fluctuations that can also influence rooting depth (Hanslin ef al. 2017). Another important factor is likely the wetland soil type and stratigraphy. Moore ef al. (2012) re- ported that sandy mineral soils may inhibit deep pen- etration of roots. Long Point has a sand mineral soil beneath an organic horizon of variable thickness; the sand soil may have limited rooting depth for all spe- cies in our study. Because P. australis produces significantly more below ground biomass in the same depth of rhizos- phere as resident vegetation communities, we expect that root processes such as enhanced gas diffusion in the rhizosphere, oxidation of waterlogged anoxic soils (Armstrong and Armstrong 1988; Bart and Hartmann 2000), and the release of allelochemicals (Rudrappa et al. 2007) provide P. australis a compet- itive advantage and contribute to its invasion success. Yet clearly, given the equivalent rooting depths of P. australis invaded and uninvaded marsh, these com- munities experience a common rooting depth limit. This conclusion is further supported by our observa- tion that meadow marsh, despite producing less be- low ground biomass than cattail marsh, nonetheless roots at an equivalent depth Conclusion Below ground biomass in P. australis invaded marsh significantly exceeded that in resident com- munities of meadow marsh and cattail marsh, af- ter accounting for water depth, but rooting depths were equivalent. Consequently, root densities must be greater in P. australis invaded marsh, potentially contributing to its invasion success in Long Point. Because P. australis did not root more deeply than resident vegetation in our freshwater coastal marsh study system, concerns around invasion mobilizing deep pools of otherwise inactive carbon or metals may be generally unwarranted. The novel quantita- tive data presented in this study increases our un- derstanding of P. australis invasion in freshwater lacustrine coastal marsh habitat and establishes the hypothesis of common limits to rooting depth in in- vaded and uninvaded sites that should be tested in other study systems. Author Contributions Conception & Design: R.C.R., S.J-Y., and C.L.; Field Work: S.J-Y.; Lab Work: C.L. and S.J-Y.; Data LEI ET AL.. BELOW GROUND CHANGES OF EUROPEAN REED 369 Analysis & Interpretation: R.C.R., S.J-Y., and C.L.; Writing — First Draft: C.L.; Writing — Review & Editing: R.C.R., S.J-Y., and C.L.; Funding Acquisition: RGR: Acknowledgements This work was supported by NSERC Discovery Grant #RGPIN 2014-03846 to R.C.R., Mitacs Accel- erate, the Nature Conservancy of Canada, and the Ontario Graduate Scholarship program. We would like to thank Ministry of Natural Resources and Forestry Aylmer District and Ontario Parks for pro- viding site access. We are grateful to Courtney Robichaud, Jessie Pearson, Graham Howell, Taylor Blackwell, Madison Brook, Bailey Dhanani, Megan Jordan, Lauren Koiter, Cindy Luu, Christine Nielsen, and Alina Steele for their participation in conduct- ing the field and lab work necessary for this study. We thank Dr. Roland Hall for his insight and feedback on an early draft of this paper and Dr. Paul Catling for useful feedback during the review process. Literature Cited Able, K.W., S.M. Hagan, and S.A. Brown. 2003. Mechan- isms of marsh habitat alteration due to Phragmites: re- sponse of young-of-the-year mummichog (Fundulus heteroclitus) to treatment for Phragmites removal. Estuaries 26: 484—494. https://doi.org/10.1007/bf02823725 Albert, A., J. Brisson, F. Belzile, J. Turgeon, and C. Lavoie. 2015. Strategies for a successful plant invasion: the re- production of Phragmites australis in north-eastern North America. Journal of Ecology 103: 1529-1537. https:// doi.org/10.1111/1365-2745.12473 Armstrong, J., and W. Armstrong. 1988. Phragmites aus- tralis—a preliminary study of soil-oxidizing sites and internal gas transport pathways. New Phytologist 108: 373-382. https://do1.org/10.1111/).1469-8137.1988 tb04177.x Ball, H., J. Jalava, T. King, L. Maynard, B. Potter, and T. Pulfer. 2003. The Ontario Great Lakes coastal wet- land atlas: a summary of information (1983-1997). Environment Canada, Peterborough, Ontario, Canada. Bart, D., and J.M. Hartman. 2000. Environmental deter- minants of Phragmites australis expansion in a New Jersey salt marsh: an experimental approach. Oikos 89: 59-69. https://doi.org/10.1034/).1600-0706.2000.890 107.x Bolton, R.M., and R.J. Brooks. 2010. Impact of the sea- sonal invasion of Phragmites australis (Common Reed) on turtle reproductive success. Chelonian Conservation and Biology 9: 238-243. https://doi.org/10.2744/CCB- 0793.1 Bourgeau-Chavez, L.L., K.P. Kowalski, M.L. Carlson Mazur, K.A. Scarbrough, R.B. Powell, C.N. Brooks, B. Huberty, L.K. Jenkins, E.C. Banda, D.M. Gal- braith, Z.M. Laubach, and K. Riordan. 2013. Map- ping invasive Phragmites australis in the coastal Great Lakes with ALOS PALSAR satellite imagery for deci- sion support. Journal of Great Lakes Research 39: 65— 77. https://doi.org/10.1016/j.jglr.2012.11.001 370 Carson, B.D., S.C. Lishawa, N.C. Tuchman, A.M. Monks, B.A. Lawrence, and D.A. Albert. 2018. Harvesting in- vasive plants to reduce nutrient loads and produce bio- energy: an assessment of Great Lakes coastal wetlands. Ecosphere 9: 1-16. https://doi.org/10.1002/ecs2.2320 Catling, P.M., and G. Mitrow. 2005. A prioritized list of the invasive alien plants of natural habitats in Canada. Canadian Botanical Association Bulletin 38: 55-57. Catling, P.M., and G. Mitrow. 2011. Major invasive alien plants of natural habitats in Canada. 1. European Com- mon Reed, Phragmites australis (Cav.) Trin. ex Steud. subsp. australis. Canadian Botanical Association Bul- letin 44: 52-61. Cook, C. 2016. Impacts of invasive Phragmites australis on diamondback terrapin nesting. M.Sc. thesis, College of William and Mary, Williamsburg, Virginia, USA. Duke, S.T., S.N. Francoeur, and K.E. Judd. 2015. Ef- fects of Phragmites australis invasion on carbon dyna- mics in a freshwater marsh. Wetlands 35: 311-321. https://doi.org/10.1007/s13157-014-0619-x Faufier, A.C., J. DuSek, H. Cizkova, and M. Kazda. 2016. Diurnal dynamics of oxygen and carbon dioxide con- centrations in shoots and rhizomes of a perennial in a constructed wetland indicate down-regulation of below ground oxygen consumption. AoB PLANTS 8: 1-15. https://doi.org/10.1093/aobpla/plw025 Granéli, W., S.E.B. Weisner, and M.D. Sytsma. 1992. Rhizome dynamics and resource storage in Phragmites australis. Wetlands Ecology and Management 1: 239— 257. https://doi.org/10.1007/BF00244929 Greenberg, D.A., and D.M. Green. 2013. Effects of an in- vasive plant on population dynamics in toads. Conser- vation Biology 27: 1049-1057. https://doi.org/10.1111/ cobi.12078 Hanslin, H.M., T. Mzehlum, and A. Szebe. 2017. The re- sponse of Phragmites to fluctuating subsurface water levels in constructed stormwater management systems. Ecological Engineering 106: 385-391. https://do1.org/ 10.1016/j.ecoleng.2017.06.019 Hirtreiter, J.N., and D.L. Potts. 2012. Canopy structure, photosynthetic capacity and nitrogen distribution in ad- jacent mixed and monospecific stands of Phragmites australis and Typha latifolia. Plant Ecology 213: 821— 829. https://do1.org/10.1007/s11258-012-0044-2 Jacob, D.L., and M.L. Otte. 2003. Conflicting processes in the wetland plant rhizosphere: metal retention or mo- bilization? Water, Air and Soil Pollution: Focus 3: 91— 104. https://doi.org/10.1023/a:1022138919019 Jobbagy, E.G., and R.B. Jackson. 2004. The uplift of soil nutrients by plants: biogeochemical consequences across scales. Ecology 85: 2380-2389. https://do1.org/ 10.1890/03-0245 Jung, J.A., D. Rokitnicki-Wojcik, and J.D. Midwood. 2017. Characterizing past and modelling future spread of Phragmites australis ssp. australis at Long Point Peninsula, Ontario, Canada. Wetlands 37: 961-973. https://www.doi.org/10.1007/s13157-017-0931-3 Keller, B.E.M. 2000. Plant diversity in Lythrum, Phrag- mites, and Typha marshes, Massachusetts, U.S.A. Wet- lands Ecology and Management 8: 391—401. https://doi. org/10.1023/a:1026505817409 THE CANADIAN FIELD-NATURALIST Vol. 133 Mal, T.K., and L. Narine. 2004. The biology of Canadian weeds. 129. Phragmites australis (Cav.) Trin. ex Steud. Canadian Journal of Plant Science 84: 365-396. https:// doi.org/10.4141/PO1-172 Minchinton, T.E., J.C. Simpson, and M.D. Bertness. 2006. Mechanisms of exclusion of native coastal marsh plants by an invasive grass. Journal of Ecology 94: 342— 354. https://doi.org/10.1111/).1365-2745.2006.01099.x Ministry of Natural Resources and Forestry. 2017. Invasive Phragmites control at Long Point Region and Rondeau Provincial Park implementation plan. Minis- try of Natural Resources and Forestry, Aylmer District, Ontario Parks, Southwest Zone. Moore, G.E., D.M. Burdick, C.R. Peter, and D.R. Keirstead. 2012. Belowground biomass of Phragmites australis in coastal marshes. Northeastern Naturalist 19: 611-626. https://do1.org/10.1656/045.019.0406 Ouellet-Plamondon, C.M., J. Brisson, and Y. Comeau. 2004. Effect of macrophyte species on subsurface flow wetland performance in cold climate. Pages 8-15 in Proceedings of the 2004 Self-Sustaining Solutions for Streams, Wetlands, and Watersheds Conference, St. Paul, Minnesota, September 2004. American Society of Agricultural Engineers, St. Joseph, Minnesota, USA. https://doi.org/10.13031/2013.17368 Packer, J.G., L.A. Meyerson, H. Skalova, P. PySek, and C. Kueffer. 2017. Biological Flora of the British Isles: Phragmites australis. Journal of Ecology 105: 1123— 1162. https://doi.org/10.1111/1365-2745.12797 Ravit, B., J.G. Ehenfeld, and M.M. Héggblom. 2006. Effects of vegetation on root-associated microbial com- munities: a comparison of disturbed versus undisturbed estuarine sediments. Soil Biology and Biochemistry 38: 2359-2371. https://doi.org/10.1016/j.soilbio.2006.02.012 Robichaud, C.D., and R.C. Rooney. 2017. Long-term ef- fects of a Phragmites australis invasion on birds in a Lake Erie coastal marsh. Journal of Great Lakes Re- search 43: 141-149. https://doi.org/10.1016/j.jglr.2017.03. 018 Rothman, E., and V. Bouchard. 2007. Regulation of car- bon processes by macrophyte species in a Great Lakes coastal wetland. Wetlands 27: 1134-1143. https://doi. org/10.1672/0277-5212(2007)27[1134:rocpbm]2.0.co;2 Rudrappa, T., J. Bonsall, J.L. Gallagher, D.M. Seliskar, and H.P. Bais. 2007. Root-secreted allelochemicals in the noxious weed Phragmites australis deploys a reac- tive oxygen species response and microtubule assembly disruption to execute rhizotoxicity. Journal of Chemical Ecology 33: 1898-1918. https://doi.org/10.1007/s10886- 007-9353-7 Saltonstall, K. 2002. Cryptic invasion by a non-native gen- otype of the common reed, Phragmites australis, into North America. Proceedings of the National Academy of Sciences 99: 2445-2449. https://doi.org/10.1073/pnas. 032477999 Tulbure, M.G., and C.A. Johnston. 2010. Environmental conditions promoting non-native Phragmites australis expansion in Great Lakes coastal wetlands. Wetlands 30: 577-587. https://doi.org/10.1007/s13157-010-0054-6 van Wijk, M.T., M. Williams, L. Gough, S.E. Hobbie, 2019 and G.R. Shaver. 2003. Luxury consumption of soil nutrients: a possible competitive strategy in above- ground and below-ground biomass allocation and root morphology for slow-growing arctic vegetation? Jour- nal of Ecology 91: 664-676. https://do1.org/10.1046/ j.1365-2745.2003.00788.x Wilcox, K.L., S.A. Petrie, L.A. Maynard, and S.W. Meyer. 2003. Historical distribution and abundance of Phragmites australis at Long Point, Lake Erie, Ontario. Journal of Great Lakes Research 29: 664—680. https:// doi.org/10.1016/S0380-1330(03)70469-9 Windham, L. 2001. Comparison of biomass production and SUPPLEMENTARY MATERIAL: LEI ET AL.. BELOW GROUND CHANGES OF EUROPEAN REED 37] decomposition between Phragmites australis (common reed) and Spartina patens (salt hay grass) in brackish tidal marshes of New Jersey, USA. Wetlands 21: 179-188. https://doi.org/10.1672/0277-5212(2001)021[0179:cob pad]2.0.co;2 Windham, L., and J.G. Ehrenfeld. 2003. Net impact of a plant invasion on nitrogen-cycling processes within a brackish tidal marsh. Ecological Application 13: 883— 896. https://doi.org/10.1890/02-5005 Received 16 May 2019 Accepted 14 February 2020 FicurE S1. Total below ground biomass with water depth for European Reed (Phragmites australis) invaded sites (solid line and black triangles; nm = 29) and uninvaded sites (dashed line and white circles; n = 29). TABLE S1. Results table for GLM predicting total below ground biomass with and without interaction term. TABLE S82. Results table for GLM predicting rooting depth with and without interaction term. The Canadian Field-Naturalist Flathead Catfish (Pylodictis olivaris) reproduction in Canada CoLin ILLES!*, JULIA E. Coto’, NICHOLAS E. MANDRAK’, and Davip M. MArson! ‘Asian Carp Program, Fisheries and Oceans Canada, 867 Lakeshore Road, Burlington, Ontario L7S 1A1 Canada "Department of Biological Sciences, University of Toronto Scarborough, 1265 Military Trail, Toronto, Ontario MIC 1A4 Canada “Corresponding author: colin.illes@dfo-mpo.gc.ca Illes, C., JE. Colm, N.E. Mandrak, and D.M. Marson. 2019. Flathead Catfish (Py/odictis olivaris) reproduction in Canada. Canadian Field-Naturalist 133(4): 372-380. https://do1.org/10.22621/cfn.v13314.2323 Abstract Eleven Flathead Catfish (Py/odictis olivaris), representing at least five age classes, were collected between 2016 and 2018 in the lower Thames River, Ontario, Canada. The capture of two juveniles (total lengths 78 mm and 82 mm), the first records of juveniles in Canada, is a strong indication that reproduction has occurred. Previous records were thought to be individu- als that dispersed from known populations in American waters of Lake Erie. Flathead Catfish is currently designated as data deficient by the Committee on the Status of Endangered Wildlife in Canada. These new findings may provide sufficient data to reconsider the conservation status of this species. Key words: Flathead Catfish; Pylodictis olivaris; reproduction; Great Lakes; Lake St. Clair; Thames River; juvenile; young- of-year Introduction Flathead Catfish (Pylodictis olivaris) is found throughout the Mississippi River basin and lower Laurentian Great Lakes (Page and Burr 2011); how- ever, It is uncertain whether the species is native to the Great Lakes basin (Fuller and Whelan 2018). It is a benthic fish species preferring turbid (Lee and Terrell 1987; Hesse 1994), warm water (Becker 1983) in low-gradient, moderate to large rivers (Lee and Terrell 1987), and is commonly associated with woody debris, undercut banks, and substrate depres- sions throughout its range (Becker 1983; Hesse 1994; Grussing et al. 1999; Jackson 1999; Daugherty and Sutton 2005a). Flathead Catfish reach sexual ma- turity between three and six years of age when fish are 375-539 mm in total length (TL; Minckley and Deacon 1959; Perry and Carver 1977). Reproduction occurs in June and July when water temperatures reach at least 22.2°C (Becker 1983). Flathead Catfish use depressions and natural cavities to construct nests (Cooper 1983; Cross 1967) and females lay up to 31 579 eggs (Becker 1983). A detailed description of the life history of Flathead Catfish was reported by Goodchild (1993). Flathead Catfish is taxonomically and morpholog- ically different from all other catfish species in the Great Lakes basin. Differences include its protruding lower jaw, ventrally compressed head, large adipose fin, and backward extensions of the premaxillary tooth patches (although Stonecat [Noturus flavus] shares the latter characteristic). Flathead Catfish has a varying amount of mottled pigmentation on the body, and the upper lobe of the caudal fin has a pale tip (Figure 1), although these traits can be absent or less obvious in larger fish. Flathead Catfish has a slightly forked caudal fin in contrast to the deeply forked cau- dal fin of Channel Catfish (/ctalurus punctatus). Flathead Catfish has a short anal fin with a ray count of 13-18 (Trautman 1981), which differen- tiates it from Channel Catfish (25-28), Yellow Bullhead (Ameiurus natalis, 24—27), Brown Bullhead (Ameiurus nebulosus, 20-23) and, in some cases, Black Bullhead (Ameiurus melas, 17—21; Scott and Crossman 1998); all anal ray counts include rudi- mentary rays. Furthermore, Flathead Catfish has ser- rations on both edges of its pectoral spines whereas Channel Catfish and Brown and Black Bullheads have serrations only on the posterior edge. Madtoms (Noturus spp.) could be confused with juvenile Flathead Catfish, but are distinguished by a con- nected adipose fin and caudal fin, which are separate in Flathead Catfish. In the Great Lakes, Flathead Catfish has been re- corded in the Lake Erie, Lake St. Clair, Lake Huron, Lake Michigan, and Lake Superior basins. Since 1890, when Flathead Catfish was first recorded in Lake Erie, it has been documented in seven tribu- 372 ©The Ottawa Field-Naturalists’ Club 2019 ILLES ET AL.: FLATHEAD CATFISH REPRODUCTION IN CANADA a3 FicurE 1. Juvenile Flathead Catfish (Pylodictis olivaris), 78 mm total length, captured on 31 August 2016 in the lower Thames River, Ontario, Canada. Photo: Colin Illes. taries and is believed to have spread to the Lake St. Clair (Goodchild 1993; COSEWIC 2008) and Lake Huron basins where it has been recorded in six tribu- taries since the first records in 1989 and 1991, respec- tively (Fuller and Whelan 2018). In Lake Michigan, Flathead Catfish was first recorded in 1922 and has since been documented in 11 tributaries (Fuller and Whelan 2018). In addition, there is a single record of a Flathead Catfish in the Lake Superior basin, cap- tured in a pond in the Au Train River watershed and believed to be an unauthorized release (Fuller and Whelan 2018). A detailed description of the histori- cal and current distribution of Flathead Catfish in the American Great Lakes basin is provided by Fuller and Whelan (2018). Whether Flathead Catfish is native to the Great Lakes basin is not known because of poorly doc- umented historical records. Historical publica- tions variously mention (e.g., Trautman 1957) and do not mention (e.g., Evermann 1902) the presence of Flathead Catfish in the lower Great Lakes. Based on a review of the literature and capture data, Fuller and Whelan (2018) concluded that Flathead Catfish is not native to the Great Lakes basin, with the possi- ble exception of a small population documented since 1890 in the Huron River, Lake Erie (Trautman 1957). Conversely, Roth et al. (2012) indicated that Flathead Catfish is native to the Erie and Michigan basins. The origin of several other fishes in the Great Lakes basin is also unclear. Much like Flathead Catfish, Gizzard Shad (Dorosoma cepedianum), and Bigmouth Buffalo Uctiobus cyprinellus) have uncer- tain origins in Lake Erie and may have spread from the Mississippi River basin into Lake Erie where pop- ulations have continually expanded because of warm- ing temperatures (Miller 1957; Goodchild 1993). These fishes are considered native to the Great Lakes basin (e.g., Lee et a/. 1980; Trautman 1981; Scott and Crossman 1998; Roth et al. 2012), despite a lack of historical records and vouchered specimens. This has been attributed to misidentification with other species (e.g., Alewife [A/osa pseudoharengus|, Smallmouth Buffalo [/ctiobus bubalus]), and confusion with early introductions of Bigmouth Buffalo (Miller 1957; Trautman 1981). Gizzard Shad and Bigmouth Buffalo were first recorded in Lake Erie in 1848 and 1878, re- spectively (Miller 1957; Trautman 1981). Roth ef al. (2012) identified only three species of questionable native status in the Great Lakes basin: Ghost Shiner (Notropis buchanani), questionably native to the Erie and Huron basins; and Brindled Madtom (Noturus miurus) and Orangethroat Darter (Etheostoma spec- tabile) to the Michigan basin. In Canada, Flathead Catfish has been collected only in the Great Lakes basin with records limited to the western basin of Lake Erie and Lake St. Clair (COSEWIC 2008). The first Canadian capture of a Flathead Catfish was in Lake Erie, in 1978; it was caught west of Point Pelee, 3.2 km north of the tip, in a commercial trap net (Crossman and Leach 1979). Subsequently, three additional single specimens were captured in the Point Pelee area in 1986, 2005, and 2011 (COSEWIC 2008; Ontario Ministry of Natural Resources and Forestry [OMNRF] unpubl. data). All three fish were recorded in commercial trap nets west of Point Pelee and south of Sturgeon Creek within 8 km of each other. In 1989, Flathead Catfish was first captured by commercial long line in Lake St. Clair, 3.2 km from the mouth of the Thames River (Royal Ontario Museum unpubl. data). In 2001 and 2003, two additional Flathead Catfish were captured in Lake St. Clair in the St. Luke’s Bay area, 10 km north of the Thames River mouth (Figure 2), both in a commer- cial trap net (OMNREF unpubl. data). Based on the four known single specimens captured in Canadian waters between 1978 and 2001, the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) could not determine whether Flathead Catfish was native to Canada or a vagrant and, thus, assessed it as data de- ficient (COSEWIC 2008). Because of its preference for hard-to-sample hab- itats (e.g., beneath woody debris and structured sub- strate in deep water), low population abundance, and solitary behaviour, Flathead Catfish has been notori- 374 THE CANADIAN FIELD-NATURALIST Vol. 133 83°W 82°W Lake H a e Huron 43°N A3°N Lake St. Clair MICHIGAN ONTARIO 42°N 42°N ee Lake Erie 83°W Kilometres 20 82°W FiGurE 2. Capture locations of Flathead Catfish (Pylodictis olivaris) in Canadian waters of the Great Lakes basin, a. 2016-2018 and b. 1979-2018. Source: Unpublished data from Fisheries and Oceans Canada, the Ontario Ministry of Natural Resources and Forestry, and the Royal Ontario Museum, sourced under the Open Government Licence Ontario. ously difficult to assess in river systems (Stauffer e¢ al, 1996; Vokoun and Rabeni 1999). This has led to limited targetted sampling and knowledge about the Species, especially in the Great Lakes (Daughtery and Sutton 2005a). To our knowledge, there has been only one population estimate for Flathead Catfish in the Great Lakes basin, conducted in the lower St. Joseph River, Michigan, which estimated an abundance of 5453 individuals (Daughtery and Sutton 2005b). In this study, we report recent records of Flathead Catfish that indicate reproduction in the Canadian waters of the Great Lakes basin and discuss implica- tions of these records for future management. Methods The Thames River, a tributary of Lake St. Clair, is a large, turbid river with a high diversity of fish and mussel species, 25 of which are at risk (Cudmore ef al. 2004). The Thames River watershed has been im- pacted by agriculture and urban and rural develop- ment (Cudmore ef a/. 2004). In addition to supporting several imperilled species, the river is highly suita- ble for the reproduction of four species of invasive Asian carp (Cudmore et al. 2017) and, therefore, is sampled routinely by Fisheries and Oceans Canada’s Asian Carp Program (Colm et al. 2019a). This sampling occurred between May and No- vember, 2013-2018, using seven gear types to tar- get adult and juvenile Asian carp, while also collect- ing baseline fish community data (Marson ef al. 2014, 2016, 2018; Colm et a/. 2018, 2019a,b). Sampling ef- fort in the lower Thames River during this period is summarized in Table 1. Flathead Catfish were captured inthe lower Thames River using three gear types: boat electrofishing (n = 2019 ILLES ET AL.: FLATHEAD CATFISH REPRODUCTION IN CANADA 375 TABLE 1. Summary of sampling effort in the lower Thames River, 2013-2018, as part of the early detection surveillance efforts of Fisheries and Oceans Canada’s Asian Carp Program. Boat electrofishing Hoop net Mini-fyke net Year No. Effort, No. Effort, No. Effort, sites h sites h sites h 2013 4 0.4 0 0.0 0 0.0 2014 19 all 3 112.4 0 0.0 2015 33 5.6 7 Sil. 0 0.0 2016 25 46 0 0.0 6 130.4 DOV], 22 3.8 10 460.1 7 155.9 2018 28 6.0 9 397.4 16 355.1 Sources: Marson et al. 2014, 2016, 2018; Colm et al. 2018, 2019a,b. 4), hoop nets (n = 2), and trammel nets (n= 5). Before 2018, the boat electrofisher was dual-boom, 6.4 m in length, and fitted with a 7.5 Generator Powered Pul- sator (Smith-Root, Vancouver, Washington, USA). In 2018, the boat electrofisher used in sampling was 7.3 m in length, dual-boom, and fitted with an Infinity Box (Midwest Lake Electrofishing, Polo, Missouri, USA). Two sizes of hoop nets were used: 1.5 m in di- ameter, 6.1 m in length, with 2.5-cm square mesh; and 0.91 m in diameter, 4.57 m in length, with 2.5-cm square mesh. Trammel nets were 183 m in length, 4.3 m in height, with 10.1-cm bar mesh and 45.7-cm outer wall panels. Trammel nets were often used in combi- nation with boat electrofishing as this is an effective method for targetting Grass Carp (Ctenopharyngodon idella), a species of Asian carp, in the Great Lakes basin (D.M.M. pers. obs.); however, fishes captured with the two gear types were processed separately. All gear types and the scope of sampling (includ- ing other locations in the Great Lakes basin) are de- scribed in Colm et al. (2019a). Seine net Trap net Trammel net No. Effort, No. Effort, No. Effort, sites hauls sites h sites h 2 6 0 0.0 2 1.8 0 0 4 91.4 5) 1.8 0 0 12 244.4 13 9.1 0 0 7 153.6 8 5.4 0 0 10 214.4 9 23 2 5 13 293.8 14 12.4 Results During 2016-2018, 11 Flathead Catfish (Table 2) were captured in three locations in the lower Thames River, near the mouth of Jeannettes Creek, Kent County, Tilbury Township (42.329°N, 82.421°W) (Figure 2a). No Flathead Catfish were detected in 2013-2015, despite sampling in similar areas to 2016— 2018. In August 2016, we recorded three Flathead Catfish, with TL 78-566 mm, at two locations. All three fish were captured using a boat electrofisher near shore in close proximity to woody debris on a clay—silt substrate (Table 2). The first location had an undercut bank with a single cluster of woody debris; the second had abundant large woody debris, includ- ing trees, logs, and branches, and a water depth of ~1 m at the bank. In June 2017, a Flathead Catfish measuring 365 mm TL was captured with a hoop net in a new location with abundant submerged logs and branches. This individual was caught farthest downstream, 1.3 km from the Thames River mouth. In September 2017, two Flathead Catfish measur- ing 833 mm and 815 mm TL were captured using a TABLE 2. Capture data for Flathead Catfish (Pylodictis olivaris). Capture locations shown in Figure 2a. tone Temp., Turbidity, Max. site Coarse woody Pecauion: OM Date length, oC NTU depth, m Substrate debris, Y/N Gear type of catalogue mm capture no. 30 Aug. 2016 566 26.29. 28.71 4.2 Clay-silt nv Boat electrofishing 1 101500 30 Aug. 2016 82 2717 27.44 2.0 Clay-silt B Boat electrofishing 2 105705 31 Aug. 2016 78 26207 ~ 2507. 2.1 Clay-—silt bi Boat electrofishing 2 101375 20 June 2017 365 24.61 16.04 2.8 Clay-silt ay: Hoop net 3 109946 13 Sept. 2017 833 20.34 23.70 4] Clay—silt Y Trammel net 1 NA 14 Sept. 2017 815 21.44 11.30 2.0 Clay-silt ¥ Boat electrofishing 2 NA 26 June 2018 820 23:71 23.11 5.0 Clay-—silt ¥ Hoop net 2 NA 27 Sept. 2018 697 20.31 STL 40 Silt—clay Ni Trammel net with 2 NA boat electrofishing 27 Sept. 2018 765 20.22 20.86 3.8 Silt—clay Y; Trammel net with 1 NA boat electrofishing 27 Sept. 2018 743 20.22 20.86 3.8 Silt—clay 2 Trammel net with 1 NA boat electrofishing 2 Oct. 2018 388 13297 2onls 4.0 Clay-silt ¥ Trammel net with 3 NA Source: Fisheries and Oceans Canada unpubl. data. boat electrofishing Note: NTU = nephelometric turbidity units, ROM = Royal Ontario Museum. 376 trammel net and boat electrofisher (Table 2). The 815- mm individual was the farthest upstream record, 6.3 km from the Thames River mouth. In 2018, five ad- ditional Flathead Catfish were collected, measuring 388-820 mm TL. The 820-mm individual was cap- tured using a hoop net at the deepest recorded capture of 5 m. The other four Flathead Catfish were captured in trammel nets used in combination with boat elec- trofishing (Table 2). Four specimens of Flathead Catfish captured in 2016 and early 2017 were preserved in 10% buffered formalin, stored in 70% ethanol, and catalogued at the Royal Ontario Museum (Table 2). Digital voucher photos were taken of the remaining seven fish before they were released. Using length-at-age data from the literature (Mayhew 1969; Young and Marsh 1990; Kwak et al. 2006; Sakaris et al. 2006), we estimate that the 11 Flathead Catfish were from five different age classes (Figure 3). Discussion We report the first indication of Flathead Catfish reproduction in the Canadian waters of the Great Lakes basin. Over six consecutive years of sampling, 11 individuals were detected in the lower Thames River, Ontario. To our knowledge, no length-at-age data are available for Flathead Catfish in the Great Lakes basin. Flathead Catfish collected on 30 August 2016 and 31 August 2016 with TLs of 78 mm and 82 mm, respectively, are assumed to be young-of-year. In addition to being the first recorded juveniles in Canada, these are the first records of Flathead Catfish from a river system in Canada; previous detections were in large bays. In the first year of growth, Flathead Catfish has been documented to reach 100 mm TL in Ohio (Trautman 1981) and 145 mm TL in Arizona (Young and Marsh 1990). Daugherty and Sutton (2005b) re- corded Flathead Catfish measuring 87 mm and 93 6 5 Frequency wf N a KP SS KF MK Total length (mm) Figure 3. Length—frequency distribution of Flathead Catfish (Pylodictis olivaris) captured in the lower Thames River, 2016—2018, by Fisheries and Oceans Canada’s Asian Carp Program. THE CANADIAN FIELD-NATURALIST Vol. 133 mm TL, which were assumed to be young-of-year, while sampling the lower St. Joseph River, Michigan, June through September 2002-2003. Historical records of Flathead Catfish captured in Canada before 2001 were speculated to be individ- uals that dispersed from a known population in the Huron River, Lake Erie, and gained access to Lake St. Clair through the Detroit River (Goodchild 1993; COSEWIC 2008). The juvenile Flathead Catfish cap- tured in our study would not likely be able to dis- perse from a Lake Erie tributary upstream through the strong-flowing Detroit River, providing further support for the likelihood that these individuals were the result of reproduction in the Thames River. Few studies have examined the movement of juvenile Flathead Catfish. Travnichek (2004) found a relation- ship between Flathead Catfish size and movement in the Missouri River: as size increased so did dis- tance travelled. In their study, Flathead Catfish that were 305-380 mm TL travelled an average of 4.6 km in up to two years after tagging (Travnichek 2004). Stocking is an unlikely alternative method of intro- duction of the individuals captured here, as there is no documented stocking of Flathead Catfish in Ontario and, in the United States, Flathead Catfish are most often stocked as adults (Guier ef a/. 1981; Jenkins and Burkhead 1994). Introduction through the live fish trade is also unlikely, as there is no record that live Flathead Catfish have been imported into Canada (Mandrak et al. 2014) and surveys of six live fish mar- kets and 20 pet stores in the Great Lakes region did not report Flathead Catfish (Rixon et a/. 2005). Flathead Catfish may be more abundant in the Canadian waters of the Great Lakes basin than cur- rently known, and its range is likely not fully docu- mented because of its cryptic behaviour and difficulty to sample (Goodchild 1993; Fuller and Whelan 2018). Continued Asian carp surveillance will further docu- ment the Flathead Catfish population in the Thames River. This work is also being conducted in 35 other locations in the Great Lakes basin and may provide information on new locations of Flathead Catfish populations. The potential impacts of Flathead Catfish in the Great Lakes basin are unknown and should be fur- ther investigated to determine how this species should be managed. Pine et al. (2005) found that Flathead Catfish is primarily piscivorous, feeding on the most abundant fishes in rivers, which could lead to a change in local food-web structures. The pres- ence and increased abundance of Flathead Catfish has been associated with decreases in the abundance of sunfishes (Lepomis spp.; Davis 1985; Thomas 1993; Bart et al. 1994; Ashley and Rachels 1998; Bonvechio et al. 2009), black basses (Micropterus spp.; Thomas 2019 1993; Bonvechio et al. 2009), redhorses (Moxostoma spp.; Bart et al. 1994), Common Carp (Cyprinus car- pio, Davis 1985; Bart et al. 1994), and bullheads (Ameiurus spp.,; Davis 1985). Flathead Catfish may have some ecological benefits in the Great Lakes ba- sin, as a predator of the invasive Common Carp and as a known host for Mapleleaf mussel (Quadrula quadrula, Howard and Anson 1922), which has been listed as special concern under the Canadian Species at Risk Act (SARA Registry 2019). The Thames River is thought to have the largest population of Mapleleaf in southwestern Ontario, and recent records of Flathead Catfish overlap with records of Mapleleaf from the lower Thames River in 2005 (COSEWIC 2016). Flathead Catfish has seasonally varying home ranges and movement patterns (Daugherty and Sutton 2005a), which are important characteristics of hosts that facilitate the genetic mixture of freshwater mussel populations in rivers (Elderkin et al. 2007). The distribution of freshwater fishes is often re- stricted by temperature. Mandrak (1989) determined that Flathead Catfish had low potential for future ex- pansion into the Great Lakes basin because of ther- mal restrictions. With climate change, the water tem- perature of the Great Lakes is expected to increase 2-3°C in southern Ontario and 3—4°C in north- ern Ontario by 2065 (Gula and Peltier 2012). Such an increase will benefit warm-water species, such as Flathead Catfish, by increasing recruitment suc- cess (Casselman 2002; Chu et al. 2005; Hansen et al. 2017). This increase in recruitment has the potential to expand the range of Flathead Catfish and lead to a greater abundance of the species in the Great Lakes basin (Casselman 2002). Our research suggests that a better understand- ing of the potential ecological impacts and improved distribution modelling of Flathead Catfish in Canada is required. With climate change, many species are likely to undergo range expansions, bringing new threats to already imperilled native species. With lim- ited resources, managers must balance this with the threats of new (or existing) invasive species that have a greater potential for damage (Rahel and Olden 2008; Rolls et al. 2017). In Canada, there is a need for clear management objectives for these species undergoing natural “invasions”, that include consistent classifica- tion and terminology and a framework to prioritize them (Chu et al. 2005; Rahel and Olden 2008). The capture of Flathead Catfish representing at least five age classes, including young-of-year fish, is a strong indication that reproduction has oc- curred in the lower Thames River. With the recent captures reported here, there may now be suffi- cient data for Flathead Catfish to be re-assessed by COSEWIC. Additional research targetting Flathead ILLES ET AL.: FLATHEAD CATFISH REPRODUCTION IN CANADA Sia Catfish is recommended to (i) better understand the distribution of this species in Canada, (11) evaluate the most effective gear for detection, (i11) estimate abun- dance, and (iv) understand the movement and habi- tat-use patterns in the Canadian waters of the Great Lakes basin. Author Contributions Writing — Original Draft: C.I. and J.E.C.; Writing — Review & Editing: C.I., J.E.C., N.E.M., and D.M.M.; Methodology: C.L., J.E.C., and D.M.M.; Visualization: CI. and J.E.C. Acknowledgements We thank the Fisheries and Oceans Canada Asian Carp Program for providing funding and sampling data and the Asian Carp Program field surveillance staff for collecting the data. The Royal Ontario Mu- seum shared historical Flathead Catfish capture re- cords from Ontario. Andy Cook, Ontario Ministry of Natural Resources and Forestry, provided additional information on an unpublished capture record. Sara Thomas, Michigan Department of Natural Resources, provided knowledge and insights into records from American waters of the Great Lakes basin. We also thank an anonymous reviewer, Mark Poesch, Fran¢ois Chapleau, and Dwayne Lepitzki for provid- ing edits and suggestions, which improved the paper. Literature Cited Ashley, K.W., and R.T. Rachels. 1998. Changes in red- breast sunfish population characteristics in the Black and Lumber rivers, North Carolina. Pages 29-38 in Proceedings of the Fifty-second Annual Conference Southeastern Association of Fish and Wildlife Agencies. Edited by A.G. Eversole, K.C. Wong, and R.W. Luebke. Orlando, Florida, USA. Bart, H.L., M.S. Taylor, J.T. Harbaugh, J.W. Evans, S.L. Schleiger, and W. Clark. 1994. New distribution records of Gulf Slope drainage fishes in the Ocmulgee River system, Georgia. Pages 4—9 in Southeastern Fishes Council Proceedings. Edited by M.M. Stevenson and G.R. Sedberry. Southeastern Fishes Council Inc., Charleston, South Carolina, USA. Becker, G.C. 1983. Fishes of Wisconsin. University of Wisconsin Press. Madison, Wisconsin, USA. Bonvechio, T.F., D. Harrison, and B. Deener. 2009. Population changes of sportfish following flathead cat- fish introduction in the Satilla River, Georgia. Pages 133-139 in Proceedings of the Sixty-third Annual Con- ference Southeastern Association of Fish and Wildlife Agencies. Edited by A.G. Eversole, K.C. Wong, and B. Davin. Atlanta, Georgia, USA. Casselman, J.M. 2002. Effects of temperature, global ex- tremes, and climate change on year-class production of warmwater, coolwater, and coldwater fishes in the Great Lakes Basin. Pages 39-60 in Fisheries in a Changing 378 Climate. Symposium 32. Edited by N.A. McGinn. American Fisheries Society, Bethesda, Maryland, USA. Chu, C., N.E. Mandrak, and C.K. Minns. 2005. Potential impacts of climate change on the distributions of sev- eral common and rare freshwater fishes in Canada. Diversity and Distributions 11: 299-310. https://do1.org/ 10.1111/,.1366-9516.2005.00153.x Colm, J., D. Marson, and B. Cudmore. 2018. Results of Fisheries and Oceans Canada’s 2016 Asian carp early de- tection field surveillance program. Canadian manuscript report 3147. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Colm, J., D. Marson, and B. Cudmore. 2019a. Results of Fisheries and Oceans Canada’s 2017 Asian carp early detection field surveillance program. Canadian man- uscript report 3168. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Colm, J., D. Marson, and B. Cudmore. 2019b. Results of Fisheries and Oceans Canada’s 2018 Asian carp early detection field surveillance program. Canadian manu- script report 3168-1. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Cooper, E.L. 1983. Fishes of Pennsylvania and the North- eastern United States. Pennsylvania State University Press, University Park, Pennsylvania, USA. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2008. Update COSEWIC Sta- tus Report on Flathead Catfish Pylodictis olivaris. COSEWIC, Ottawa, Ontario, Canada. COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2016. COSEWIC assessment and status report on the Mapleleaf Quadrula qua- drula (Great Lakes—Upper St. Lawrence population, Saskatchewan-Nelson Rivers population) in Canada. COSEWIC, Ottawa, Ontario, Canada. Cross, F.B. 1967. Handbook of Fishes of Kansas. Museum of Natural History, University of Kansas, Lawrence, Kanas, USA. Crossman, E.J., and J.H. Leach. 1979. First Canadian re- cord of Flathead Catfish. Canadian Field-Naturalist 93: 179-180. Accessed 17 March 2020. https://www.bio diversitylibrary.org/page/28063496. Cudmore, B., L.A. Jones, N.E. Mandrak, J.M. Dett- mers, D.C. Chapman, C.S. Kolar, and G. Conover. 2017. Ecological risk assessment of Grass Carp (Cteno- pharyngodon idella) for the Great Lakes basin. Canadian Science Advisory Secretariat Science Research Docu- ment 2016/118. Fisheries and Oceans Canada, Ottawa, Ontario, Canada. Cudmore, B., C.A. Mackinnon, and S.E. Madzia. 2004. Aquatic species at risk in the Thames River watershed, Ontario. Fisheries and Oceans Canada, Burlington, Ontario, Canada. Accessed 15 May 2019. https://waves- vagues.dfo-mpo.gc.ca/Library/316802.pdf. Daugherty, D.J., and T.M. Sutton. 2005a. Seasonal move- ment patterns, habitat use, and home range of flathead catfish in the Lower St. Joseph River, Michigan. North American Journal of Fisheries Management 25: 256— 269. https://doi.org/10.1577/M03-252.2 THE CANADIAN FIELD-NATURALIST Vol. 133 Daugherty, D.J., and T.M. Sutton. 2005b. Population abundance and stock characteristics of flathead cat- fish in the lower St. Joseph River, Michigan. North American Journal of Fisheries Management 25: 1191-— 1201. https://doi.org/10.1577/M03-251.1 Davis, R.A. 1985. Evaluation of flathead catfish as a pred- ator in a Minnesota lake. Investigational report 384. Division of Fish and Wildlife, Minnesota Department of Natural Resources, Saint Paul, Minnesota, USA. Accessed 5 March 2019. http://www.nativefishlab.net/ library/textpdf/13980.pdf. Elderkin, C.L., A.D. Christian, C.C. Vaughn, J.L. Metcalfe-Smith, and D.J. Berg. 2007. Population ge- netics of the freshwater mussel, Amblema plicata (Say 1817) (Bivalvia: Unionidae): evidence of high dispersal and post-glacial colonization. Conservation Genetics 8: 355-372. https://doi.org/10.1007/s10592-006-9175-0 Evermann, B.W. 1902. List of fishes known to occur in the Great Lakes or their connecting water. Bulletin of the United States Fish Commission 21: 95-96. Fuller, P.L., and G.E. Whelan. 2018. The flathead cat- fish invasion of the Great Lakes. Journal of Great Lakes Research 44: 1081-1092. https://doi.org/10.1016/j. jglr.2018.07.001 Goodchild, A.C. 1993. Status of the Flathead Catfish, Plyodictis olivaris, in Canada. Canadian Field-Natu- ralist 107: 410—416. Accessed 17 March 2020. https:// www. biodiversitylibrary.org/page/34810552. Grussing, M.D., D.R. DeVries, and R.A. Wright. 1999. Stock characteristics and habitat use of catfishes 1n reg- ulated sections of four Alabama rivers. Pages 15-34 in Proceedings of the Fifty-third Annual Conference Southeastern Association of Fish and Wildlife Agencies. Edited by A.G. Eversole, K.C. Wong, and P. Mazik. Baton Rouge, Louisiana, USA. Guier, C.R., L.E. Nichols, and R.T. Rachels. 1981. Bio- logical investigation of flathead catfish in the Cape Fear River. Pages 607—621 in Proceedings of the Thirty-fifth Annual Conference Southeastern Association of Fish and Wildlife Agencies. Edited by J. Sweeney and L. Nielsen. Tulsa, Oklahoma, USA. Gula, J., and W.R. Peltier. 2012. Dynamic downscaling over the Great Lakes basin of North America using the WRF regional climate model: the impact of the Great Lakes system on regional greenhouse warming. Journal of Climate 25: 7723-7742. https://doi.org/10.1175/JC LI-D-11-00388.1 Hansen, G.J.A., J.S. Read, J.F. Hansen, and L.A. Win- slow. 2017. Projected shifts in fish species dominance in Wisconsin lakes under climate change. Global Change Biology 23: 1463-1476. https://doi.org/10.1111/ gcb.13462 Hesse, L.W. 1994. The status of Nebraska fishes in the Missouri River, flathead catfish, Plydoictis oliva- ris, and blue catfish, /ctalurus furcatus (Ictaluridae). Transactions of the Nebraska Academy of Sciences 21: 89-98. Howard, A.D., and B.J. Anson. 1922. Phases in the para- sitism of the Unionidae. Journal of Parasitology 9: 68— 82. https://doi.org/10.2307/3271139 2019 Jackson, D.C. 1999. Flathead catfish: biology, fisheries, and management. Pages 23-35 in Catfish 2000: Proceedings of the International Ictalurid Symposium. Edited by E.R. Irwin, W.A. Hubert, C.F. Rabeni, H.L. Schramm, Jr., and T. Coon. American Fisheries Society, Bethesda, Maryland, USA. Jenkins, R.E., and N.M. Burkhead. 1994. Freshwater Fishes of Virgina. American Fisheries Society. Be- thesda, Maryland, USA. Kwak, T.J., D.S. Waters, and W.E. Pine, IIT. 2006. Age, growth, and mortality of introduced flathead catfish in Atlantic Rivers and a review of other populations. North American Journal of Fisheries Management 26: 73-87. https://doi.org/10.1577/M04-144.1 Lee, D.S., C.R. Gilbert, C.H. Hocutt, R.E. Jenkins, D.E. McAllister, and J.R. Stauffer, Jr. 1980. Atlas of North American Freshwater Fishes. North Carolina State Mu- seum of Natural History, Raleigh, North Carolina, USA. Lee, L.A., and J.W. Terrell. 1987. Habitat suitability in- dex models: flathead catfish. Biological report 82(10, 152). United States Fish and Wildlife Service, National Ecology Research Center, Fort Collins, Colorado, USA. Mandrak, N.E. 1989. Potential invasion of the Great Lakes by fish species associated with climatic warming. Jour- nal of Great Lakes Research 15: 306-316. https://doi. org/10.1016/S0380-1330(89)71484-2 Mandrak, N.E., C. Gantz, L.A. Jones, D. Marson, and B. Cudmore. 2014. Evaluation of five freshwater fish screening-level risk assessment protocols and applica- tion to non-indigenous organisms in trade in Canada. Canadian Science Advisory Secretariat Research Docu- ment 2013/122. Fisheries and Oceans Canada, Burling- ton, Ontario, Canada. Marson, D., J. Colm, and B. Cudmore. 2018. Results of Fisheries and Oceans Canada’s 2015 Asian carp early de- tection field surveillance program. Canadian manuscript report 3146. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Marson, D., E. Gertzen, and B. Cudmore. 2014. Results of the Burlington 2013 Asian carp early detection field monitoring program. Canadian manuscript report 3054. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Marson, D., E. Gertzen, and B. Cudmore. 2016. Results of Fisheries and Oceans Canada’s 2014 Asian carp early de- tection field surveillance program. Canadian manuscript report 3103. Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, Burlington, Ontario, Canada. Mayhew, J.K. 1969. Age and growth of flathead catfish in the Des Moines River, Iowa. Transactions of the American Fisheries Society 98: 188—121. https://doi.org/ 10.1577/1548-8659(1969)98[118:A AGOFC]2.0.CO;2 Miller, R.R. 1957. Origin and dispersal of the Alewife, A/soa pseudoharengus, and the Gizzard Shad, Dorosoma ce- pedianum, in the Great Lakes. Transactions of the American Fisheries Society 86: 97-111. https://doi.org/ 10.1577/1548-8659(1956)86[97:0A DOTA]2.0.CO;2 Minckley, W.L., and J.E. Deacon. 1959. Biology of the Flathead Catfish in Kansas. Transactions of the Ameri- can Fisheries Society 8: 344-355. ILLES ET AL.: FLATHEAD CATFISH REPRODUCTION IN CANADA 379 Page, L.M., and B.M. Burr. 2011. Peterson Field Guide to Freshwater Fishes. Second Edition. Houghton Mifflin, Boston, Massachusetts, USA. Perry, W.G., and D.C. Carver. 1977. Length at maturity, total-collarbone length, and dressout for Flathead Cat- fish and length at maturity of Blue Catfish, southwest Louisiana. Pages 529-537 in Proceedings of the Thirty- first Annual Conference Southeastern Association of Fish and Wildlife Agencies. Edited by R.W. Dimmick and J.A. Grover. Knoxville, Tennessee, USA. Pine, III, W.E., T.J. Kwak, D.S. Waters, and J.A. Rice. 2005. Diet selectivity of introduced Flathead Catfish in coastal rivers. Transactions of the American Fisheries Society 134: 901-909. https://do1.org/10.1577/T04-166.1 Rahel, F.J., and J.D. Olden. 2008. Assessing the ef- fects of climate change on aquatic invasive species. Conservation Biology 22: 521-533. https://doi.org/10. 1111/4).1523-1739.2008.00950.x Rixon, C.A.M., LC. Duggan, N.M.N. Bergeron, A. Ricciardi, and H.J. Macisaac. 2005. Invasion risks posed by the aquarium trade and live fish markets on the Laurentian Great Lakes. Biodiversity and Conser- vation 14: 1365-1381. https://doi.org/10.1007/s10531-0 04-9663-9 Rolls, R.J., B. Hayden, and K.K. Kahilainen. 2017. Con- ceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish. Ecology and Evolution 7: 4109-4128. https://doi.org/10. 1002/ece3.2982 Roth, B.M., N.E. Mandrak, T.R. Hrabik, G.G. Sass, and J. Peters. 2012. Fishes and decapod crustaceans of the Great Lakes basin. Pages 105—135 in Great Lakes Policy and Management. Second Edition. Edited by WW. Tay- lor and A. Lynch. Michigan State University Press, East Lansing, Michigan, USA. Sakaris, P.C., E.R. Irwin, J.C. Jolley, and D. Harrison. 2006. Comparison of native and introduced flathead cat- fish populations in Alabama and Georgia: growth, mor- tality, and management. North American Journal of Fisheries Management 26: 867—874. https://doi.org/10. 1577/M05-135.1 SARA (Species at Risk Act) Registry. 2019. Species profile, Mapleleaf. Government of Canada. Accessed 14 Febru- ary 2020. https://species-registry.canada.ca/index-en. html#/species?key words=Mapleleaf. Scott, W.B., and E.J. Crossman. 1998. Freshwater Fishes of Canada. Revised Edition. Galt House Publishing, Oakville, Ontario, Canada. Stauffer, K.W., R.C. Binder, B.C. Chapman, and B.D. Koenen. 1996. Population characteristics and sampling methods of flathead catfish Plyodictis olivaris in the Minnesota River: final report. Minnesota Department of Natural Resources, Division of Fish and Wildlife, Section of Fisheries, Saint Paul, Minnesota, USA. Thomas, M.E. 1993. Monitoring the effects of intro- duced flathead catfish on sport fish populations in the Altamaha River, Georgia. Pages 531-538 in Proceedings of the Forty-seventh Annual Conference Southeastern Association of Fish and Wildlife Agencies. Edited by A.G. Eversole, K.C. Overacre, and M. Konikoff. Atlan- ta, Georgia, USA. 380 Trautman, M.B. 1957. The Fishes of Ohio with Illustrated Keys. Ohio State University Press, Columbus, Ohio, USA. Trautman, M.B. 1981. The Fishes of Ohio with Illustrated Keys. Revised Edition. Ohio State University Press, Co- lumbus, Ohio, USA. Travnichek, V.H. 2004. Movement of flathead catfish in the Missouri River: examining opportunities for man- aging river segments for different fishery goals. Fish- eries Management and Ecology 11: 89-96. https://doi. org/10.1046/}.1365-2400.2003.00377.x Vokoun, J.C., and C.F. Rabeni. 1999. Catfish sampling THE CANADIAN FIELD-NATURALIST Vol. 133 in rivers and streams: a review of strategies, gears, and methods. Pages 271-286 in Catfish 2000: Proceedings of the International Ictalurid Symposium. Edited by E.R. Irwin, W.A. Hubert, C.F. Rabeni, H.L. Schramm, Jr., and T. Coon. American Fisheries Society, Bethesda, Maryland, USA. Young, K.L., and P.C. Marsh. 1990. Age and growth of flathead catfish in four southwestern rivers. California Fish and Game 76: 224—233. Received 26 July 2019 Accepted 14 February 2020 The Canadian Field-Naturalist Book Reviews Book Review Editor’s Note: The Canadian Field-Naturalist is a peer-reviewed scientific journal publishing papers on ecology, behaviour, taxonomy, conservation, and other topics relevant to Canadian natural history. In line with this mandate, we review books with a Canadian connection, including those on any species (na- tive or non-native) that inhabits Canada, as well as books covering topics of global relevance, including climate change, biodiversity, species extinction, habitat loss, evolution, and field research experiences. Currency Codes: CAD Canadian Dollars, USD United States Dollars, EUR Euros, AUD Australian Dollars, GBP British Pounds. ENTOMOLOGY Buzz, Sting, Bite: Why We Need Insects By Anne Sverdrup-Thygeson. 2019. Simon and Schuster. 235 pages, 35.00 CAD, Cloth. Buzz, Sting, Bite is another entry into the growing list of accessible popular science books written as passion projects by an academic re- searcher. A professor of conservation biology at the Norwegian University of Environmental and Life Sci- oe ences and a scientific ad- nversow visor to the Norwegian In- stitute for Nature Research, Sverdrup-Thygeson’s spe- cialty is the ecological role of insects in trees and forests, but the book covers arthropods and their eco- logical roles more broadly. In addition to a multi- tude of interesting facts, it includes some discussion of broader conservation ecology, such as habitat con- nectivity, extinction debt, and endangered species. As promised by the tagline “why we need insects”, the work also delves into humanity’s ties to the in- sect world, from 13th century Chinese cricket fights to termites eating their way through stashes of bank notes. Organized into nine main chapters, the scope is broad and about what you would expect from the out- set: anatomy, mating, agricultural food systems, the ecological role of detritovores, and insect-human interactions. While each chapter has a stated theme, they are further divided into multiple sections and subsections. Overall quite intuitive and well man- aged, this structure does pose narrative challenges and can become disjointed at times as topics begin BUZZ STING BITE i, Why We Need Insects to blur together. One key advantage of this bite- sized-piece approach is that like many books writ- ten for popular audiences, it makes for easy reading; this book may not pull you in for an all-night read- ing binge but it is well designed to be picked up at your leisure. Artfully translated by Lucy Moffatt from the orig- inal Norwegian 2018 publication, Buzz, Sting, Bite in- cludes some truly excellent explanations and turns of phrase. Although there are a few notable oversimpli- fications when discussing the natural history of insect groups and genera (e.g., bumble bees), the writing 1s at its best when it focusses on the truly weird and won- derful. Chapter 7, From Silk to Shellac: Industries of Insects, was by far my personal favourite, galloping across time periods and cultural traditions to bring together everything from oak gall wasps and histor- ical records, silk production, bulletproof vests, and the Aztec and Mayan traditions of breeding cochi- neal bugs. To my repertoire of offbeat insect-based cocktail conversation I can now add the link between shellac, phonograph records, and a 1942 restriction ordered by the United States government on the re- cord industry to reduce shellac consumption by at least 70%—for this I am forever grateful. The main text is complimented by black and white illustrations by artist Tuva Sverdrup-Thygeson, one at the beginning of each chapter matched to its over- all theme. These are welcome additions, as is the list of eight other author-recommended popular science insect books found under Further Reading follow- ing the Acknowledgments section. Although no in- text citations are provided, a bibliography of sorts is 381 382 found in the 20-page Sources section which is organ- ized by chapter and includes full citations of journal papers, reports, books, and popular science articles. The text ends with a detailed Index, so when you in- evitably want to refresh the details on a specific fact or anecdote it is at your fingertips. One of the author’s objectives in writing this book is to shine a spotlight on the creepy-crawly things of the world and shower them with the praise and appre- ciation that they deserve. In highlighting their value to human societies and their intrinsic ‘cool’ factor (even going so far as to use that rarest punctuation mark, the exclamation point, on several occasions), the author is largely successful. Although I doubt that those with a serious bug phobia will be drawn to this book, the range and variety of topics covered means that there is probably something here for every- THE CANADIAN FIELD-NATURALIST Vol. 133 one. Human-insect-avian interactions? Take Greater Honeyguide (/ndicator indicator) birds and _ their collaboration with the Yao people of Mozambique. Urban ecology and localized natural selection events? Have a side of mosquito speciation by station area of the London Underground. Want to hop on the in- sect eating bandwagon? You’ll find it espoused here, if only briefly. Reading Buzz, Sting, Bite, | was re- minded why I enjoy reading books on broad topics written by good writers—the more I learn, the more I want to know. This book provides more breadth than depth when it comes to bug love but is an excellent jumping off point for those who want to dive deeper, and a great toe dip for any who may otherwise hesi- tate to even approach the water’s edge. HEATHER A. CRAY Halifax, NS, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 BooK REVIEWS 383 Butterflies: Their Natural History and Diversity. Second Edition By Ronald Orenstein. Photography by Thomas Marent. 2020. Firefly Books. 24.95 USD / CAD, Paper. A quick internet search ap- pears to confirm the easy notion that butterflies must be the most popular of the insects. I think of them as the birds of the insect world, often colourful, active, and highly visible. And, as with birds, the internet is full of books, posters, calendars, etc. related to butterflies. One might wonder at the need for yet another book, but given the Sn of the topic, it comes as no surprise. And this one deliv- ers the goods in an informative, accessible way. The photographs catch the eye first. Swiss photo- grapher Thomas Marent is a well-travelled wildlife photographer who, starting young, has about 40 years experience in shooting pictures of various forms of wildlife that have been featured in a number of books, including the first edition of this one, an earlier book with Orenstein, and an earlier one yet of his own. The photos are consistently gorgeous, crisp in their detail, and beautifully presented. It would be very easy—and a big mistake—to treat this as a picture book! Ronald Orenstein is a Canadian zoologist/ornithologist, lawyer, wildlife conservationist, and prolific author/editor of natural history books. He admits in the Acknowledgements that he is “not an entomologist” (p. 224) but the text reveals an enviable capacity for digesting the lat- est research. The book opens with a lengthy intro- ductory outline of lepidopteran natural history that ranges from the origins of the term ‘butterfly’ and their cultural significance through their evolution- ary history and brief description of the six families into which they are now organized. The book covers wing formation and function, mimicry, mating and reproduction, host plants, development from egg to adult phases, issues of conservation, and much more. Orenstein isn’t shy about using scientific names and terms—androconia, for example—that are always defined in the text. We learn some surprising things, such as why but- terfly flight is erratic (pp. 9-10), the various types and roles of wing scales and the genetic coding that pro- BUTTERFLIES THEIR NATURAL HISTORY AND DIVERSITY duces their colours (p. 11), that mimicry in a particu- lar species can vary 1n time and place, the well-known Viceroy being an example (p. 13). Nuptial gifts and sperm competition (p. 16), pollination, migration, and DNA-based discoveries all receive concise, re- search-based accounts. One of the most interesting things I learned was that these lovely insects that can be such innocent symbols of beauty and grace are ca- pable of cannibalism and manipulative deception in symbiotic relations with other animals such as ants. Chapters 1 through 6 discuss the six families and their subfamilies, each of which receives brief intro- ductions. The photographs of species come into their own here, each identified with scientific name and lo- cation, followed by brief and informative comments on topics such as distribution, habitat, caterpillar stages, and toxicity. The next four chapters are the- matic, profusely illustrating and adding to the themes of the introduction: Butterfly Wings (Chapter 7), Butterfly Life History (Chapter 8), What Butterflies Eat (Chapter 9), and Butterflies in Their Environment (Chapter 10). The 11th and concluding chapter, Myriads of Moths, reminds us that “butterflies are moths” (p. 6) after all. Moth species “outnumber but- terflies by at least fifteen to one” so this chapter is “a miscellany, not a survey...” (p. 185). And a hand- some survey it is, covering some spectacular exam- ples, the caterpillars being particularly fascinating. The book concludes with a page of Further Reading that lists books and websites, plus a URL for the “400+ papers consulted...” (p. 219), and an Index. One odd thing: the book has two covers, the new one you can see here, and a reproduction of the origi- nal cover; it is this one, not the new cover, that says Second Edition. A minor puzzle for a worthy book. Books on butterflies seem to be written for profes- sional lepidopterists or for kids, with many popular ‘picture books’ in between. This book is one of the comparatively few that focus on natural history for the interested generalist who has some background in the topic. Orenstein and Marent have created a fine addition to such a reader’s library, one that informs while pointing the way to further study. BARRY COTTAM Ottawa, ON, and Corraville, PE, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 384 HERPETOLOGY THE CANADIAN FIELD-NATURALIST Vol. 133 The Field Herping Guide: Finding Amphibians and Reptiles in the Wild By Mike Pingleton and Joshua Holbrook. 2019. University of Georgia Press. 264 pages, 26.95 USD, Paper. An increasing number of r people are interested in amphibians and reptiles, or | ‘herps’, and this interest in- | cludes wanting to see them in nature. Many of these species can be somewhat challenging to find and a guide to finding herps is a good idea. The Field Herp- ing Guide does just this as well as discussing issues one should keep in mind to keep the herpers and the herps safe. It should be stressed that this is not a book about how to conduct scientific surveys of amphibi- ans or reptiles or how to design field ecology studies, this is a guide to finding herps for fun. The book consists of nine chapters with lots of col- our photographs. The chapter titles are a good indica- tion of the topics the book covers: Getting Started; Understanding Herp Behavior; Finding Herps; Catching and Handling Herps; Safety in the Field; Ethics and Etiquette, Rights and Responsibilities; Classification, Taxonomy, and Species Identification; Citizen Science and Data Collection; and Herp Photography. Several appendices on topics such as diseases, various kinds of public lands (mainly from an American point of view), internationally known herp hotspots, herp education, and the history of field herping round out the book. Is this a valuable book? The book is easy read- ing, but still contains a lot of information. Much of the advice seems very general, such as sometimes it is too hot for herps to be active, or often herps are active during or after it rains. Given the wide range of species covered, from salamanders to snakes, it is hard to generalize about herps. The authors do a good job of tackling each group of species, but even here the diversity is greater than many people real- ize. Salamanders can be completely aquatic and never leave the water, live along and in streams, depend upon temporary wetlands for breeding, or live in for- ests with no need of aquatic habitats. Overall, the authors provide useful advice on the diversity of life- styles and guidance for how and when to survey for different sub-groups of species. Unfortunately, the book also has many problems. A book that covers searching for venomous snakes should emphasize safety. The authors discourage people from catching venomous snakes and provide cautions about getting too close when photographing them, but then include a photo of someone in shorts and sandals with a snake hook and a venomous snake (p. 11). This is not the kind of lax safety precautions the authors should be encouraging. And despite urg- ing people not to catch venomous snakes the authors provide several methods for capturing venomous snakes (pp. 113-116). I also caught a surprising number of factual er- rors in the book. In the section on crocodilians, the authors give the distribution of Morelet’s Crocodile (Crocodylus moreletii) as being limited to Mexico and Guatemala but omit Belize (p. 89). In the sec- tion on frogs, it is incorrectly stated that cricket frogs (Acris spp.) are ranids or true frogs, when, in fact, they are hylids or treefrogs (p. 93). The authors state that Wood Frog (Lithobates sylvaticus) is the only herp in Alaska (p. 94), but this is not even remotely accurate as there are five other native amphibians. There are also a few things the authors could have stressed more. Near the beginning of the book the au- thors mention that insect repellant can be toxic to am- phibians (p. 18) but this fact is not mentioned again in the section on catching frogs (p. 126). Nor is there any mention of sunscreen on hands which can also be toxic to amphibians. I hope the authors prepare a second edition which corrects these things. While this book is not going to teach experienced herpetologists much about search- ing for herps, it is a great introduction to field herp- ing for those who are keen about herps but don’t have much experience. Even with the errors in this book it could still be a valuable resource at school libraries where it could kindle passion in a young reader. DAVID SEBURN Ottawa, ON, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 ORNITHOLOGY Gulls of the World: A Photographic Guide BooK REVIEWS 385 By Klaus Malling Olsen. 2018. Princeton University Press. 368 pages, 65 maps, and 600+ colour pictures, 45.00 USD, Cloth. As has been frequently noted, gulls can be a real pain to identify because, not only are many of the spe- cies very similar, but they exhibit changes of plumage with age to a greater degree than any comparable group of birds. Consequently, they 9 deserve, and have received, | much attention in the form of identification guides spe- cific to the group, starting with Peter Grant S (1982) classic, Gulls: A Guide to Identification. They con- tinue to attract enormous attention from birders, especially now that hybridization among species is known to be extensive. Facebook groups and ‘twit- terati’ agonise over the identity of individual birds (... probably a Western x Glaucous-winged ... almost certainly a second winter Thayer’s Gull ... ), some- times long after the bird has flown off into the sunset. Fifteen years ago, Klaus Malling Olsen, along with the artist Hans Larsson, produced a monumen- tal, 608-page guide to the gulls of the northern hem- isphere (Olsen and Larsson 2003) which dealt with their identification, voice, moult, plumage, and dis- tribution, including detailed range maps. The cur- rent book is a revision and expansion of the earlier book, although with much less detail on topics other than identification. In place of Larsson’s plates, the book is illustrated entirely with photographs, which, as pointed out by another reviewer, are an improve- ment on those in the first book. In fact, the book com- prises an unmatched collection of gull portraits and, as such, is an unmatched resource for identifying gulls in the field. Given the global spread of e-Bird since the ear- lier book, you might have expected an improvement in range maps as well, but I did not find that to be the case. The colour code in the current volume con- sists of yellow for breeding range, blue for wintering range, and green for “if no wintering area shown, oc- currence all year” (p. 29). In fact, I found no green areas on any maps. Consequently, where breeding and wintering areas overlap, as for American Herring Gull (Larus smithsonianus) on the Great Lakes, the KLAUS MALLING OLSEN reader cannot tell where the northern limit of the win- tering area is. Some rather strange errors in the maps have been perpetuated from the earlier book, includ- ing the breeding colony of Black-legged Kittiwakes (Rissa tridactyla) at Cape Cod (unnoticed so far by North American ornithologists) and the swath of Ivory Gulls (Pagophila eburnea) supposedly breed- ing across the western Queen Elizabeth Islands. Because of extensive hybridization among gull species, their taxonomy is contentious. For exam- ple, Olsen treats Thayer’s and Iceland Gulls (Larus thayeri, Larus glaucoides) and American and Euro- pean Herring gulls (Larus smithsonianus, Larus ar- gentatus) as different species, whereas the American Ornithological Society now regards Thayer’s as a subspecies of Iceland (L. g. thayeri) and continues to treat North American Herring Gulls as conspecific with their European counterparts. Since Grant’s book in 1982, four species dealt with here have been carved out of his “Herring Gull”. One small reservation I have about treating this book as the last word on gull identification (a reser- vation I also hold about the opinions of many experts that I read on the web) is that few of the contentious identifications are backed up by genetic material. Consequently, I cannot see how many of the identi- fications can be treated as better than ‘best guesses’. One of the problems of a book like this is that some- one might find a gull in the field identical to one of Olsen’s pictures and consequently feel confident in the identification. But what if Olsen was wrong? Peter Adriaens and Amar Ayyash, on the American Birding Association website, give a list of errors that they found, including several identifications that they consider erroneous (http://blog.aba.org/errata-gulls- of-the-world). Nothing is perfect and we need to keep that in mind. In science, all is provisional. Literature Cited Grant, P.J. 1982. Gulls: A Guide to Identification. T. & A.D. Poyser, Calton, England. Olsen, K.M., and H. Larsson. 2003. Gulls of Europe, Asia and North America. A&C Black, London, United Kingdom. ToNy GASTON Ottawa, ON, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 386 ZOOLOGY THE CANADIAN FIELD-NATURALIST Vol. 133 The Flying Zoo: Birds, Parasites, and the World They Share By Michael Stock. 2019. University of Alberta Press. 260 pages, 29.99 USD, Paper Michael Stock’s book, The Flying Zoo: Birds, Para- sites, and the World They Share, provides an intrigu- ing glimpse into the lives of birds and their parasites, which are usually looked © upon with disgust and dis- > missed as worthless vermin. _ However, parasites may provide benefits to their hosts, and Stock’s narra- tive breathes new life into “ the world of these often-misunderstood organisms. The author asks “How has this weird association be- tween one organism (a bird) and its fellow travelers (parasites) become normal? What special adaptations have parasites had to evolve to be able to find, colo- nize, and survive in or on their hosts? ... How have hosts evolved to survive with their ‘zoological gar- den’?” (p. 4). These questions, and many more, are examined and explored with vigour and enthusiasm. The book is divided into 10 chapters: A World on a Bird; Lice: It’s a Beautiful Life; Fleas: The Circus in the Zoo; Tough Ticks; Mites: Little Things Mean a Lot; Flying Zoo Flies; The Worms that ate the Bird; Oddities in the Flying Zoo; Flying Zoo Behaviour; and Environmental Impacts: The Future of the Flying Zoo. Also included are a Notes section, Further Reading References, and an Index. More than 30 highly detailed pen and ink illustrations of the parasites in question are also dispersed throughout the book. The book is a joy to read; the author crafts a fas- cinating journey into the lives of birds and their para- sites using current research cases, vivid descriptions, and subtle humor. Co-evolutionary themes are com- monplace and connect ecology, biology, adaptation, and survival into a seamless narrative. The reader travels the world, from Madagascar to the Americas, exploring the various interactions between parasite and host. Some amazing information comes to light from Stock’s research: who knew that fleas could sing or that there is a specialized moth that drinks the tears of sleeping birds? One of the perks of the book is that the author defines various biological terms (some- times breaking down the Latin or Greek root words), a major help to those first encountering the term or a refresher for seasoned naturalists and biologists. All of the classic parasitic groups are covered, in- cluding fleas, ticks, lice, flies, and worms. However, peripheral species are also addressed, such as bed bugs (but for birds), moths, leeches, and strange critters called tongue worms. Figure 1.1 on p. 3 il- lustrates the parasitic relationship with birds well: it shows a Common Pigeon (Columba livia) surrounded by various parasitic species that may occur on and within a single bird, from roundworms, flukes, and tapeworms occurring inside the bird to mites, ticks, fleas, and lice occurring on the outside, each occu- pying a specialized niche (hence the idea of a “flying zoo’). One of the more fascinating topics Stock ex- plores is the niche theory, which states that in order to reduce direct competition, species evolved to occupy different habitats or feeding behaviours. For example, a single bird may support several species of lice, but these lice live in different parts of the bird, such as on various locations of the wing (either on the feathers or inside the quill), on the head, or near the skin. In addi- tion, these lice evolved different body shapes in order to avoid being detected or removed by the bird during the preening process. It is unwise to assume parasites are worthless crea- tures with no value, and Stock provides several exam- ples. Worms, such as blood flukes, have evolved ways to not be attacked by the host bird’s immune system by down-regulating the immune response. Humans with auto-immune diseases may benefit in the future when we figure out how flukes and other parasites al- ter host immune systems (p. 150). Leeches, in another example, have a protein anticoagulant in their saliva called hirudin. The anticoagulant is now commer- cially produced and used to treat people with cardi- ovascular problems (p. 166). In addition, sometimes parasites provide advantages to their hosts. For ex- ample, wild Mute Swans (Cygnus olor) have a mature community of co-evolved helminths (worms). Mute Swans in a zoo environment, on the other hand, are not exposed to their usual worm parasites and were infected with two rare tapeworm species causing ma- jor infections and significantly diseased birds. The normal worm parasites are apparently a benefit to the swan by preventing harmful helminths from infecting the host. The co-evolved relationship between para- site and host seems to lead to a peaceful co-existence (p. 143). In another example, feather mites may bene- fit hosts by eating bacteria and fungi trapped in preen gland oil. These bacteria and fungi, in large numbers, may make a bird look unhealthy or diseased, but the 2019 mites, by consuming these organisms, allow a bird to appear to have bright and healthy plumage, aiding in their reproductive success (p. 182). Co-evolution between hosts and parasites is not novel. A few “rules” have been established by re- searchers exploring the idea. The first, known as Fahrenholz’s Rule, claims “that parasite evolution- ary histories, or phylogenies, should mirror the his- tories of their hosts” (p. 142); that is, hosts that are re- lated evolutionarily may harbour the same parasites. A second rule, called Manter’s Rule, states “that long associations between hosts and parasites should lead to strong host specificity” (p. 142) and that “para- sites should speciate more slowly than their hosts” (p. 24). The third rule, Eichler’s Rule, states that “a large taxonomic group of hosts ... should have more gen- era and species of parasites than a small taxonomic group” (p. 24). The fourth rule, Szidat’s Rule, claims “more recent or specialized host groups should have more recent or specialized parasites while more prim- itive or generalized hosts should have more primitive or generalized parasites” (p. 24). Finally, Harrison’s Rule states that “large-bodied species of hosts should BooK REVIEWS 387 have large-bodied parasites” (p. 24). Indeed, Stock explores these relationships throughout the book. Overall, the book is a must-read for those inter- ested in the intricate and interwoven world of birds and their parasites. The author emphasizes that it would be a mistake for anyone interested in avian bi- ology to ignore that parasites are a real and significant part of the lives of birds. Parasites influence many as- pects of the lives of our feathered friends, from sex- ual selection to healthy co-evolutionary relationships. A bird parasite may be harmful, beneficial, or indif- ferent, and any single parasite can fulfill any one of these roles to all three. Host-parasite studies will continue to lead to more questions and puzzles, es- pecially with the looming climate change crisis, and Stock has provided a good starting point on this jour- ney with his book The Flying Zoo. Acknowledgement: I thank Susan Hagen for im- proving the manuscript. Howarb O. CLARK, JR. Colibri Ecological Consulting, LLC, Fresno, California, USA ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 388 Orca: The Whale Called Killer. Fifth Edition By Erich Hoyt. 2019. Firefly. 320 pages, 24.95 USD, Paper. Orca: The Whale Called Killer is a really great read. Erich Hoyt has been study- ing whales for a long time, and his knowledge of the Killer Whale (or Orca) shines through in this book. Hoyt leads readers through his first three summers (1973-— 1975) documenting North- ern Resident Killer Whales around Johnstone Strait, northern Vancouver Island. Hoyt and his colleagues were filming, photographing, and recording the underwater vocalizations of Killer Whales to make documentaries on them. At this time, very little was known about Killer Whales. For exam- ple, now we know that there are four different types of Killer Whales in British Columbia (BC): the northern and southern populations of resident, salmon-eating ecotypes; the transient, mammal-eating ecotype; and the offshore shark specialist ecotype. But in 1973, biologists did not know that these Killer Whales were different. The book focusses on the timeline of Hoyt’s exploits in the field, including how he learned new things about Killer Whales during his adven- tures. This book is partly set up like a field notebook or diary, with frequent excerpts from Hoyt’s field notes, which I found an effective style to portray the story. This is the fifth edition of the book, but ac- cording to Hoyt, the last substantial update occurred in the 1990 version (third edition), so this new edi- tion adds information gleaned about Killer Whales over the past 30 years. This new edition has a new in- troduction detailing important events that have hap- pened with Killer Whales since the 1990 version of this book. It also includes an expanded afterword, epilogue, and bibliography. Hoyt’s tales of whale watching in the wild are also interwoven with the looming reality of Orca capture events that were happening concurrently. At this time, Killer Whales in BC and Washington State were actively being captured and sold to aquaria world- THE WHALE CALLED KILLER FIFTH EDITION + UPDATED AND EXPANDED THE CANADIAN FIELD-NATURALIST Vol. 133 wide. In many ways, the live capture events of Killer Whales and the early days of Orcas in captivity are what sparked Hoyt’s interest in spending entire sum- mers on the water to learn about the wild whales that were barely known by science at the time. Hoyt’s first years in the field also happened at the same time and in the same locations as famed Killer Whale biolo- gist Michael Biggs, who collected incredibly impor- tant information for the Canadian government about all of the Killer Whales along the coast of BC, which helped lead to the end of the live capture of Killer Whales in Canada. The historical context of this book is one of its great features. My favourite part of this book is the way that Hoyt brings everything together in the final chapter. Hoyt was a huge proponent for an ecological reserve that was established for these Killer Whales in Robson Bight, and in this final chapter he discusses a lot of the rationale for why and how that process actually happened. Since Hoyt’s early days of studying Killer Whales, he has become a global proponent for marine protected areas as a tool for conserving whales, and his early work on this ecological preserve on northern Vancouver Island clearly paved the way for his future work on marine protected areas. This book would be a great read for any natural- ists interested in learning about Killer Whales, both the natural history of populations in BC, but also the history surrounding their conservation and protec- tion in Canada. This may be of particular interest to those who have been following the recent efforts of Fisheries and Oceans Canada to study and protect Southern Resident Killer Whales, which are closely related to the Northern Resident Killer Whales that Hoyt followed in this book. For those interested in a comparison between Killer Whales in the wild ver- sus those in captivity, this book also provides a lot of useful context. WILLIAM D. HALLIDAY Wildlife Conservation Society Canada, Whitehorse, YT, and Department of Biology, University of Victoria, Victoria, BC ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 BooK REVIEWS 389 Mammals of Prince Edward Island and Adjacent Marine Waters By Rosemary Curley, Donald F. McAlpine, Dan McAskill, Kim Riehl, and Pierre-Yves Daoust. 2019. Island Studies Press. 354 pages, 49.95 CAD, Paper. Ok, go ahead, judge this book by its cover ... it is quite stunning! A Red Fox (Vulpes vulpes), seemingly just aroused from its slum- ber to look at the photog- rapher ... its tail wrapped around itself, while resting | on the snow ... what a per- fect shot to entice a shopper to take a copy off the book- store shelf! This is a thorough book—over 1000 references were used! The introduction provides a background and synopsis of Prince Edward Island’s mammals, covering both extirpations and (re)introductions. Large-scale factors influencing mammals, including climate change and white-nose syndrome, are intro- duced; these are treated in more depth further in the book. Here, domestic animals are given mention, and dismissed from further representation in the book. This book covers 57 species of mammal, essen- tially split evenly between the marine and terrestrial environments. I believe one is missing, but I'll defer that discussion. Each account includes a colour illus- tration of the animal, a range map (North American distribution, or beyond), and a diagram of the skull from three perspectives (dorsal, palatal, and lateral). Sometimes, there is also a photograph. For most non- volant, terrestrial species, at least one trackway, and an accompanying more-detailed illustration of a hind and fore footprint, are included. Five of these track- ways appear only as series of irregular grey shapes, clearly a printing error, for which there was no ex- cuse; one hopes that a second printing clears this up. The text for each account is very well organ- ized and the writing is clear and consistent, not a small feat for a book with so many authors. Short sections include description, range, and status (now and earlier) whereas most of the accounts encompass the species’ ecology, often running several pages. History on the island is detailed, which, when appro- priate, includes introduction and extirpation dates and details of these events. The missing species from this book is_ the Domestic (free roaming) Cat (Felis catus). Other in- troduced species are included—Bobcat (Felis ru- fus), Brown Rat (Rattus norvegicus), Eastern Gray Squirrel (Sciurus carolinensis)—so why not the infi- nitely problematic free roaming house cat? In Prince Edward Island, just like other jurisdictions, there are not only individuals who let their cats run amok, there are still those misguided people who promote sup- ported colonies of these wildlife destroyers. Omitting the Domestic Cat from this book was a missed oppor- tunity for further education. This book is well-suited to people with a general interest (an extensive glossary was included, and will be much appreciated), but adept naturalists will still learn a lot. The previously mentioned voluminous ref- erence section will serve as a start to finding more in- formation for mammal enthusiasts of any level. RANDY LAUFF Biology, St. Francis Xavier University Antigonish, NS, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 390 OTHER THE CANADIAN FIELD-NATURALIST Vol. 133 Surviving Global Warming: Why Eliminating Greenhouse Gases Isn’t Enough By Roger A. Sedjo. 2019. Prometheus Books. 245 pages, 24.00 USD, Cloth, 22.50 USD, E-book. Dr. Roger Sedjo is a Senior gy GLOBAL Fellow Emeritus at the en- vironmental think tank Re- sources for the Future in Washington, DC. He spe- cializes in forestry and pol- icy, holds several honour- ary degrees, awards, and fellowships, and shared the 2007 Nobel Peace Prize for his work on the Intergov- ernmental Panel on Cli- mate Change (IPCC) Cli- mate Assessments. Despite these qualifications, I struggled through this book. The core argument is straightforward: climate change is inevitable and will have dramatic, unavoid- able impacts on human society. Even our most am- bitious mitigation solutions will not stop this inevi- tability, so we must invest in adaptation solutions. For anyone engaged in the climate change conversa- tion, this is not a new idea: most government climate change strategies in Canada recommend both miti- gation and adaptation measures. Climate change ad- aptation is not controversial, so I am puzzled by Dr. Sedjo’s insistence that to justify adaptation efforts, he must discredit the need for mitigation. Sedjo dedicates the first third of the book to scru- tinizing what he calls “Al Gore’s theory of global warming” (Chapter 1, Al Gore and the Greenhouse Gas Theory: Plan A), that is, the theory that recent cli- mate change has been caused by increases in green- house gas (GHG) emissions from humans. He seems to think that if he can convince the reader that the cli- mate is changing at least in part from natural causes, then the reader will also be convinced that mitigation is a waste of time: if GHG emissions are not the en- tire problem, GHG reduction cannot be the whole so- lution. I cannot help but recall Joel Pett’s well-known political cartoon from 2009 depicting delegates at a climate summit: “What if it’s a big hoax and we cre- ate a better world for nothing?” In order to cast doubt on “Gore’s theory”, the au- thor spends quite a bit of time discussing evidence of natural climate change, including the existence of the medieval warming period in the climate record and the role of solar cycles (see Chapter 2, Natural Climate Change: GHG’s Are Not the Whole Answer). He writes that “solar energy is not currently viewed as a major contributor to today’s warming by the WHY ‘.. ELIMINATING GREENHOUSE IPCC. However, solar factors are still not yet well un- derstood” (p. 37). Out of curiosity, I googled “Is the sun causing climate change?” The first result, from the National Aeronautics and Space Administration (NASA), starts off this way: “No. The Sun can influ- ence the Earth’s climate, but it isn’t responsible for the warming trend we’ve seen over the past few decades” (National Aeronautics and Space Administration 2020). Around this point I started to lose patience for his deep-dives into the medieval warming period and frustrating lack of understanding when it comes to basic climate science, for example: “How rapidly will land-based glaciers melt, and will future snows offset much of that melting?” (p. 18). (The answer is no—warming temperatures will offset any possible increases in snowfall because the melting will out- pace the rate of accumulation [National Snow & Ice Data Centre 2020].) There is so much repetition of the same poorly referenced material that I often had the disorienting feeling that I had read the same para- graph multiple times. The bulk of the book—Chapters 4 through 8— is dedicated to “Plan B: The Adaptation Solution”. Some of Sedjo’s ideas are reasonable: for example, he writes about the importance of coastal habitat protec- tion to buffer sea level rise (p. 103). But as an ecolo- gist, I find many of his ideas disturbing. In his discus- sion on the relative albedos of different surface types, he writes: “So, Mother Nature being complicated, those who are cutting down the Amazon rainforest could be seen by some as countering global warming instead of aggravating” (p. 153). I’m still not clear if that is supposed to be a joke or not. The section on geoengineering (Chapter 5) is a lit- any of potential projects that sound rather extreme: carbon capture and storage, seeding the atmosphere with sulphur dioxide, or “moderating the atmosphere with calcium carbonate particles” (p. 145). This in- sistence on adaptation over mitigation confuses me, because, even from a strictly economic viewpoint, minimizing our GHG emissions now will make adap- tation in the future cheaper because there will be less carbon in the atmosphere. The only answer I came up with is a fear of the drastic changes that must occur to transition away from a fossil fuel economy. In the final pages, Sedjo states that natural gas could be the remarkable solution that we need; it’s a bit anti-climactic. Even Sedjo admits that “in the long term, it is only a part of a more environmentally friendly energy transition. The question is: a transi- 2019 Book REVIEWS 391 tion to what?” (p. 208). Now that sounds like the first global-warming/. line of a book that we need today. National Snow & Ice Data Center. 2020. Quick facts on ice sheets. Accessed 2 April 2020. https://nsidc.org/ Literature Cited cryosphere/quickfacts/icesheets.html. National Aeronautics and Space Administration. 2020. Is the sun causing global warming? Accessed 2 April 2020. EMMA BOCKING https://climate.nasa.gov/faq/14/is-the-sun-causing- Halifax, NS, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). B92 THE CANADIAN FIELD-NATURALIST Vol. 133 How to Walk on Water and Climb up Walls: Animal Movement and the Robots of the Future By David L. Hu. 2018. Princeton University Press. 240 pages, 24.95 USD, Cloth or E-book. Albert Einstein (1955: 64) once said that “The im- portant thing is not to stop questioning. Curiosity has its own reason for existing. One cannot help but be in awe when he contemplates the mysteries of eternity, of life, of the marvelous structure of reality. It is enough if one tries merely to comprehend a little of this mystery every day”. One can keep thinking about curiosity towards all as- pects of life while reading How to Walk on Water and Climb up Walls: Animal Movement and the Robots of the Future. In his book, David L. Hu, an associate professor of mechanical engineering and biology and adjunct pro- fessor of physics at Georgia Institute of Technology, tells us about his research and through that teaches us how to maintain curiosity and approach research questions. Specifically, in his research he tries to find and focus on the principles of animals’ movements and apply these to robots. However, the book does not cover all animal motions, but generally those on which the author has done experimentations. For ex- ample, in the first chapter he shows us how the wa- ter strider’s motion and ability to stand on water has inspired a water-walking robot. And in the sec- ond chapter we learn about the principles of crawl- ing animals’ (snakes and sandfish) movements. Next, Hu tells us about animals (e.g., jellyfish) that use their body parts to influence the flow of fluids for their own advantage. We learn about surface structure of ani- mal bodies such as sharkskin (Chapter 4), and body structures (Chapter 5) that could be used to develop machines that are capable of moving underwater or on land with decreased energy expenditure or to de- sign wearable devices—exoskeletons—that could lower the energy costs of human walking. In the sixth chapter we learn how insects deal with collisions and how engineers are inspired by these in- sects as potential applications to robots. For example, how mosquitoes survive being struck by raindrops, How to Walk on Water and Climb up Walls bees survive crashing into obstacles such as flowers and plants, and cockroaches squeeze themselves into very narrow spaces. Then the author tells us how ani- mals automatically respond to their surroundings, us- ing the examples of flies overcoming turbulence dur- ing their flight and the cockroach’s ability to measure its distance from the obstacles during quick running (Chapter 7). Finally, we learn about ants’ ability to link their bodies to create a flow like liquid, form bridges, or spring back like a solid. How to Walk on Water and Climb up Walls is in- teresting for those curious minds learning how one can do experimentation, as throughout the book Hu details the steps of his experiments and how he has overcome the problems during the process of experi- mentation. This book is for general readers interested in scientific inquiries as it teaches the way one should pursue them. It is full of colourful stories, a joyous read for curious minds, making it an easy read for laymen and even school students. The book is written from an engineering rather than a biological perspective and one may speculate about evolutionary and adaptive mechanisms while reading the book. Hu writes that “animal motion is all around us. It is the principal way animals get things done in the world. How did such a diversity of animal movements come about?” (p. 4), but the book does not tell us how these animals might have adapted and does not therefore generalize for other species. Altogether, the book is for general readers inter- ested in learning how scientific inquiry should work and how a scientist thinks and does experiments. That is a very interesting, fun, thought-provoking, story-based, and amusing book for undergraduate and high school students interested in physics, robot- ics, fluid mechanics, mechanical engineering, and re- lated disciplines. Literature Cited Einstein, A. 1955. Old man’s advice to youth: ‘never lose a holy curiosity’. Life Magazine 1955 (May): 64. FARID PAZHOOHI Department of Psychology, University of British Columbia, Vancouver, BC, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 2019 BooK REVIEWS 393 Frog Pond Philosophy: Essays on the Relationship Between Humans and Nature By Strachan Donnelley. 2018. University Press of Kentucky. 248 pages, 38.17 CAD, Paper. If you like baseball, Frog Pond Philosophy will in- trigue you. If you like hunt- ing and fishing, particu- larly fly fishing, Frog Pond Philosophy will appeal to you. If you like thinking about the natural world and how we relate to it, Frog Pond Philosophy will inter- est you. If you like music, sitting by the water in springtime, and listening to frog song, Frog Pond Philosophy will be certain to charm you—particularly the essay that inspires the title of the book. It was one of my favour- ite pieces in this collection of writings by the late Strachan Donnelley (1942-2008), an environmental philosopher and bioethicist who focussed on studying the intricacies of human-nature relations. Donnelley was also the founder and first president of the Center for Humans and Nature (https://www. humansandnature.org), an initiative which portrays itself as exploring and promoting human responsi- bilities in relation to nature and the whole commu- nity of life. The Center’s website describes Donnelley as rejecting reductionist, silo thinking, and bring- ing together ideas from many corners, including biology, ecology, economics, engineering, poetry, the arts, and philosophy. This breadth of perspec- tives is reflected in the subject matter of Frog Pond Philosophy essays, which span over four decades of work, and range from hard-core philosophical inter- pretations to reminiscences on personal encounters and outdoor experiences. No wonder it took the editors—daughter Ceara Donnelley and colleague Bruce Jennings—so many years to publish the book. It is clearly a labour of love and respect. As Ceara explains in the Editor’s Afterword, her father spent the final months of his life re-reading, mulling over, and assembling the man- uscript from new and previously published pieces. Shaping the final collection was obviously a task that could not be hurried. The editors organized the essays into four sec- tions. The first two are introductory and more gen- erally reflective; the last two are more intensely FROG POND Essays on the Relationship Between Humans and Nature STRACHAN DONNELLEY Edited by Crara Donnelley and Bruce Jennings Foreword by Frederick L. Kirschenmann philosophical. The pieces vary in length, many of them short. The content of the first two sections, aptly named Two Preludes and A Guide for the Naturally Perplexed, was the most compelling, and the essays were easy to read and comprehend. Frog Pond Philosophy was my favourite essay—partly because I love frog song and partly because the content lines were simple. The image of Homo sapiens singing alongside innumer- able other organisms in a great planetary frog pond adds to the essay’s appeal, along with the closing par- agraph of the essay where Donnelley calls for insight from “bullfrog philosophers” in the “urgent busi- ness” of saving “our earthly frog pond” (p. 35). In the essay Bottom Lines and the Earth’s Future, he de- scribes the similarly urgent business of replacing the prevailing economic bottom line with an ecological “nature alive” (p. 51) bottom line—another important message in the current global context of biodiversity loss and climate instability. The intensely philosophical essays in the final two sections of the book delve into diverse philosophical traditions. Connelly revisits them from different an- gles with the purpose, in the words of Jennings, of “thinking ‘humans’ and ‘nature’ together” (p. 219) and overcoming the separation of “human being” from “the rest of natural being” (p. 220) encouraged by dominant modern philosophical and scientific the- ory. I confess that I was less motivated to read those pieces thoroughly. I was easily discouraged from wading through them in detail because of their dense- ness and complexity. But you might not be. The baseball and fly-fishing analogies introduced early in the book did not, unfortunately, resonate with me. They had, in fact, the opposite effect of aliena- tion, and the feeling did not dissipate easily. But that might not happen to you. I was pleased to learn that proceeds from the sale of Frog Pond Philosophy are being donated by the University Press of Kentucky to the Center for Humans and Nature. That fact, along with the essays in the first two sections of the book, and the sec- ond if you are so inclined, would make it worth the purchase. RENATE SANDER-REGIER Environmental Studies, University of Ottawa, Ottawa, ON, Canada ©The author. This work is freely available under the Creative Commons Attribution 4.0 International license (CC BY 4.0). 394 NEw TITLES Prepared by Barry Cottam THE CANADIAN FIELD-NATURALIST Vol. 133 Please note: Only books marked 7 or * have been received from publishers. All other titles are listed as books of potential interest to subscribers. Please send notice of new books to the Book Review Editor. +Available for review *Assigned Currency Codes: CAD Canadian Dollars, AUD Australian Dollars, USD United States Dollars, EUR Euros, GBP British Pounds. BIOLOGY Biology of Floral Scent. Edited by Natalia Dudareva and Eran Pichersky. 2019. CRC Press. 346 pages, 176.00 USD, Cloth, 59.96 USD, Paper, 48.72 USD, E-book. Biology of Plant Volatiles. Second Edition. By Eran Pichersky and Natalia Dudareva. 2020. CRC Press. 408 pages, 140.00 USD, Cloth. The Call of Carnivores: Travels of a Field Biol- ogist. By Hans Kruuk. 2019. Pelagic Publishing. 200 pages, 176 colour illustrations, and 55 drawings, 33.73 CAD, Paper. Cry Wolf: Inquest into the True Nature of a Predator. By Harold R. Johnson. 2020. University of Regina Press. 160 pages, 16.95 CAD, Paper. The Invertebrate Tree of Life. By Gonzalo Giribet and Gregory D. Edgecombe. 2020. Princeton Uni- versity Press. 608 pages, 85.00 USD, Cloth. Sexual Selection: A Very Short Introduction. By Marlene Zuk and Leigh W. Simmons. 2018. Oxford University Press. 160 pages, 11.95 USD, Paper. BOTANY Darwin’s Most Wonderful Plants: A Tour of His Botanical Legacy. By Ken Thompson. 2019. Uni- versity of California Press. 256 pages, 25.00 USD, Cloth, 18.00 USD, E-book. *Flora of Florida, Volume VII: Dicotyledons, Oro- banchaceae through Asteraceae. By Richard P. Wunderlin, Bruce F. Hansen, and Alan R. Franck. 2020. University Press of Florida. 492 pages, 70.00 USD, Cloth. Mosses of the Northern Forest: A Photographic Guide. The Northern Forest Atlas Guides Series. By Jerry Jenkins. 2020. Cornell University Press, Comstock Publishing Associates. 176 pages, 1435 colour photos, and 1321 diagrams, 16.95 USD, Paper, 11.95, USD, Fold-out Chart. Plant Names: A Guide to Botanical Nomenclature. Fourth Edition. By Roger Spencer and Rob Cross. 2020. CSIRO Publishing. 168 pages, 44.99 AUD, Paper. Also available as an E-book. Plants on Islands: Diversity and Dynamics on a Continental Archipelago. By Martin L Cody. 2019. University of California Press. 270 pages, 70.00 USD, Cloth, 57.95 USD, E-book. Plant Systematics. Third Edition. By Michael G. Simpson. 2019. Academic Press. 774 pages, 105.00 USD, Paper, 89.25 USD, E-book. Tree Story: The History of the World Written in Rings. By Valerie Trouet. 2020. Johns Hopkins University Press. 256 pages, 27.00 USD, Cloth. Wild Urban Plants of the Northeast: A Field Guide. By Peter Del Tredici. Foreword by Steward T.A. Pickett. 2020. Comstock Publishing Associates. 428 pages and 965 colour photos, 34.95 USD, Paper, 16.99 USD, E-book. CLIMATE CHANGE The Citizen’s Guide to Climate Success: Over- coming Myths that Hinder Progress. By Mark Jac- card. 2020. Cambridge University Press. 292 pages, 68.95 CAD, Cloth, 22.95 CAD, Paper, 16.00 CAD, E-book. Entertaining Futility: Despair and Hope in the Time of Climate Change. By Andrew McMurry. 2018. Texas A&M University Press. 224 pages, 27.00 USD, Paper. Also available as an E-book. Waters of the World: The Story of the Scientists Who Unraveled the Mysteries of Our Oceans, Atmosphere, and Ice Sheets and Made the Planet Whole. By Sarah Dry. 2019. UCP. 368 pages, 30.00 USD, Cloth, 18.00 USD, E-book. ENTOMOLOGY *Butterflies: Their Natural History and Diversity. Second Edition. By Ronald Orenstein. Photography by Thomas Marent. 2020. Firefly Books. 224 pages, 24.95 CAD / USD, Paper. Courtship and Mating in Butterflies. By R.J. Cannon. 2020. CABI. 384 pages, 135.00 USD, Cloth. 2019 Desert Navigator: The Journey of an Ant. By Rudiger Wehner. 2020. Harvard University Press. 400 pages, 18 colour photos, and 153 colour illustra- tions, 59.95 USD, Cloth. The Insect and Spider Collections of the World. Second Edition. By Ross H. Arnett, Jr, G. Allan Samuelson, and Gordon M. Nishida. 2020. CRC Press. 316 pages, 119.00 USD, Cloth, 58.50 USD, E-book. Insect Metamorphosis: From Natural History to Regulation of Development and Evolution. By Xavier Belles. 2020. Academic Press. 304 pages, 120.00 USD, Paper or E-book. Nature Underfoot: Living with Beetles, Crabgrass, Fruit Flies, and Other Tiny Life Around Us. By John Hainze. Illustrated by Angela Mele. 2020. Yale University Press. 254 pages, 28.00 USD, Cloth. Extraordinary Insects: Weird. Wonderful. Indis- pensable. The Ones Who Run Our World. By Anne Sverdrup-Thygeson. 2019. HarperCollins, Mud- lark Imprint. 320 pages, 14.99 GBP, Cloth, 9.99 GBP, Paper, 5.99 GBP, E-book. Spiders of the World: A Natural History. By Nor- man I. Platnick. 2020. Princeton University Press. 240 pages, 29.95 USD, Cloth. HERPETOLOGY On the Backs of Tortoises: Darwin, the Galapagos, and the Fate of an Evolutionary Eden. By Elizabeth Hennessy. 2019. Yale University Press. 336 pages, 30.00 USD, Cloth. Reptiles and Amphibians of New Zealand. By Dylan Van Winkel, Marleen Baling, and Rod Hitch- mough. 2020. Princeton University Press, Princeton Field Guides. 368 pages, 35.00 USD, Paper. Secrets of Snakes: The Science beyond the Myths. W.L. Moody, Jr., Natural History Series. By David A. Steen. 2019. Texas A&M Press. 184 pages and 103 colour photos, 25.00 USD, Flexbound. Also available as an E-book. ORNITHOLOGY Barn Owls: Evolution and Ecology with Grass Owls, Masked Owls and Sooty Owls. By Alexandre Roulin. Illustrated by Laurent Willenegger. 2020. Cambridge University Press. 300 pages and 50 colour illustrations, 68.95 CAD, Cloth, 48.00 USD, E-book. Bird Love: The Family Life of Birds. By Wenfei Tong. 2020. Princeton University Press. 192 pages and 220 colour photos, 29.95 USD, Cloth or E-book. NEw TITLES 395 Birds in Minnesota. Revised and Expanded Edi- tion. By Robert B. Janssen. Foreword by Carrol L. Henderson. 2020. University of Minnesota Press. 624 pages, 315 colour plates, and 1100 maps, 34.95 USD, Paper. In Search of Meadowlarks: Birds, Farms, and Food in Harmony with the Land. By John M. Marz- luff. 2020. Yale University Press. 352 pages, 28.00 USD, Cloth. Silent Spring Revisited. By Conor Mark Jameson. 2019. Bloomsbury USA. 288 pages, 20.00 USD, Paper. Also available as an E-book. Waterfowl of Eastern North America. Second Edition. By Chris G. Earley. 2020. Firefly Books. 168 pages and 400 colour photographs, 19.95 CAD, Paper. What It’s Like to Be a Bird: From Flying to Nest- ing, Eating to Singing—What Birds Are Doing, and Why. By David Allen Sibley. 2020. Knopf Publishing Group. 240 pages and 430 colour illustra- tions, 35.00 USD, Cloth. ZOOLOGY Bat Basics: How to Understand and Help These Amazing Flying Mammals. By Karen Krebbs. 2019. Adventure Publications. 140 pages, 22.50 CAD, Paper. Becoming Wild: How Animal Cultures Raise Families, Create Beauty, and Achieve Peace. By Carl Safina. 2020. Henry Holt and Co. 384 pages, 29.99 USD, Cloth, 14.99 USD, E-book. Published in England as Becoming Wild: How Animals Learn to be Animals by Oneworld Publications. Felines of the World: Discoveries in Taxonomic Classification and History. By Giovanni Giuseppe Bellani. 2019. Academic Press. 486 pages, 89.95 CAD, Paper or E-book. Guide to the Identification of Marine Meiofauna. Edited by Andreas Schmidt-Rhaesa. 2020. Verlag Friedrich Pfeil. 607 pages, 68.00 EUR, Cloth. Humans and Lions: Conflict, Conservation and Coexistence. By Keith Somerville. 2019. Routledge. 234 pages, 120.00 GBP, Cloth, 34.99 GBP, Paper. Also available as an E-book. Invasive Wild Pigs in North America: Ecology, Impacts, and Management. Edited by Kurt C. VerCauteren, James C. Beasley, Stephen S. Ditch- koff, John J. Mayer, Gary J. Roloff, and Bronson K. Strickland. Foreword by Dale L. Nolte. 2020. CRC Press. 480 pages, 88 colour and 37 black and white 396 illustrations, 200.00 USD, Cloth, 79.95 USD, Paper or E-book. Invertebrate Embryology and Reproduction. By Fatma Mahmoud El-Bawab. 2020. Academic Press. 931 pages, 165.00 CAD, Paper or E-book. Investigation and Monetary Values of Fish and Freshwater Mollusk Kills. American Fisheries Society, Special Publication 35. Edited by Robert I. Southwick and Andrew J. Loftus. 2017. 165 pages, 79.00 USD, Hardcopy or PDF download. Mammalogy: Adaptation, Diversity, Ecology. Fifth Edition. By George A. Feldhamer, Joseph F. Merritt, Carey Krajewski, Janet L. Rachlow, and Kelley M. Stewart. 2020. Johns Hopkins University Press. 744 pages, 600+ photos, maps, and illustrations, 124.95 USD, Cloth. Also available as an E-book. Pangolins: Science, Society and Conservation. Edited by Daniel W.S. Challender, Helen Nash, and Carly Waterman. 2019. Academic Press. 658 pages, 110.00 CAD, Cloth or E-book. Supernavigators: Exploring the Wonders of How Animals Find Their Way. David Barrie. 2019. The Experiment. 336 pages, 25.95 USD, Cloth. The North Atlantic Right Whale: Past, Present, and Future. By Joann Hamilton-Barry. 2019. Nimbus Publishing. 104 pages, 19.95 CAD, Paper. Whales of the Southern Ocean: Biology, Whaling and Perspectives of Population Recovery. Advances in Polar Ecology Volume 5. By Yuri Mikhalev. 2020. Springer Nature. 382 pages, 179.99 USD, Cloth, 139.00 USD, E-book. OTHER +A Sand County Almanac and Sketches Here and There. By Aldo Leopold. Introduction by Barbara Kingsolver. 2020. Oxford University Press. 240 pages, 15.95 CAD, Paper. Ahab’s Rolling Sea: A Natural History of “Moby- Dick”. By Richard J. King. 2019. University of Cali- fornia Press. 464 pages, 30.00 USD, Cloth, 18.00 USD, E-book. Amber Waves: The Extraordinary Biography of Wheat, from Wild Grass to World Megacrop. By Catherine Zabinski. 2020. University of California Press. 216 pages, 24.00 USD, Cloth or E-book. Charles Darwin’s Barnacle and David Bowie’s Spider: How Scientific Names Celebrate Adven- turers, Heroes, and Even a Few Scoundrels. By Stephen B. Heard. Illustrations by Emily S. Damstra. THE CANADIAN FIELD-NATURALIST Vol. 133 2020. Yale University Press. 256 pages, 28.00 USD, Cloth. The Chemical Age: How Chemists Fought Famine and Disease, Killed Millions, and Changed Our Relationship with the Earth. By Frank A. von Hip- pel. 2020. University of California Press. 368 pages, 29.00 USD, Cloth or E-book. Down by the Eno, Down by the Haw: A Wonder Almanac. By Thorpe Moeckel. 2019. Mercer Univer- sity Press. 136 pages, 16.00 USD, Paper. Environments of Empire: Networks and Agents of Ecological Change. Edited by Ulrike Kirchberger and Brett M. Bennett. 2020. 278 pages, 90.00 USD, Cloth, 29.95 USD, Paper, 22.99 USD, E-book. Great Lakes Rocks: 4 Billion Years of Geologic History in the Great Lakes Region. By Stephen E. Kesler. 2020. University of Michigan Press. 368 pages and 100 illustrations, 80.00 USD, Cloth, 29.95 USD, Paper. The Greater Gulf: Essays on the Environmental History of the Gulf of St Lawrence. Edited by Claire Elizabeth Campbell, Edward MacDonald, and Brian Payne. 2020. McGill-Queens University Press. 384 pages, 120.00 CAD, Cloth, 34.95 CAD, Paper. Kingdom of Frost: How the Cryosphere Shapes Life on Earth. By Bjorn Vassnes. Translated by Lucy Moffatt. 2020. Greystone Books. 240 pages, 32.95 CAD, Cloth. Nature and Value. Edited by Akeel Bilgrami. 2020. Columbia University Press. 312 pages, 105.00 USD, Cloth, 35.00 USD, Paper or E-book. Nature beyond Solitude: Notes from the Field. By John Seibert Farnsworth. Foreword by Thomas Lowe Fleischner. 2020. Comstock Publishing Associates. 216 pages, 19.95 USD, Paper. Nature’s Best Hope: A New Approach to Conser- vation that Starts in Your Yard. By Douglas W. Tallamy. 2020. Timber Press, Inc. 256 pages, 29.95 USD, Cloth, 14.99 USD, E-book. North America’s Galapagos: The Historic Chan- nel Islands Biological Survey. By Corinne Heyning Laverty. Foreword by Torben C. Rick. 2019. Univer- sity of Utah Press. 384 pages, 29.95 USD, Paper, 24.00 USD, E-book. Origins of Darwin’s Evolution. Solving the Species Puzzle Through Time and Place. By J. David Archi- bald. 2020. Columbia University Press. 208 pages, 30.00 USD, Paper. Cloth and E-book editions pub- lished in 2017. 2019 Our Wild Calling: How Connecting with Animals Can Transform Our Lives—and Save Theirs. By Richard Louv. 2019. Algonquin Books. 320 pages, 37.95 CAD, Cloth. Phylogenetic Ecology: A History, Critique, and Remodeling. By Nathan G. Swenson. 2020. Uni- versity of California Press. 240 pages, 120.00 USD, Cloth, 40.00 USD, Paper. Also available as an E-book. Plastic Waste and Recycling: Environmental Im- pact, Societal Issues, Prevention, and Solutions. Edited by Trevor M. Letcher. 2020. Academic Press. 686 pages, 100 colour illustrations, and 200 black and white illustrations, 250.00 USD, Paper or E-book. Primer of Ecological Restoration. By Karen D. NEw TITLES 397 Holl. 2020. Island Press. 224 pages, 35.00 USD, Paper or E-book. Reef Life: An Underwater Memoir. By Callum Roberts. 2020. Pegasus Books. 368 pages, 28.95 USD, Cloth, 18.99 USD, E-book. Restigouche: The Long Run of the Wild River. By Philip Lee. 2020. Goose Lane Editions. 288 pages, 22.95 CAD, Paper. Re-Bisoning the West: Restoring an American Icon to the Landscape. By Kurt Repanshek. 2019. Torrey House Press. 200 pages, 18.95 USD, Paper. The Salmon Way: An Alaska State of Mind. 2019. By Amy Gulick. Braided River. 192 pages, 29.95 USD, Cloth. The Canadian Field-Naturalist News and Comment Compiled by Amanda E. Martin Upcoming Meetings and Workshops This will be an unusual Upcoming Meetings and Workshops entry. That seems appropriate, given the unusual times in which we find ourselves living. At the time of writing, the novel coronavirus disease, COVID-19, has been detected in most countries, with 1 699 595 confirmed cases and 106 138 fatali- ties (World Health Organization 2020). In response to this threat, many countries have enacted measures to slow the spread of the virus and to avoid overwhelm- ing our healthcare systems. This includes measures to limit non-essential travel and in-person gatherings. This has led to some events being cancelled (e.g., American Society for Mammalogists annual meet- ing; https://mammalmeetings.org/) or delayed (e.g., Québec RE?’ Conference; http://www.re3-quebec 2020.org/). Others have quickly changed the for- mat of their event, from in-person to online (see list- ings below). As the COVID-19 situation continues to evolve, we expect that the situation for meeting and workshop organizers will too. Thus, we encourage readers to refer to the meeting/workshop webpages for the most up-to-date information. We wish our readers, their colleagues, friends, and family the best of health during these difficult times. Literature Cited World Health Organization. 2020. WHO COVID-19 dashboard. Accessed 13 April 2020. https://who. sprinklr.com/. Ontario Ecology, Ethology, and Evolution Colloquium The Ontario Ecology, Ethology, and Evolution Col- loquium to be held as an online meeting, 9 May 2020. More information is available at https://oe3c.com/. North American Regional Association of the International Association for Landscape Ecology Annual Meeting The annual meeting of the North American Regional Association of the International Association for Landscape Ecology to be held as an online meet- ing, 11-14 May 2020. Registration is currently open. More information is available at http://www.ialena. org/annual-meeting.html. Canadian Botanical Association/L’Association Botanique du Canada Annual Meeting The annual meeting of the Canadian Botanical Asso- ciation/L’Association Botanique du Canada to be held as an online meeting, 1—2 June 2020. More in- Botany 2020 Botany 2020 to be held 18-22 July 2020 at the Dena’ina Center, Anchorage, Alaska. Registration is formation is available at https://abc-cba2020.uqat.ca/ index.php. currently open. More information is available at http:// 2020.botanyconference.org/. Mycological Society of America Annual Meeting The annual meeting of the Mycological Society of | ference is: ‘Mycology in the Swamp’. Registration America to be held 19-22 July 2020 at the University of Florida, Gainesville, Florida. The theme of the con- is currently open. More information is available at https://msafungi.org/2020-annual-meeting/. North American Congress for Conservation Biology The 5th biennial North American Congress for Conservation Biology to be held 26-31 July 2020 at the Sheraton Downtown Hotel, Denver, Colorado. The theme of the conference is: ‘Cross- ing Boundaries: Innovative Approaches to Conser- vation’. Registration is currently open. More infor- mation is available at https://scbnorthamerica.org/ index. php/naccb-2020/. 398 Index to Volume 133 Compiled by William Halliday Achyranthes japonica, 56 Alaska, Adak, 49 Aleutian Islands, 49 Amchitka, 49 Attu, 49 Brooks Range, 151 Alberta, Central, 309 North-Central, 1 Northeastern, 189 Albinism, 113 Alces americanus, 329 Ambystoma laterale, 43 Ammodytes personatus, 144 Amphibians, 43, 101, 193, 196 Anderson, R.B., 1 Annual Reports of OFNC Committees for October 2017—September 2018, 90-95 Ant(s), 309 Araceae, 139 Archaeology, 332 Area(s), Protected, 43, 206, 313 Attaction, Conspecific, 235 Axanthism, 196 Balaenoptera acutorostrata, 144 Barbarea orthoceras, 118 stricta, 118 verna, 118 Barber-Meyer, S.M., 343 Bat, Hoary, 125 Batch, 325 Bay, Hudson Bear, Brown, 151 Beaver, American, 332 Behaviour, 235 Hunting, 16 Singing, 28 Swimming, 25 Bellan, L., 305 Biogeography, 156, 206 Bioindicators, 206 Biomass, Below Ground, 364 Bird(s), 20, 28, 167, 235, 301, 305, 352 Cavity-nesting, 352 Paridae, 28 Blaney, C.S., 118 Body, Fat, 34 Bondrup-Nielsen, S., 329 Boudreau, M.J., 329 Boyle, S.P., R. Dillon, J.D. Litzgus, D. Lesbarreres. Desiccation of herpetofauna on roadway exclu- sion fencing, 43-48 Brassicaceae, 118 Braun, C.E., W.P. Taylor, S.M. Ebbert, L.M. Spitler. Monitoring Rock Ptarmigan (Lagopus muta) populations in the Western Aleutian Islands, Alaska, 49-55 Breeding, 20, 301 Time of, 352 British Columbia, 156 Cormorant Island, 144 Haida Gwaii, 352 Hecate Strait, 263 Kamloops, 28 Kelowna, 28 Kootenays, 221 Williams Lake, 28 Brodo, I., 96 Brunton, D.F. A practical technique for preserving spe- cimens of duckmeal, Wolffia (Araceae), 139- 143 Bullfrog, American, 43, 196 California, Sierra Nevada Mountains, 34 Callospermophilus lateralis, 34 Cambaridae, 160 Cameron, M.D., 151 Campomizzi, A.J., Z.M. Lebrun-Southcott, K. Rich- ardson. Conspecific cues encourage Barn Swal- low (Hirundo rustica erythrogaster) prospect- ing, but not nesting, at new nesting structures, 235-245 Canis, 16 latrans var., 329 lupus, 60, 343 Cannings, S.G., T.S. Jung, J.-H. Skevington, I. Du- clos, S. Dar. A reconnaissance survey for Col- lared Pika (Ochotona collaris) in northern Yu- kon, 130-135 Carnivore, 16 Castor canadensis, 332 Catfish, Flathead, 372 Cavity-nesters, 352 399 400 Cavity-nesting, 352 Cedar, K., 160 Cetacean, 144, 263 Chaff-flower, Japanese, 56 Chapman, C.J.,C.S. Blaney, D.M. Mazerolle. Winter- cresses (Barbarea W.T. Aiton, Brassicaceae) of the Canadian Maritimes, 118-124 Chelydra serpentina, 216 Chickadee, Mountain, 28 Chrosomus neogaeus, 105 Chrysemys picta, 216 Chub, Creek, 325 Hornyhead, 325 Cipriani, J., 96 Clupea pallasi, 144 Cobb, T.P., 189 Colm, J.E., 372 Colouration, 301 Leucistic, 301 Communication, 28 Competition, Interspecific, 105 Condition, Body, 34 Conductivity, Total Body Electrical, 34 Conservation, 118, 206, 235 Coyotes, Eastern, 329 Crayfish, 160 Cruciferae, 118 Cucumaria frondosa, 113 Cues, Social, 235 Curley, R., D.L. Keenlyside, H.E. Kristmanson, R.L. Dibblee. A review of the historical and current status of American Beaver (Castor canadensis) on Prince Edward Island, Canada, 332-342 Cyprinidae, 325 Dace, Blacknose, 325 Finescale, 105 Northern Pearl, 325 Dar, S., 130 Decapoda, 160 Den, 1 Deer, Decoy, 16 White-tailed, 16, 246, 343 DeMaynadier, P.G., 196 Depredation, 60 Design, BACI Dibblee, R.L., 332 Diet, 151, 329 Dillon, R., 43 Dispersal, 136, 332 THE CANADIAN FIELD-NATURALIST Vol. 133 Distribution, 60, 130, 156, 160, 189, 199, 221 Diversity, Colour, 113 Dorval, H.R., R.T. McMullin. Lichens and allied fun- g1 of Sandbar Lake Provincial Park, Ontario, 206-215 Duck, 167 Duclos, I., 130 Dynamics, Population, 253 Ebbert, S.M., 49 Ecology, Road, 43, 101 Editors’ Report for Volume 132 (2018), 298-300 Effects, Ecosystem, 364 Electrofishing, 372 Emydoidea blandingii, 216 Erethizon dorsatum, 25 Excavators, 352 Expansion, Host, 309 Range, 309 Extension, Range, 56, 156, 160, 189, 199, 221, 309 Fat, 34 Feeding, 144 Fencing, Exclusion, 43 Festarini, A., 305 Fire, 313 Prescribed, 253 Fish, 105; 154.325, 372 Fishing, 151 Fitness, 301 Floristics, 118 Forbes, G.J., 270 Forest, Boreal, 1, 206 Great Lakes-St. Lawrence, 206 Formica podzolica, 309 Forsyth, R.G., 313 Forsyth, R.G., J. Kamstra. Roman Snail, Helix poma- tia (Mollusca: Helicidae), in Canada, 156-159 Frog(s), 43 American Bull, 43, 196 Gray Tree, 43 Green, 43, 196 Northern Leopard, 43, 193 Wood, 43 Fungi, 206 Gable, D.P., 16 Gable, T.D., D.P. Gable. Wolf (Canis sp.) attacks life- like deer decoy: insights into how wolves hunt deer? 16-19 Gartersnake, Common, 43 2019 Gaston, A.J., 352 Gaston, A.J. Birds of Mansel Island, northern Hud- son Bay, 20-24 Gaston, A.J., N.G. Pilgrim, V. Pattison. Humpback Whale (Megaptera novaeangliae) observa- tions in Laskeek Bay, western Hecate Strait, in spring and early summer, 1990-2018, 263-269 Gastropod(s), 156, 313 Freshwater, 189 Terrestrial, 221 Glasier, J.R.N., 309 Glass, 305 Glon, M.G., 160 Goldfish, 105 Gosselin, I., 305 Grantham, M., 313 Grieves, L.A., 301 Grit, 305 Ground, Feeding, 144 Ground Squirrel, Golden-mantled, 34 Grouse, Sharp-tailed, 253 Gulo gulo, | Habitat, 1, 144 Marine, 136, 144 Nesting, 235 Restoration, 253 Hamel, C.D., 313 Hamel, J.-F., 113 Hanrahan, C., 96 Harpalejeunea molleri subsp. integra, 199 Hart, J., 60 Haughian, S.R., T.H. Neily. Harpalejeunea molleri subsp. integra (R.M. Schuster) Damsholt new to Atlantic Canada, 199-205 Haughton, J., 304 Helix pomatia, 156 Hepatic, 199 Heron, J., 221 Herring, Pacific, 144 Hibernation, 34 Hinchliffe, R.P., C. Tebby, T.P. Cobb. First recorded co-occurrence of Valvata lewisi Currier, 1868 and Valvata lewisi ontariensis Baker, 1931 (Gastropoda: Valvatidae) from Alberta, Can- ada, with notes on morphometric and genetic variability, 189-192 Hirundo rustica erythrogaster, 235 History, 332 Holothuroidea, 113 Hotspot, 101 Hunter, M.L., Jr., 193 Hybognathus hankinsoni, 325 Hyla versicolor, 43 INDEX TO VOLUME 133 401 Illes, C., J.E. Colm, N.E. Mandrak, D.M. Marson. Flat- head Catfish (Pylodictis olivaris) reproduction in Canada, 372-380 Impact, Environmental, 305 iNaturalist, 160 Index, Condition, 34 Urbanization, 28 Information, Public, 235 Insectivore, Aerial, 235 Institute, Alberta Biodiversity Monitoring, 189 Introduction, Intentional, 105 Invasive, 56 Inventory, 221 Isbell, F., 60 Island(s), Adak, 49 Aleutian Islands, 49 Amchitka, 49 Attu, 49 Campobello, 136 Cormorant, 144 East Sister, 56 Erie, 56 Mansel, 20 Middle, 56 Jokinen, M.E., S.M. Webb, D.L. Manzer, R.B. Ander- son. Characteristics of Wolverine (Gulo gulo) dens in the lowland boreal forest of north-cen- tral Alberta, 1-15 Joly, K., 151 Jones, C.D., M.G. Glon, K. Cedar, S.M. Paiero, P.D. Pratt, T.J. Preney. First record of Painted Mud- bug (Lacunicambarus polychromatus) in Onta- rio and Canada and the significance of iNatu- ralist in making new discoveries, 160—166 Jung, T.S., 130 Jung, T.S. Behaviour of a porcupine (Erethizon dor- satum) swimming across a small boreal stream, 25-27 Juvenile, 372 Kamstra, J., 156 Kamstra, J. Japanese Chaff-flower, Achyranthes ja- ponica (Amaranthaceae), on the Erie islands, an invasive plant new to Canada, 56—59 Keefe, D.G., R.C. Perry, G.R. McCracken. First re- cords of Finescale Dace (Chrosomus neogaeus) in Newfoundland and Labrador, Canada, 105— 112 Keenlyside, D.L., 332 Kelt, D.A., 34 4Q2 Kinley, T.A. Seasonal movements of White-tailed Deer (Odocoileus virginianus) in the Rocky Mountains of British Columbia, 246—252 Kreuzberg, E., 101 Kristmanson, H.E., 332 Krueger, J., 60 Lacunicambarus polychromatus, 160 Lagopus muta, 49 Lake(s), Erie, 56, 364 Great, 372 St. Clair, 372 Williams, 28 Lasiurus cinereus, 125 LaZerte, S.E., K.L.D. Marini, H. Slabbekoorn, M.W. Reudink, K.A. Otter. More Mountain Chicka- dees (Poecile gambeli) sing atypical songs in urban than in rural areas, 28-33 Lebrun-Southcott, Z.M., 235 Eee," 12 305 Lei, C., S.J. Yuckin, R.C. Rooney. Rooting depth and below ground biomass in a freshwater coastal marsh invaded by European Reed (Phragmites australis) compared with remnant uninvaded sites at Long Point, Ontario, 364-371 Lejeuneaceae, 199 Lemnoideae, 139 Lepage, D. Minutes of the 140" Annual Business Meet- ing (ABM) of the Ottawa Field-Naturalists’ Club, 8 January 2019, 88-89 Lepitzki, D., A. Martin. Editors’ Report for Volume 132 (2018), 298-300 Lesbarreéres, D., 43 Leucism, 301 Leucistic, 301 Lichens, 206 Lindemann, S.B., A.M. O’Brien, T.B. Persons, P.G. DeMaynadier. Axanthism in Green Frogs (Lithobates clamitans) and an American Bull- frog (Lithobates catesbeianus) in Maine, 196— 198 Lindemann, S.B., D.E. Putnam, M.L. Hunter, Jr., T.B. Persons. Spotless bursni pattern in Northern Leopard Frog (Lithobates pipiens) in Maine, 193-195 Lithobates catesbeianus, 43, 196 clamitans, 43, 196 pipiens, 43, 193 sylvaticus, 43 Litter, Anthropogenic, 305 Litzgus, J.D., 43 THE CANADIAN FIELD-NATURALIST Vol. 133 Liverwort, 199 Livestock, 60 Lowlands, | Luxilus cornatus, 325 Maine, 193, 196 Mallard, 167 Mammals, 1, 16, 25, 34, 60, 125, 144, 151, 246, 263, 329, 332, 343 Management, Prairie, 253 Mandrak, N.E., 372 Manitoba, Southeastern, 313 Manzer, D.L., 1 Margariscus nachtriebi, 325 Marini, K.L.D., 28 Maritimes, 118 Marsh, Coastal, 364 Marson, D.M., 372 Martin, A., 298 Mass, Body, 34 Mazerolle, D.M., 118 McAlpine, D.F. Occurrence of the rare marine littoral millipede, Thalassisobates littoralis (Diplopo- da: Nematosomatidae), in Canada, 136-138 McAlpine, D.F., G.J. Forbes. A Tribute to Rudolph Franck Stocek, 1937-2018, 270-275 McCracken, G.R., 105 McCurdy-Adams, H., 216 McLachlan Hamilton, K., 96 McMillan, C.J., 144 McMullin, R.T., 206 Mech, L.D., S.M. Barber-Meyer. Sixty years of White- tailed Deer (Odocoileus virginianus) yarding in a Gray Wolf (Canis /upus)—deer system, 343— 351 Mech, L.D., F. Isbell, J. Krueger, J. Hart. Gray Wolf (Canis Iupus) recolonization failure: a Minne- sota case study, 60-65 Megaptera novaeangliae, 263 Mercier, A., 113 Metal, 305 Migration, 343 Millipede, Marine, 136 Minnesota, 60 Northeastern, 343 Minnow(s), 325 Brassy, 325 Minutes of the 140" Annual Business Meeting (ABM) of the Ottawa Field-Naturalists’ Club, 8 Janu- ary 2019, 88-89 Mitigation, Road-effect, 43, 216 Mollusca, 156, 189, 221, 313 Monitoring, Population, 49 2019 Montgomery, E.M., T. Small, J.-F. Hamel, A. Mer- cier. Albinism in Orange-footed Sea Cucum- ber (Cucumaria frondosa) in Newfoundland, 113-117 Moore, K., 352 Moose, 329 Morningstar, D., A. Sandilands. Summer movements of a radio-tagged Hoary Bat (Lasiurus cinere- us) captured in southwestern Ontario, 125—129 Mortality, 43, 167 Motus, 125 Mountains, 151 Nahoni, 130 Richardson, 130 Sierra Nevada, 34 Movement, Seasonal, 125, 246, 263 Summer, 125 Mudbug, Paintedhand, 160 Muntz, E.M., 329 Murphy, R.K., K.A. Smith. Sharp-tailed Grouse (7Zym- panuchus phasianellus) population dynamics and restoration of fire-dependent northern mixed-grass prairie, 253-262 Neily, T.H., 199 Nesting, 235, 305 Cavity, 352 Nestling, 305 New Brunswick, Campobello Island, 136 Newfoundland and Labrador, 113 Exploits River, 105 Newt, Eastern, 101 Nicolai, A., R.G. Forsyth, M. Grantham, C.D. Hamel. Tall grass prairie ecosystem management—a gastropod perspective, 313-324 Nocomis biguttatus, 325 North Atlantic, 113 North Dakota, 253 Notophthalmus viridescens, 101 Nova Scotia, 199 Cape Breton, 329 Nunavut, Mansel Island, 20 O’Brien, A.M., 196 Occurrence, Seasonal, 263 Oceanography, 263 Ochotona collaris, 130 Odocoileus virginianus, 16, 246, 343 Oncorhynchus spp., 151 Ontario, 156 Eastern, 216, 305 INDEX TO VOLUME 133 403 Killarney, 16 Lake Erie, 56 Long Point, 364 Northwestern, 206 Southern, 43, 235, 301, 325 Southwestern, 125, 160, 364, 372 Ornithology, 20, 28, 167, 235, 301, 305, 352 Otter, K.A., 28 Ovaska, K., L. Sopuck, J. Heron. Survey for terrestri- al gastropods in the Kootenay region of British Columbia, with new records and range exten- sions, 221-234 Paiero, S.M., 160 Paper, 305 Parasitism, Dulotic, 309 Park, Cape Breton Highlands National, 329 Daadzaii Van Territorial, 130 Gates of the Arctic National Park and Preserve, 151 Gatineau, 101 Herring Cove Provincial, 136 Jarvis Bay Provincial, 309 Killarney Provincial, 16 Kootenay National, 246 Nv unlit Njik (Fishing Branch) Territorial, 130 Presqu’ile Provincial, 43 Sandbar Lake Provincial, 206 Pattison, V., 263 PettyeR.C. 105 Persons, T.B., 193, 196 Phragmites australis, 364 Piercey, R.S., 144 Pika, Collared, 130 Pilgrim, N.G., 263 Pilgrim, N.G., J.L. Smith, K. Moore, A.J. Gaston. Nest site characteristics of cavity-nesting birds on a small island, in Haida Gwaii, British Colum- bia, Canada, 352-363 Plant(s), 56, 118, 139, 199 Plastic, 305 Poecile gambeli, 28 Polyergus bicolor, 309 Population, 49, 253, 263 Porcupine, North American, 25 Power, J.W.B., M.J. Boudreau, E.M. Muntz, S. Bon- drup-Nielsen. High reliance on a diet of Moose (Alces americanus) by Eastern Coyotes (Canis latrans var.) in Cape Breton Highlands Nation- al Park, Nova Scotia, Canada, 329-331 404 Prairie, Northern Mixed-grass, 253 Tall Grass, 313 Pratt, P.D., 160 Predation, 16, 343 Predator-prey, 16 Preney, T.J., 160 Preparation, Herbarium Specimen, 139 Prince Edward Island, 332 Prospecting, 235 Ptarmigan, Rock, 49 Putnam, D.E., 193 Pylodictis olivaris, 372 Quebec, Gatineau, 101 Quinn, N.W.S. Batch spawning in five species of min- nows (Cyprinidae) from Ontario, Canada, 325— 328 Range, Distributional, 130 Summer, 246 Winter, 246 Recolonization, 60 Record, New, 118 New Distribution, 160, 221 New Provincial, 156, 160 Reed, European, 364 Reintroduction, 60 Relations, Predator-prey, 343 Reproduction, 372 Reptiles, 43, 216 Residuals, Mass-length, 34 Restoration, Habitat, 235 Reudink, M.W., 28 Review, 332 Rhinichthys artatulus, 325 Rhizomes, 364 Richardson, K., 235 River, Klondike, 25 Thames, 372 Robin, American, 301 Rood, S.B., A. Willcocks. Duckling mortality at a riv- er weir, 167-171 Rooney, R.C., 364 Roots, 364 Salamander, 101 Blue-spotted, 43 Salmon, 151 Sand Lance, Pacific, 144 THE CANADIAN FIELD-NATURALIST Vol. 133 Sandilands, A., 125 Scat, 329 Sea Cucumber, Orange-footed, 113 Seburn, D.C., E. Kreuzberg, L. Viau. Roadkill of East- ern Newts (Notophthalmus viridescens) in a protected area in Quebec, 101-104 Seburn, D.C., H. McCurdy-Adams. Do turtle warning signs reduce roadkill? 216—220 Selection, Sexual, 301 Semotilus atromaculatus, 325 Shiner, Common, 325 Signs, Turtle Warning, 216 Wildlife, 216 Site, Nest, 352 Skevington, J.H., 130 Slabbekoorn, H., 28 Slugs, Terrestrial, 313 Small, T., 113 Smith, J.L., 352 Smith, K.A., 253 Snail(s), Freshwater, 189, 313 Roman, 156 Terrestrial, 156, 221, 313 Snakes, 43 Common Garter, 43 Snow, 1 Songs, Atypical, 28 Sopuck, L., 221 Sorum, M.S., K. Joly, M.D. Cameron. Use of salmon (Oncorhynchus spp.) by Brown Bears (Ursus arctos) in an Arctic, interior, montane environ- ment, 151-155 Sosiak, C.E., M. West, J.R.N. Glasier. First record and new host record of the obligate dulotic ant, Polyergus bicolor (Hymenoptera: Formicidae), in Alberta, Canada, 309-312 Spawning, Batch, 325 Species, Anthropochorus, 136 Exotic, 105 Invasive, 56, 364 Rare, 313 Spitler, L.M., 49 Squirrel, Golden-mantled Ground, 34 Ground, 34 Status, Population, 49, 332 Storage, 139 Strait, Hecate, 263 Structures, Nesting, 235 Stuart, M., 305 2019 Survey, Acoustic, 28 Aerial, 130 Camera Trap, | Electrofishing, 372 Fyke Net, 105 Gill Net, 105 Hoop Net, 372 Live Trap, | Minnow Pot, 105, 325 Passive Acoustic, 28 Point Count, 43 Road, 101 Scat, 329 Trammel Net, 372 Transect, 329, 352 Visual, 43, 101, 130, 144, 151, 206, 263, 329, 352 Swallow, Barn, 235 Tree, 305 Tachycineta bicolor, 305 Taylor, W.P., 49 Tebby, C., 189 Telemetry, GPS, 1 Radio, 125, 343 Thalassisobates littoralis, 136 Thamnophis sirtalis, 43 Thorn, R.G., 301 Towers, J.R., C.J. McMillan, R.S. Piercey. Sighting rates and prey of Minke Whales (Balaenoptera acutorostrata) and other cetaceans off Cormo- rant Island, British Columbia, 144—150 Treefrog, Gray, 43 Trees, Wildlife, 352 Trends, Population, 263 Tribute, 270 Turdus migratorius, 301 Turtle, 216 Blanding’s, 216 Painted, 216 Snapping, 216 Tympanuchus phasianellus, 253 Urbanization, 28 Ursus arctos, 151 Use, Habitat, 144 INDEX TO VOLUME 133 405 Valvata, 189 lewisi, 189 lewisi ontariensis, 189 Valvatidae, 189 Van Vuren, D.H., 34 Variant, Blue Colour, 196 Pattern, 193 Viau, L., 101 Vocalization, 28 Walsh, S., J. Haughton, L. Bellan, I. Gosselin, A. Festa- rini, D. Lee, M. Stuart. Occurrence of anthropo- genic litter in nestling Tree Swallows (Zachy- cineta bicolor), 305-308 Waterfowl, 167 Watson, J.A., 34 Webb, S.M., 1 Weir, River, 167 Wells, C.P., J.A. Watson, D.A. Kelt, D.H. Van Vuren. Body mass as an estimate of female body con- dition in a hibernating small mammal, 34—42 West, M., 309 Wetlands, 364 Whale, Humpback, 263 Minke, 144 Willcocks, A., 167 Wintercresses, 118 Wolf, 16 Gray, 60, 343 Wolffia, 139 Wolverine, 1 Yard, Deer, 343 Yarding, 343 Young-of-year, 372 Yukin, S.J., 364 Yukon, Klondike River, 25 Zitani, N.M., L.A. Grieves, R.G. Thorn. A successful- ly breeding, partially leucistic American Robin (Turdus migratorius), 301-304 Zurbrigg, E., I. Brodo, J. Cipriani, C. Hanrahan, K. McLachlan Hamilton. The Ottawa Field-Nat- uralists’ Club Awards for 2018, presented Feb- ruary 2019, 96-99 406 Index to Book Review Botany Brunton, D.F. “Flora of Florida Volume 6 (Dicoty- ledons, Convolvulaceae through Paulownia- ceae)” by R.P. Wunderlin, B.F. Hansen, and A.R. Franck, 2019, 70 Brunton, D.F., M.J. Oldham. “Michigan Ferns & Ly- cophytes: A Guide to Species of the Great Lakes Region” by D.D. Palmer, 2018, 68-69 Clarkin, O. “Identification of Trees and Shrubs in Winter Using Buds and Twigs” by Bernd Schulz, 2018, 71-72 Cray, H.A. “Seaweed Chronicles” by Susan Hand Shetterly, 2018, 276 Crins, W.J. “Sedges of the Northern Forest: A Pho- tographic Guide” by Jerry Jenkins, 2019, 172-173 Climate Change Brooks, R. “The Uninhabitable Earth: Life After Warming” by David Wallace-Wells, 2019, 66— 68 Entomology Cottam, B. “Butterflies: Their Natural History and Diversity. Second Edition” by Ronald Oren- stein, Photography by Thomas Marent, 2020, 383 Cray, H.A. “Buzz, Sting, Bite: Why We Need Insects” by Anne Sverdrup-Thygeson, 2019, 381-382 Lauff, R. “Field Guide to Flower Flies of Northeast- ern North America” by Jeffrey H. Skevington, Michelle M. Locke, Andrew D. Young, Kevin Moran, William J. Crins, and Stephen A. Mar- shall, 2019, 73 Herpetology Seburn, D. “The Field Herping Guide: Finding Am- phibians and Reptiles in the Wild” by Mike Pingleton and Joshua Holbrook, 2019, 384 Ornithology Clark, H.O., Jr. “Feed the Birds: Attract and Identi- fy 196 Common North American Birds” by Chris Earley, 2019, 279-280 Curry, B. “The Handbook of Bird Families” by Jona- than Elphick, 2018, 74-75 Foster, R.F. “Ospreys: The Revival of a Global Rap- tor” by Alan F. Poole, 2019, 175 Gaston, T. “Gulls” by John C. Coulson, 2019, 174 THE CANADIAN FIELD-NATURALIST Vol. 133 Gaston, T. “Gulls of the World: A Photographic Guide” by Klaus Malling Olsen, 2018, 385 Lein, M.R. “Birds of Saskatchewan” by Alan R. Smith, C. Stuart Houston, and J. Frank Roy, 2019, 277-278 Other Bocking, E. “Surviving Global Warming: Why Elimi- nating Greenhouse Gases Isn’t Enough” by Roger A. Sedjo, 2019, 390-391 Burke, T. “How to Give Up Plastic: A Guide to Changing the World, One Plastic Bottle at a Time” by Will McCallum, 2018, 283 Burke, T. “Plastic Soup: An Atlas of Ocean Pollution” by Michiel Roscam Abbing, 2019, 285 Cottam, B. New Titles, 80-83, 184-186, 289-293, 394-397 Cottam, B. “To Speak for the Trees: My Life’s Jour- ney from Ancient Celtic Wisdom to a Healing Vision of the Forest” by Diana Beresford- Kroeger, 2019, 181-182 Cottam, B. “The Overstory: A Novel” by Richard Powers, 2018, 183 Cottam, B. “Mama’s Last Hug: Animal and Human Emotions” by Frans de Waal, 2019, 286-287 Gaston, T. “North Pole: Nature and Culture” by Mi- chael Bravo, 2019, 288 Hartzell, S.M. “The Environment: A History of the Idea” by Paul Warde, Libby Robin, and Sver- ker Sorlin, 2018, 76 Hunter, M.L., Jr. “The Great Himalayan National Park: The Struggle to Save the Western Hi- malayas” by Sanjeeva Pandey and Anthony J. Gaston, 2019, 77—78 Pazhoohi, F. “How to Walk on Water and Climb up Walls: Animal Movement and the Robots of the Future” by David L. Hu, 2018, 392 Sander-Regier, R. “Frog Pond Philosophy: Essays on the Relationship Between Humans and Na- ture” by Strachan Donnelley, 2018, 393 Zoology Clark, H.O., Jr. “The Flying Zoo: Birds, Parasites, and the World They Share” by Michael Stock, 2019, 386-387 Cottam, B. “Mammal Tracks and Sign: A Guide to North American Species. Second Edition” by Mark Elbroch with contributions by Casey McFarland, 2019, 178-179 Cray, H.A. “Bats: An Illustrated Guide to All Spe- 2019 cies” by Marianne Taylor, Photography by Merlin D. Tuttle, 2018, 177 Gibson, J.F. “The New Beachcomber’s Guide to the Pacific Northwest” by J. Duane, 2019, 176 Gibson, J.F. “A Field Guide to Marine Life of the Pro- tected Waters of the Salish Sea” by Rick M. Harbo, 2019, 176 Gibson, J.F. “A Field Guide to Marine Life of the Outer Coasts of the Salish Sea and Beyond” by Rick M. Harbo, 2019, 176 Halliday, W.D. “The North Atlantic Right Whale: Disappearing Giants. Revised and Updated Edition” by Scott Kraus, Marilyn Marx, Heather Pettis, Amy Knowlton, and Kenneth Mallory, 2019, 281 INDEX TO VOLUME 133 407 Halliday, W.D. “Orca: The Whale Called Killer. Fifth Edition” by Erich Hoyt, 2019, 388 Lauff, R. “Mammals of Prince Edward Island and Adjacent Marine Waters” by Rosemary Cur- ley, Donald F. McAlpine, Dan McAskill, Kim Riehl, and Pierre-Yves Daoust, 2019, 389 Way, J. “Return of the Wolf: Conflict and Coexis- tence” by Paula Wild, 2018, 78-79 Way, J. “The Rise of Wolf 8: Witnessing the Triumph of Yellowstone’s Underdog” by Rick Mc- Intyre, 2019, 180-181 Way, J. “Yellowstone Cougars: Ecology Before and During Wolf Restoration” by Toni K. Ruth, Polly C. Buotte, and Maurice G., 2019, 282— 283 THE CANADIAN FIELD-NATURALIST Volume 133, Number 4 A successfully breeding, partially leucistic American Robin (7urdus migratorius) Nina M. ZITANI, LEANNE A. GRIEVES, and R. GREG THORN Occurrence of anthropogenic litter in nestling Tree Swallows (Jachycineta bicolor) STEPHANIE WALSH, JENNIFER HAUGHTON, LEE BELLAN, ISABELLE GOSSELIN, AMY FESTARINI, DAvID LEE, and MARILYNE STUART First record and new host record of the obligate dulotic ant, Polyergus bicolor (Hymenoptera: For- micidae), in Alberta, Canada CHRISTINE E. SOSIAK, MARI WEST, and JAMES R.N. GLASIER Tall grass prairie ecosystem management—a gastropod perspective ANNEGRET NICOLAI, ROBERT G. FORSYTH, MELISSA GRANTHAM, and Cary D. HAMEL Batch spawning in five species of minnows (Cyprinidae) from Ontario, Canada NORMAN W.S. QUINN High reliance on a diet of Moose (Alces americanus) by Eastern Coyotes (Canis latrans var.) in Cape Breton Highlands National Park, Nova Scotia, Canada JASON W.B. Power, MICHAEL J. BOUDREAU, ERICH M. MuntTz, and SOREN BONDRUP-NIELSEN A review of the historical and current status of American Beaver (Castor canadensis) on Prince Edward Island, Canada ROSEMARY CURLEY, DAvID L. KEENLYSIDE, HELEN E. KRISTMANSON, and RANDALL L. DIBBLEE Sixty years of White-tailed Deer (Odocoileus virginianus) yarding in a Gray Wolf (Canis lupus)— deer system L. Davip MEcH and SHANNON M. BARBER-MEYER Nest site characteristics of cavity-nesting birds on a small island, in Haida Gwaii, British Colum- bia, Canada NEIL G. PILGRIM, JOANNA L. SmiTH, KEITH Moore, and ANTHONY J. GASTON Rooting depth and below ground biomass in a freshwater coastal marsh invaded by European Reed (Phragmites australis) compared with remnant uninvaded sites at Long Point, Ontario CALVIN LEI, SARAH J. YUCKIN, and REBECCA C. ROONEY Flathead Catfish (Pylodictis olivaris) reproduction in Canada COLIN ILLEs, JULIA E. Com, NICHOLAS E. MANDRAK, and Davip M. MARSON 2019 301 305 309 313 52D 329 332 343 392 364 372 (continued inside back cover) ISSN 0008-3550 TABLE OF CONTENTS (concluded) Volume 133, Number 4 Book Reviews ENTOMOLOGY: Buzz, Sting, Bite: Why We Need Insects—Butterflies: Their Natural History and Diversity. Second Edition HERPETOLOGY: The Field Herping Guide: Finding Amphibians and Reptiles in the Wild ORNITHOLOGY: Gulls of the World: A Photographic Guide ZooLoGy: The Flying Zoo: Birds, Parasites, and the World They Share—Orca: The Whale Called Killer. Fifth Edition—Mammals of Prince Edward Island and Adjacent Marine Waters OTHER: Surviving Global Warming: Why Eliminating Greenhouse Gases Isn’t Enough—How to Walk on Water and Climb up Walls: Animal Movement and the Robots of the Future—Frog Pond Philosophy: Essays on the Relationship Between Humans and Nature NEw TITLES News and Comment Upcoming Meetings and Workshops Ontario Ecology, Ethology, and Evolution Colloquium—North American Regional Association of the International Association for Landscape Ecology Annual Meeting—Canadian Botanical Association/ L’Association Botanique du Canada Annual Meeting—Botany 2020—Mycological Society of America Annual Meeting—North American Congress for Conservation Biology Index Mailing date of the previous issue 133(3): 24 March 2020 2019 381 384 385 386 390 394 398 399