CONTRIBUTIONS
S IDA to Botany
VOLUME 14 NUMBER 1 JUNE 1990
CONTENTS
A new species of Viola (Violaceae) from the Guadalupe Mountains, Trans-Pecos Texas. A. Michael Powell and Brent Wauer.
An explanation for the discrepancy in the chromosome count of the redbud (Cercts canadensis, Leguminosae).
Will H. Blackwell.
Crataegus secreta (Rosaceae), a new species of hawthorn from the Edwards Plateau, Texas. J. B. Phipps.
The alpine-subalpine flora of northeastern México. J. Andrew McDonald.
Identification of the plants illustrated and described in Catesby’s Natural History of the Carolinas, Florida and the Bahamas. Robert L. Wilbur.
The Clematis virginiana (Ranunculaceae) complex in the southeastern United States. Frederick B. Essig.
Cyperus entrerianus (Cyperaceae), an overlooked species in temperate North America. Richard Carter.
Extension of native range of Sabal mexicana (Palmae) in Texas to include Central Coast. Landon Lockett and Robert W. Read.
Synopsis of Carex section Lupulinae (Cyperaceae) in
A,
Texas. Stanley D. Jones and Stephan L. Hate
(Continued on back cover)
US ISSN 0036-1488
Sipa, CONTRIBUTIONS TO BOTANY Founded by Lloyd H. Shinners, 1962
Publisher Wm. E Mahler BRIT/SMU Herbarium Dallas, Texas, 75275
Editor Associate Editor Barney L. Lipscomb John W. Thieret BRIT/SMU Herbarium Northern Kentucky University Dallas, Texas, 75275 Highland Heights, Kentucky, 41076
Guidelines for contributors are available upon request.
Subscription: $15.00/$22.50 (U.S.) per year; numbers issued twice a year.
© Sida, Contributions to Botany, Volume 14, Number 1, pages | — 140. Copyright 1990 by Wm. E Mahler
A NEW SPECIES OF VIOLA (VIOLACEAE) FROM THE GUADALUPE MOUNTAINS, TRANS-PECOS TEXAS
A. MICHAEL POWELL
bce y Biology, Sul Ross State University Alpine, TX 79832, U.S.A.
BRENT WAUER
ies ene National Park, HC 6 a Box 400 lt Flat, TX 79847-9400, U.S
ABSTRACT e only yellow-flowered Viola known to exist in the Guadalupe Mountains, Texas, is described as V. guadalupensis. The rock-dwelling new species is known from a single, small limestone formation on the East Rim of the mountains, and is an immediate candi- date for a status. The new species is related to V. vallicola and V. nuattallii of northern New México and western United States, and is also similar to V. painteri of northern México.
RESUMEN La unica Viola con flores amarillas en las montafas Guadalupes, Texas, se describe como V. guadalupensis. La especie, que vive (existe) entre las piedras, se reconoce de una sola, ee formaci6n de piedra caliza en la Orilla al Este de las montanas, y es candidata mediate para estado 0 posicidn de peligro. La nueve especie esta relacionada a V. vallicola y Vv nuttallii del norte de nuevo México es el oeste de los Estados Unidos, y es también semejante a V. painter del norte de Méxic
During the course of photographic studies of plants in Guadalupe Mountains National Park, Ranger Brent Wauer discovered an undescribed yellow-flowered violet growing in one small rock formation along the East Rim of the Guadalupe Mountains. Photographic, ecologic, and other data in addition to those presented below for the new species of Viola are includ- ed in the extensive photographic collection of plants housed in Guadalupe Mountains National Park headquarters at Pine Springs.
Viota guadalupensis A.M. Powell and B. Wauer, sp.nov. Fig. 1.
Plantae perennes quasi glabrae usque = 10 cm altae. Laminae foliorum ovatae vel ovati- lanceolatae, 1.2 —2.4 cm longae, 0.7 — 1.2 cm latae, glabrae vel trichomatibus paucis secus venas paginarum infernorum, margines integrae vel parce crenalae in dimidio ade Corolla flava, petala 7 — 10 mm longae, petalum infernum venis prominentibus brunneis; styli ca 1.5
mm longi capitati. Fructus stramineus 3.0—-4.5 mm oe Semina ee ta 2 mm ae
Sipa 14(1):1—6. 1990.
Perennial in small openings of /:mestone rock face, plants to 10 cm tall. Stems glabrous, 1 — 4 cm long (those parts collected from rock openings). Leaves caulescent; petioles glabrous, 2—6 cm long; stipules 3-11 mm long, 0.5 — 1.8 mm wide, lanceolate to ovate- or oblong-lanceolate, or linear, whitish to greenish and thin, sparingly glandular-fimbriate; blades ovate to triangular-ovate or ovate-lanceolate, 1.2 —2.4 cm long, 0.7 —1.3 em wide, glabrous or with a few short hairs especially along veins underneath, the margins entire or with 1—3 crenations on lower half, apex acute to rounded, the tip rather obscurely callused, the base broadly cuneate to roun- ded or truncate. Flowers borne among or above the upper leaves, pedicels- peduncles 3.5 —6 cm long, glabrous, slender; sepals glabrous, linear to linear-lanceolate, 3.5 —5 mm long, margins scarious, the base truncate or rounded to subauriculate; corollas yellow, fading reddish brown, some of the petals smeared reddish-brown outside, inconspicuously brown-veined (middle veins) near base and inside on lateral petals, prominently brown- veined (middle and diverging veins) on lower (spured) petal, the petals 7 — 10 mm long, the two lateral petals bearded inside; spur | — 1.3 mm long; anthers 1.5 — 2 mm long; terminal appendages 1 — 1.4 mm long; nectari- ferous spurs ca 1 mm long; styles ca 1.5 mm long, capitate, with short hairs on 2 margins. Fruit greenish, maturing tan, glabrous, 3—4.5 mm long; seeds ovoid, ca 2 mm long, light brown, with a well developed caruncle, off-white to tanish in color, extending back along the seed from nearly one- half to almost the entire length of the seed.
Tyee: TEXAS. CuLBeRsON Co.: Guadalupe Mountains of Guadalupe Mountains Na- tional Park, N side of prominent rock face (no map name), along E Rim, 1.48 km N, 1.0 km E of the summit of Hunter Peak; ca 35 scattered plants on a 7 X 10 m rock face sheltered by vegetation; elev. 2600 m (8000 ft); collected by B. Wauer who found plants growing in “bullet-hole” openings in rock faces where roots could not be collected without damage; 12 May 1988, A.M. Powell and B. Wauer 5497 (HOLOTYPE: SRSC; tsorype: TEX).
Known only from the type collection.
Viola guadalupensis 1s named after the only mountain range in which it is known to occur. In fact, the plants are known only from one northwest- facing dolomitized limestone outcrop (with small ledges), shaded by Pseudotsuga menziesii (Douglas Fir), on the East Rim of the Guadalupe Mountains. Associated plant species on the rock outcrop include Petro- phytum caespitosum, Valeriana texana, Pinaropappus parvus, Chaetopappa hersheyt, Stipa lobata, and Carex sp., and at the base of the rock outcrop Fendlerella utahaensis, Ptelea trifoliata, Cercocarpus montanus, var. argenteus, Fendlera rupicola, Amelanchier utahensis, Physocarpus monogynus, and Cheilan- thes sp. The second author has carefully examined other seemingly suitable habitats in the Guadalupe Mountains including those along about 8 km (5
FIG. 1. Habit of Viola guadalupensis, Guadalupe Mountains, Texas.
4
mi) of the East Rim from the top of Bear Canyon to Lamar Canyon without locating any additional plants. The entire known population of V. gaada- lupensis comprises about 35 individuals in the one site.
Two other species of Vzola are known to occur in the Guadalupe Mountains, V. /ovelliana Brainerd and V. muissouriensis Greene, both blue- flowered taxa (Correll and Johnston 1970; Russell 1965; Burgess and Northington 1981). Viola guadalupensis is the only known yellow-flowered violet in the Guadalupe Mountains, and it appears to be related to V. valli- cola A. Nels. and V. nuttallii Pursh, yellow-flowered species of the western United States with closest distribution in southern Colorado or northern New Mexico (Martin and Hutchins 1984; Fabijan et al. 1987). Viola guadalupensis differs from V. vallicola and V. nuttallii by its rock-dwelling habit, leaf characters (especially the blade shape and pubescence), fruit size, and seed morphology (Table 1). Viola guadalupensis also exhibits generally smaller vegetative and floral features than V. vallicola and V. nuttalli:z. Morphological traits suggest closest relationship with the diploid V. vallicola. A chromosome count of V. guadalupensis will be very helpful in clarifying its relationship in the V. nwttallii complex (Fabijan et al. 1987, Baker 1957).
Viola guadalupensis is also similar to the yellow-flowered V. painteri Rose & House, a species of pine-fir woodlands in the Sierra Maderas de] Carmen in Coahuila, south to Oaxaca, in Mexico (Rose and House 1905, Henrick- son, pers. comm.). Viola paintert and V. barroetana Schaffner may be the only yellow-flowered violets in northern Mexico (Nesom, pers. comm., Baker 1957). Viola guadalupensis is delimited from V. painteri by its rock- dwelling habit, glabrous herbage, ovate to ovate-lanceolate leaf blades that are smaller and narrower with margins entire or sparingly crenate on the lower half, apexes acute or rounded, broadly cuneate to rounded or truncate leaf bases, shorter sepals and petals, and smaller fruits and seeds. Viola painter has herbage glabrous to pubescent, leaf blades cordate to reniform, 1 — 3(-5) cm long, 1 — 2(-4) cm wide, apexes acute, bases cordate, margins evenly crenate-serrate, fruit 7-9 mm long, and seeds ca 2.5 mm long. Baker (1957) suggests that V. va/licola may have arisen from V. barroetana although Fabijan et al. (1987) do not discuss this possibility, and we have not compared V. guadalupensis with V. barroetana.
ACKNOWLEDGMENTS
We are grateful to Guy Nesom who provided the Latin translation, in- formation about the general distribution of yellow-flowered violas in Mexico, and corrections to the manuscript, and to Jim Henrickson who made available the treatment of Vsola for the Chihuahuan Desert Flora.
TaBLe 1. Some distinguishing features of Viola guadalupensis and related species.
V. GUADALUPENSIS V. VALLICOLA V. NUTTALLII Substrate Rock crevices Soils Soils Leaf blades Ovate to triangular- Ovate to oblong- Mostly lanceolate, ovate or oblong- ovate, 1.9—7 cm long, 2.5—7.5 cm long, lanceolate, 1.2 — 2.4 0.9—3 cm wide, the 0.6—3.2 cm wide, , 0.7—1.3 cm __ base truncate to the base wide, the base subcordate, sparsely attenuate, broadly cuneate to to densely puberulent surfaces glabrous rounded or rarely throughout or to rather sparsely truncate, glabrous glabrous, but often puberulent throughout or with puberlent along veins especially along a few short hairs and margins if the veins along veins glabrous on the underne underneath, margins surfaces, margins seen Paine, entire or 1 — 3- usually ciliate, entire to crenate on lower entire co ea mostly crenulate on lower half Fruit 3—4.5 mm long, (5)6—8 mm long, 7—11 mm long, glabrous glabrous to densely glabrous to puberulent puberulent Seeds ca 2 mm long —. 2.7 mm long, 2.8—3 mm long, 1.2—1.4 mm wide, .2—1.5 mm wide, 1.5-—1.8 mm wide, ovoid with an off- os with a ates ovoid, with a white to tanish caruncle ca 0.8 mm jie Soh caruncle well ace and distally ca 1 mm long and developed (not flattened fae Fee
Chromosome Unknown 2n = 12 2n = 24 Number
The Spanish translation of the abstract was kindly provided by Dr. Abelardo Baeza, Professor of Languages and Literature and Director of Minority Affairs at Sul Ross State University. We thank an anonymous reviewer for supplying information about pertinent literature and con- structive advice about the manuscript. We are grateful to curator Ron Hartman (RM) who responded so quickly to our request for a loan.
REFERENCES
BAKER, M. S. 1957. Studies in western violets — VII. The Nuttalianae continued. Brittonia 9:217 — 230.
BURGESS, T. L. and D. K. NORTHINGTON. 1981. Plants of the Guadalupe Mountains and mela Caverns National Parks. Chihuahuan Desert Res. Inst., Contr. No. 107, Alpine, T
CORRELL, D. S. and a c JOHNSTON. 1970. Manual of the Vascular Plants of Texas.
Texas aes Foundation, Renner, a
FABIJAN, D. M., J. G. Packer, and K ears 1987. The taxonomy of the Viola mati complex. Gat J. Bot. 65: oo 580.
MART . R. HUTCHINS. On Spring Wildflowers of New Mexico. ca a New Mexice Press, Albuquerque.
ROSE, J. N. and H. D. House. 1905. Descriptions of three Mexican violets. Proc. U.S.
RUSSELL, N. H. oe Violets (Viola) of Central and Eastern United States: an introduc- tory survey. Sida 2: 13.
AN EXPLANATION FOR THE DISCREPANCY IN THE CHROMOSOME COUNT OF THE REDBUD (CERCIS CANADENSIS, LEGUMINOSAE)
WILL H. BLACKWELL
Department of Botany, ag U ae Oxford, OH 45056,
ABSTRACT
Varying chromosome counts of Cercis canadensis L. have been reported in the literature,
.e., 2 = Gand» = 7. Squash piven : pollen parent cells from native trees in southwestern Ohio confirm the count of » = 7. However, past segmental interchanges between nonhomologues are the assumed cause of occasional structural connections observed between two or more bivalents during meiosis. The viewing of such connected chromosomes, perhaps superficially appearing as a unit structure, might result tn an erron- eous count, such as 7 = 6.
A plant species with a low number of chromosomes usually does not present special difficulty in the determination of chromosome number un- less aneuploidy or dysploidy exists in its populations (cf. Claytonta virgi- nica, Lewis 1967). Cercis canadensis, a woody legume, is not known to be aneuploid or dysploid, yet different base chromosome numbers have been reported. Senn (1938) reported » = 6 and 2” = 12, based on both meiotic and mitotic counts at the Blandy Experimental Farm, Virginia. These counts had impact in the framework of Senn’s pioneering work on legume cytology in relation to phylogeny. However, Taylor (1967) reported » = 7 for Cercis canadensis from a “shrub of unknown origin in Dominion Arboretum. Ottawa.” Curtis (1976) subsequently reported 2 = 7 from a specimen cultivated at the Missouri Botanical Garden, and suggested that the » = 6 determination (by Senn) was incorrect. But would a diligent worker such as Senn have made an actual miscount on a species as seeming- ly simple chromosomally as the redbud? The argument takes on additional significance in light of Goldblatt’s (198 1) view of Cerczs as a diploid (2n = 14,” = 7, x = 7)relict in the subfamily Caesalpinioideae, most members of which are presumably tetraploids — often” = 14, butsomen = 12! In Cercis, the ancestral diploid condition is considered to be retained, not only in context of the caesalpinioids, but in that of the legumes as a whole. However, does variation in the chromosome number of Cerczs occur? Is it uniformly » = 7 as one would suspect, or do counts of » = 6 exist as well?
Over the course of three successive springs (centering around early
Stipa 14(1):7 — LL. 1990.
8
April), l obtained meiotic pollen parent cell (PPC) counts from four trees at different locations among the native populations of redbud in southwestern Ohio (Butler County). These specimens are vouchered in the Herbarium of Miami University (MU). Standard aceto-carmine smear methods were em- ployed in all cases. Prior to staining, flower buds were fixed in modified Carnoy’s solution (4 parts chloroform, 3 parts absolute ethanol, and | part glacial acetic acid). Buds containing appropriate meiotic stages were collected typically between 7:00 and 7:30 A.M
I was surprised by finding apparent counts of » = 7, » = 6, and even lower numbers, in all trees (Figs. 1 — 6). However, careful anaylsis by oil- immersion light microscopy, photography, and enlargement of photo- graphic prints revealed that the actual count is » = 7 (Fig. 1). This count (2 = 7) could be documented in the case of each tree. In the squashes, however, associations were rather commonly observed between two or more of the bivalents during at least the first prophase/metaphase of meiosis I. These associations may occur to an extent that chiasma-like structural con- nections exist for a time between nonhomologues (Figs. 2 — 3). These con- nections may persist, as observed between nonhomologous univalents in telophase of meiosis I (Fig. 4). The fact that some cells in a given smear show only unassociated bivalents, while others show apparently catenated chromosomes, is not altogether surprising because plants with reciprocal translocations can occasionally produce a “normal” complement of bivalents at meiosis (reported, but as rare, in Collinsia heterophylla by Garber and Dhillon 1962). Regardless, the associations in Cercis canadensis provide some evidence of a past (perhaps long past) segmental interchange (Burnham 1956) between two or more of the chromosome pairs.
Associations between nonhomologous chromosomes are known within another member of the legume family (lines of garden pea, Pisum sativum, n = 7, cf. Sansome 1932). Although similar to those of the garden pea, the associations in redbud are not as consistently present, nor necessarily as definitive. The chromosomal connections in Cercis are nonetheless in some cases striking enough to result in the appearance of a chain of four (or more) chromosomes at diakinesis (Fig. 5). This chain in redbud bears a close resemblance to the often-cited, excellent photographic illustration of a translocation chain or ring of four chromosomes described by Brown (1949) in irradiated tomatoes. That the associated chromosomes in Cercis are merely connected at a point (Figs. 2— 3) and appear at most connected as a chain (Fig. 5) and not as an actual ring is probably indicative of the past interchange of only small chromosomal segments (Burnam 1932). Nonetheless, to a cytotaxonomic investigator who had not studied the situ- ation in Cerczs, a count other than the correct one of » = 7 might be made,
BTSs: i a Os PPC’ s bok Core canadensis: Fig. 1, lace prophase of meiosis eivision I,” = 7. Figs. 2—3, I ). Fig. 4, telophase of meiosis
t f ,
i persi f ion b ival ). Fig. 5,1 sae of meiosis I; note apparent count of six due to ch me chain in involving two bncliana) fatrow). F g. 6, metaphase of meiosis I: f f« five i ibled ). Scale bar, 20 jum;
’ Tr all photographs are at same magnification.
10
especially if a cell such as that in Figure 5 would be encountered. Even if bivalents are merely “interlocked” (a condition reported as common in diploid species of Tradescantia; Sax and Anderson 1933), an opportunity for a miscount would exist. It seems plausible that Senn (1938) encountered either very closely associated, perhaps interlocked, bivalents, or else an actual reciprocal translocation, when he reported his count of 7 = 6 from PPC’s. Close spatial association of nonhomologues may give rise to natural reciprocal translocations, (suggested as a possiblility by Sax and Anderson 1933). Such translocations may be observed in somatic tissue as well as cells undergoing meiosis (Burnham 1956). Hence, Senn’s 2” (somatic) count of 12, made from anther wall tissue, is also understandable in the light of the interpretation(s) presented here. Regardless, the chromosome count of all species of Cerczs, although aii difficult to determine in the case of C. canadensis, is apparently n = 7, 2” = 14, as considered by Gold- blate (1981).
Study of the cytology of species of Cerczs other than C. canadensis might prove interesting if only to determine if nonhomologous chromosome associations exist in these as well. Since chromosomal connections due to reciprocal translocations may result in varying levels of reduced fertility (Garber 1948), a study of pollen viability (or an analysis of microspore quartets) might be undertaken as well.
ACKNOWLEDGMENTS
J acknowledge the help of Julie A. Ballenger and Michael A. Vincent for assistance with collection of some of the bud and voucher material utilized in this study. | am appreciative of the helpful suggestions of Drs. Roy C. Brown, Thomas G. Lammers, and Askell Léve during the preparation of this manuscript.
REFERENCES BROWN, S.W. 1949. The structure and meiotic behavior of the differentiated chromo- somes of tomato. Genetics 34:437 — 461. BURNHAM, C.R. 1932. An interchange in maize giving low sterility and chain con- figurations. Proc. Natl. Acad. Sci. 18:434 — 440. poe Chromosomal eae ake in plants. Bot. Rev. 22:419—552.
CURTIS, W.E i som in Grielum and Corcis. Ann. Missouri Bot. Gard. 633379 = 38 naa 136 1948. A reciprocal translocation in Sorghum versicolor Anderss. Amer. J. poe 20 F
& TS. DHILLON. 1962. The genus Collinsia. XVII. A cytogenetic study of radiation-induced reciprocal translocations in C. heterophylla. Genetics 47:461 — 467.
GOLDBLATT, P. 1981. Cytology and the phylogeny of Leguminosae. Pp. 427 — 463 in,
11
R. M. Polhill & P H. Raven eds. Advances in Legume Systematics. Part 2. Royal ee ae Kew LEWIS, , R. L. OLIVER, & Y. SUDA. 1967. ee of Claytonia virginica and its are Ann. Missouri Bot. Gard. 54:153— 1 SANSOME, E.R. 1932. Segmental interchange in oe sativum. Cytologia 3:200 — 219 SAX, K. & E. ANDERSON. elias ee 18:53 — 67 SENN, oa fee
1933. Segmental interchange in chromosomes of Trades- ii Chromosome number relationships in the Leguminosae. Bibliogr.
— 336. TAYLOR, R.L. ete In IOPB chromosome number reports XIII. Taxon 16:456.
CRATAEGUS SECRETA (ROSACEAE), A NEW SPECIES OF HAWTHORN FROM THE EDWARDS PLATEAU, TEXAS'
J. B. PHIPPS
Department of shia Sciences University of Western Ontario London, Ontario, CANADA NOA 5B7
ABSTRACT
A new species of Crataegus L. (Rosaceae), C. secreta Phipps, is described from west- central Texas in the drier, northwestern parts of the Edwards Plateau. It is now known from at least a five-county area. The new species is compared with candidates in series Virides and Molles and easily excluded from these series. Its probable affiliation is series Tenwifoliae, but this awaits further stud
The Edwards Plateau area of Texas is an important phytogeographical area. Cooler than the Rio Grande Valley to the south and the Gulf Coast Plain to the southeast, considerably more mesic than most of Trans-Pecos Texas to the west and most of the Chihuahan Desert to the south and south- west, it forms a stepping stone between the Cross-Timbers region of eastern Texas and the higher, more mesic parts of the Sierra Madre Oriental in Mexico. Its vegetation is discussed in detail tn Amos and Gehlbach (1988). The undulating uplands at around 1500 to 2500 ft. a.s.1. are cove- red by a mixture of grassland and scrubby dwarf oaks (Quercus sinuata vat. breviloba - Bigelow Oak, Q. fusiformis - Texas Live Oak and Q. texana - Texas
Red Oak) with junipers (especially J. ashei and _J. pinchotii). The valleys, which are often ravine-like, are quite rich in taxa with northern affinities. They are often more mesic, with taller trees. The ranchland ecosystems, however, are not necesarily in anywhere near their ancestral state with the decreased fire regime imposed by modern pastoralism and the introduction of cattle and goats brought about by European settlement. Goats, in parti- cular, may have mediated massive changes in the woody flora. Also to be taken into account are the elimination of the mobile bison and the presumptively large changes in deer population since large predator removal and control of screw-worm larvae.
This interesting area has been explored for hawthorns by the author in recent years with a view to helping to establish the southwesterly limits of American species of Crataegus and the northern limits of predominantly
‘Editor: Manuscript received for publication 7 Sep 1989, revised and accepted 26 Dec 1989.
Stipa 14(1):13— 19. 1990.
14
Mexican species. Crataegus, hitherto generally held to be rare in the Ed- wards Plateau, have recently been demonstrated to be locally common in this region. It 1s clear that hawthorns are a fairly general feature of the scrub oak woodland (but much less common among junipers) and are not by any means restricted to streams in the Edwards Plateau as formerly believed. Among the species encountered are C. crus-galli L., sens. lat. (locally common in the central Edwards Plateau), C. reverchonii Sarg. (primarily in the northeastern part of the Edwards Plateau), C. mollis (T.&G.) Scheele, sens. lat. (primarily on the eastern margins), C. tracy: Ashe ex Egglest. (scattered at higher elevations) and C. greggiana Egglest. (extremely rare and seemingly relict). Members of series Virides are also found around the southeastern,eastern and northeastern margins of this area
n view of the generally good state of botanical unowiledge of this region it was, therefore, a considerable surprise to encounter a distinctive and apparently unknown species of Crataegus in the Edwards Plateau. The new species has been carefully compared with the regional endemics C. wvalden- sis Sarg. (series Molles - close to or conspecific with C. greggtana) and C. desertorum Sarg. (a xeromorphic form of series Virides also described origi- nally from Uvalde) but these are quite different. Crataegus sutherlandensis, a Texas endemic from just outside the Edwards Plateau, east of San Antonio, was also considered, and also rejected as being a typical member of series Virides. Considering the somewhat isolated nature of the Edwards Plateau, attention was therefore given to regions further afield as possibly having disjunct conspecifics. However,to the immediate west (Trans-Pecos) only C. tracy? is known from the Davis Mountains (this species also occurs in the Edwards Plateau). To the south, no Mexican species is a match (Phipps, in preparation). To the north and northeast, therefore, it seemed prudent to consider species described from Oklahama, southwest Arkansas, and eastern Texas.
The new species is nevertheless in many ways closest to C. greggiana in general facies. The fine straight thorns, scrubby plant habit and small, some what lobed leaves are all suggestive, as is the red fruit. But the slight- ly pubescent, or nearly glabrate, plant parts — foliage, young shoots, in- florescence, and fruit — of the new species, are all dissimilar from C. greggiana as 1s the stamen number. Crataegus brachyphylla Sarg., from southwest Arkansas, is also an obvious candidate species for comparison but it is also more typically Mo//es in its pubescence. Moreover, in spite of its name, it has substantially larger foliage than the new species. All other members of series Mo//es in the region mentioned may be excluded by their much larger foliage and dense indumentum. Species of series Crus-galli are too generally different to warrant even cursory attention. Crataegus secreta
1
may be differentiated from the most similar Crataegus species discussed by the characters given in Table |. It will be seen that one of the most similar of those considered is C. desertorum. However, Sargent’s (1922b) assign- ment of this taxon to series Virides is undoubtedly correct. The other species mentioned in the table are all unquestionably series Mo/l/es. One is,
therefore, left with the conclusion that a distinctive new species of Crata- egus (even though a statistically unlikely phenomenon, given the number of taxa described already in North America) has been discovered.
The first collection of C. secreta may have been Palmer's in 1917 from Menard, Menard Co.; however, this specimen is sterile. The species remained uncollected until 1933 — 35 when a number of specimens from Sutton and Val Verde counties were collected by Cory and by Parks and Cory. McVaugh then collected it in Menard Co. in 1947. The next collec- tion appears to be by Marshall Enquist from Scalp Creek, Menard Co., in April 1986, a specimen of which was donated, along with a allecion of other unidentified Crataegus, to the author. The recognition that the Scalp Creek hawthorn might be a new species has spurred a minor craze in Crata- egus collecting in this Crataegus-poor region of Texas so that now at least 20 numbers of the new taxon exist in herbaria. Due to the ability of this species to resist scientific detection for so long, I am naming it Crataegus secreta. 1930's collections are also reported from Scalp Creek by R. O’Kennon from the University of Texas herbarium but these have not been located by the author.
CRATAEGUS Secreta Phipps, sp. nov. Fi
Frutex vel arbor parva, 3 — 6 m alta, spinosa; spinae rectae, + tenues, + nigrae, usque ad 4 cm longis; cortex platata. Foliae brachyblastorum ovatae, parvae, oe 2 3°¢em apoee distincte lobatae — 4 paribus venarum secondarium, serratae, in apice acutae, + truncatae pro male, laevicer DePacentes (praecipue supra) ubi juvenes, glabrescentes, petiolis pee oliae surculorum elongationum magniores, magis profunde lobatae, carentes venas in sinibus. Anthesis vernalis (in Aprilem); inflorescentia panicula subconvexa, ferens ca 4— 10 flores ee ramuloris pedicellisque sparsim pu
escentibus; lobi calycis 2— 3 mm longi, triangulati, pubescens adaxiale, marginibus + integris (glandulis stipitatis); petala orbiculares, ca 5 mm longa; stamina ca 15, antheribus purpureis; stigmata, styli et carpella (4-) 5; fructus pomum aha parum oblatus, ca | cm diametro, carine farinosi, pyrenibus 5, dorsale sulcatibus.
Bush to small tree 3 — 6 m tall, thorny; thorns straight, + fine, black- ish, to 4 cm long; bark plated. Leaves of short shoots ovate, small, short- petiolate, 2—3 cm long, distinctly lobed, with 3—4 lateral nerves, serrate, acute at the apex, + truncate below, slightly pubescent (especially above when young, glabrescent; leaves of shoots of elongation larger, more deeply lobed, lacking veins to the sinus. Flowering in spring (April), in-
16
Taste 1. Comparison of Crataegus secreta Phipps with selected congeners.
C. secreTA = C. GREGGIANA C. BRACHYPHYLLA C. UVALDENSIS C. DESERTORUM
gglest. Sarg. Sarg. (ex litt.) Sarg. (1909) (1922a) (1922b) (1922b) Series ? Tenuifoliae Molles Molles Molles Virides Distribution Tx: Menard, Tx: Edwards Ark: Tx: Uvalde Co. Tx: Uvalde Co. ason, Plateau Hempstead Co. Schleicher, (rare); utton, Mexico: Val Verde Sierra Madre Be Oriental > Jelt Davis Leaf: length 2¢ 2cm 4-5 cm cm 1.5—2 cm shape = ovate ovate broad-ovate ovate ovate to obovate pubescence subglabrous densely densely pubescent pubescent (young) pubescent pubescent Hypanthial glabrous densely densely densely glabrous pubescence pubescent pubescent pubescen Calyx lobe + entire, irregularly laciniately irregularly obscurely margin with stipi- glandular glandular glandular serrate tate glands serrate serrate serrate to glandular- serrate Stamen no. ca 15 LO 20 5— 10 20 Anther colour purple pink deep rose ? pale yellow Style no. (4-) 5 5 3 35 4-5 Fruit: diameter 10 mm 10 mm 10-12 mm 10-14 mm colour re deep red dull dark red bright red orange-red
florescence a flattish, convex panicle, bearing about 4 — 10 white flowers, branches and pedicels sparsely pubescent; calyx lobes 2—3 mm long, triangular, adaxially strigose-pubescent, margins + entire to glandular- serrate, or entire with some stipitate glands; petals + circular, about 5 mm long; stamens ca 15, anthers purple; stigmas, styles and carpels (4-) 5; fruit a red pome, slightly oblate, ca 1 cm diameter, flesh mealy; pyrenes 5, dor- sally ribbed
Type: TEXAS.: Menard Co.: creekside ca 5 mi E of Menard, 2000 ft, 15 Apr 1988, J.B ici M. eee and R. O'Kennon 6123 (HOLOTYPE: UWO; isorypes: to be disersared):
ollected in fruit is J.B. PAzpps and R. O’Kennon 6233, 1 3 Oct 1988 (UWO).
a other specimens have been collected at the above site. M. Engquist s.n., Apr 1987 (UWO), ae 9 Apr 1986 (TEX) represent earlier Galigenions of this taxon aoe the same location.
FIG. 1. Line drawing of Crataegus secreta Phipps, sp. nov. Fruiting branch, fruit and pyrenes from J.B. Phipps 6233; flowering shoot and flower parts from J.B. Phipps 6121; leaf from J.B. Phipps 6121. Scale bars 1 cm. Susan Laurie-Bourque del.
18
Other specimens examined: TEXAS: Mason Co: | km N of Katemcy, 1880 ft, scrub along creek bed, 15 Apr 1988, Phipps and O’Kennon 6127 (UWO); 14 Oct 1988, Phipps and O’Kennon 6243 (same location) (UWO); 1.6 mi S of jct. 1851 and 1222, west side of ro along Sandy Creek, A. W. Edmiston property, 27 Apr 1989, Enquist 1076 (TEX- LL. UWO). Menard Co.: Route 83, 1 mi S of jet. with Tx 29, 2000 ft, PAzpps, — and O’Kennon - a 15 Apr 1988 (UWO); Phipps and O’Kennon 6239 represents the same plant in fruit, ct 1988 (UWO); on bluffs of San Saba River near Fr. McKavett, 4 ne 1989, Teen in ae and forced, Phipps, Enquist and O’Kennon 6318 (UWO); wooded bottom of San Saba R., near crossing of Ft. McKavett Rd., 17 mi S of west of Menard, scarce, 12 May 1947 R. McVaugh 8787 (SMU). Schleicher Co.: W of Fort McKavett, 3.2 mi W of jct with hwy. 864, first crossing of Middle ee Prong of San Saba River, 19 Apr 1989,
Enquist 996 (TEX-LL, UWO). Sutton Co.: 30 mi SW of Sonora, 4 Apr 1933, V. L. Cory 5505 (TAES); Aldwell Bros. (? ranch), 21 i ie H.B. Parks and V.L. Cory 8433 and 8434 (TAES); south-west quadrant ... near Dry Devil's River, 19 Apr 1989, Enquist
717,732 (TEX-LL, UWO); SW quadrant, by Granger Draw Road near Ree of Dry Devil's River, 19 Apr 1989, Enguist 720 (TEX-LL, UWO). Val Verde C : NE quadrant, on the floodplain of the Devil’s River, 19 Apr 1989, Enquist 710 (TEX IL, U wo). 11 mi NE of Juno, 4 Jun 1939, H.B. Parks and V.L. Cory 31678 (TAES) -doubtful ID, specimen badly damaged by herbarium beetle. : ae that E, J. Palmer 11889 (A) from "low woods on the San Saba River, Menard, ard Co., May 12, 1917 "may also be C. secreta. Since this specimen is sterile, it is not ee re} eee ee C. uvaldensis. However, it is within the area of distribution for C. secreta. Enguist 1144 (UWO) from Musquiz Canyon, Jeff Davis County may represent the same species but confirmatory material is required.
I would like to propose the vernacular name ‘Plateau Hawthorn’ for Crataegus secreta.
Crataegus secreta has now been found at several sites in Menard, Mason, Schleicher, Sutton and Val Verde counties. It occurs near creek beds and in the shade of oak trees, always,so far as is understood, in rangeland. Crata- egus Secreta, although hardly common, is not believed to be under threat, due to a compatable type of land-use in its natural habitat. It should be searched for in adjacent counties.
The sertal affiliation of C. secreta is not clear. On prima facie grounds assignment to series Tenuwifoliae seems obvious. These are very ‘median’ American hawthorns with + ovate, relatively small, shallowly lobed leaves, a modest amount of pubescence, quite thorny, with thorns of median length, flowering early-midseason, with smallish flowers, fruit red of medium size, spherical, with unpitted pyrenes. These characteristics all apply to C.secreta. However, if C. secreta were to be assigned to Tenuifoliae then its somewhat xeromorphic characteristics, particularly short-petiolate leaf, and fine, straight thorns, as well as the precise leaf shape with its unusual lobing would make it the most distinctive members of the series. Furthermore, Tenwifoliae are not primarily either a southern or a xeromor- phic series. Affiliation with series Virides is however, easily rejected where the central tendencies of Vrrides are concerned: the leaves (though usually small) are usually evenly lobed (or sometimes unlobed) and the plants are
19
+ glabrate, not usually very thorny, calyx lobes + entire, with twenty stamens and ivory anthers, and small, + shiny, orange-red fruit. However, Texas Virides can be found with blood-red fruit (a deeper color than C. secreta) but these are more succulent and shiny than C. secreta. Also the sometimes lobed leaf shape in forms assigned to C. desertorum and C. suther- landensis together with a greater thorniness than typical Vzrides, may resemble C. secreta, but then the fruit and calyx are quite wrong. The aforementioned taxa in the series Virides (together with other members of series Vurides) occur around the southern and eastern margins of the plateau. Crataegus secreta also has some striking resemblances to the small- leaved Molles species C. greggiana Egglest. particularly in leaf size, thorni- ness (a very close match) and fruit characteristics (bright red when ripe, though slightly smaller than in C. greggzana), with mealy, not succulent flesh. But then the leaf shape is different and also C. secreta lacks the dense pubescence of all parts that (especially while young) so thoroughly characterize the Modlles series. C. greggiana, of course, occupies the most generally xeric habitats of all North American Crataegus and occurs, although scarcely, on the Edwards Plateau. The stamen number (15) of C. secreta 1s midway between that of C. greggiana (10) and series Varides (20). It is not unreasonable, therefore, to suppose that C. secreta is of Molles X Viri- des hybrid origin for its characters fall midway between these two series, but this hypothesis must await detailed biosystematic and morphometric analysis. If this hybrid hypothesis is true, then resemblances to series Tenuifoliae are coincidental. However, even if of hybrid origin, C. secreta has the marks of a good species, being rather uniform over a five (or six, if Jeff Davis be included) county area of distribution and not merging into any other Texan species. Therefore, for the time being, it seems wise to leave C. secreta unassigned.
ACKNOWLEDGMENTS Thanks to Wm. E Mahler (SMU) for information on Edwards Plateau ecology and for wide ranging searches by Marshall Enquist during the rediscovery phase. I would also like to thank Bob O’Kennon and Marshall Enquist for their comments on the first draft of this paper, January 1989. REFERENCES AMOS, B.B. and ER. GEHLBACH. 1988. Edwards Plateau vegetation. Baylor Univer- sity Press, Waco EGGLESTON, WW. 1909. Crataegi of Mexico and Central America. Bull. Torrey Bot. Club. 86:511. PHIPPS, J.B. (in prep.) The Crataegus (Rosaceae) of Northern Mexic SARGENT, C.S. 1922a. Notes on North American trees, IX. J. at Arbor. 3:8. 1922b. Notes on North American trees, X. J. Arnold Arbor. 3: 187, 195.
THE ALPINE-SUBALPINE FLORA OF NORTHEASTERN MEXICO
J. ANDREW McDONALD
Department of Botany, U sot, y Texas Austin, TX 78713, U.S.A
ABSTRACT
A floristic list of angiosperms found above or in association with timberline vegetation: in northeastern México 1s presented. The flora doubles the number of alpine-subalpine species previously reported for the region, and extends the known distribution of this vegetation type. Included are 170 species, representing 119 genera and 46 families.
RESUMEN
Se presenta un listado floristico de angiospermas que existen en los limites arboreos de zonas altas en el nordeste de México. Se reconoce el doble del numero de elementos alpino- subalpinos reportados para la region en trabajos anteriores, y se extiende la distribucion conocida de este tipo de vegetacion. Se reconocen 170 especies, 1 19 generos y 46 familias.
The isolated presence of timberline vegetation in northeastern México was recognized by Muller (1939), and has since been subjected to few studies. Beaman & Andresen (1966) characterized in detail the ecological and floristic aspects of Cerro Potosi, Nuevo Leon, one of several prominent peaks in ie region. Alpine mean dominated by chamaephytes and hemicryptophytes is encountered on Cerro Potosi from 3620— 3700 m, and subalpine meadow composed primarily of erect forbs and cespitose grasses is found as low as 3460 m. A unique form of subalpine vegetation dominated by dense, shrubby stands of Pznus culminicola Andresen & Bea- man often intercedes the Pinus hartwegii Benth. forests and alpine meadow as low as 3450 m. Based on the above characterizations of alpine and sub- alpine zones, and their associated elevational limits, one would suspect these vegetation types to be more widespread, as the region includes several ranges that reach from 3450 — 3700 m. Contrary to suggestions that Cerro Potosi is the sole center for alpine-subalpine vegetation in northeastern México (Beaman & Andresen, 1966), recent explorations of high elevation- al ranges revealed a more complex and widespread timberline flora.
e timberline vegetation of northeastern México includes three dis- creet centers (Fig. 1). The northern center begins 36 km east of Saltillo in the northernmost extensions of the Sierra Madre Oriental. The closely spa- ced Sierra Coahuilon, Sierra La Marta and Sierra La Viga provide refugia for alpine or subalpine elements along their ridges and upper, southern ex-
Sipa 14(1):21 — 28. 1990.
LAT. N. LONG. W ALTITUDE
1
‘SIERRA LA VIGA 25 21' 100 33' ~— 3700 m éSTERRA POTRERO DE ABREGO 25.19' 100 22' 3460 m SSTERRA COAHUILON 25.14" 100 20' ~— 3500 m 4STERRA LA MARTA 25 12' 100 22-3700 m SCERRO POTOSI 24.53' 100 15' 3700m SSTERRA PENA NEVADA 23 48! 99 51' 3640 m ’STERRA BORRADO 23 47! 99 51' 3460 m
1. Distribution of alpine-subalpine sites explored in northeastern México, including their dldtudes: latitudes and longitudes. Underlined localities included in floristic list.
posures from 3400 — 3700 m. The second center for timberline vegetation, Cerro Potosi, occurs as a singular peak 38 km to the south of Sierra La Marta. Present day maps (DETENAL, Joint Operations Graphic maps, Department of Commerce Operational Navigation Charts) generally place Cerro Potosi at 3700 m, about equal in elevation to Sierra La Marta, the closest point of alpine-subalpine contact to the north. The third and southern center for timberline vegetation is located 125 km south of Cerro Potosi, including Sierra Borrado and Sierra Pena Nevada (Fig. 1). The lat- ter peaks are generally reported to reach 3400 and 3650 m, respectively. As predicted by Muller (1939), Sierra Pefia Nevada also provides satisfactory habitat for shade intolerant, timberline species, which are distributed sporadically with stunted individuals of Pinus hartwegii along the ridges
23
and uppermost southeast and southwest exposures of the range. Sierra orrado, though excluded in the floristic list due to its lack of an es-
tablished subalpine vegetation, deserves mention since many subalpine species are encountered on its upper and relatively open, eastern exposures.
Fieldwork was undertaken during summer months from 1984 — 86. All sites were visited at least once at the beginning of the flowering season (June), during the peak of the flowering season (July — August), and dur- ing the fruiting months (September — October). In addition to the author's collections, complementary material was studied at TEX, where a signifi- cant collection of the Northeast Mexican flora has been accumulated in recent years. Near complete sets of the author’s collections are deposited at MEXU and TEX, and incomplete sets are at UAT, WIS and XAL.
While Beaman & Andresen (1966) reported 81 species for Cerro Potosi, the updated list includes 170 species for the alpine-subalpine vegetation of northeastern México. A few additional species are added to the list for Cerro Potosi, and most species previously listed as endemic to the peak are present and often prevalent in the other timberline refugia. A forthcoming study will analyze in more depth, based in part on the distributional data presented here, the phytogeographic relationships among various alpine- subalpine peaks of northern México (McDonald, in press).
Froristic List
PN PO MACO VI
AGAVACEAE Agave macroculmis Tod. xX XxX BORAGINACEAE Hackelia leonotis 1. M. Johnston Xx X Lithospermum sordidum Brand. xX XxX Onosmodium dodrantale 1.M. Johnston xX xX X CAMPANULACEAE Campanula rotundifolia L. X X X X X CAPRIFOLIACEAE Pei ee H.B.K. X X X X X pisses onan a i Xx X XxX X X Arenaria cf. lycopodioides Willd. X Xx ex Schle cht. a cf. oresbia Gree xX X X XK X oe: Lee feacel ex X X X X ay) Robins.
am = Pena Nevada, PO = Cerro Potosi, MA = Sierra La Marta, CO = Sierra Coahuilon, = Sierra La Viga
24
(Floristic List continued)
PO
MA
Stellaria ae Willd.
Silene lac
CELASTRACE EAE
Paxistima myrsinites Raf.
COMMELINACEAE
Commelina tuberosa L.
COMPOSITAE
Achillea millefolinm L.
Ageratina oretthales (B.L. Rob.) B. Turner
A iis oven (B.L. Rob.) B. Tur
rooms ‘parvifol ta Nut Astranthium beamanti De - see triplinervia H.B.K s
ais coahutlensis (A. Gray) Harcombe & Beaman Brickellta hintoniorum B. Turner
Chaetopappa
gray Cirsium i ae GC. Nesom (in prep)
Dugaldia pinetorum (Standl.) Bierner Erigeron hintoniorum Nesom (in prep) Erigeron onofrensts Nesom (in prep) Erigeron potosinus Stand.
Erigeron pubescens H.B.K.
Erigeron wellsit Nesom
—
Guaphalium hintoniorum B. Turner
(in prep) Grindelia inuloides Willd.
Helianthella eee (Hook.) Gray Heterotheca mucronata Harms ex Turner
Hieracium iynynan Blake Hymenoxys ursina Stand. Hymenopappus ae B. ‘Turner
Senecio bintoniorum B. Turner Senecto loratifolius Greenm. Senecio madrenst iA
Taraxacum officinale Weber in Wigg. Thelesperma eraminiformis (Sherff) Melchert (in prep)
> >
~ xX
~ mK mK
~
~ x
~ x
~ rx
x KK KK
~~
nx xX
ee
xx
~~ xX
x x
mx
~~
~
x x
~ x
x
(Floristic Lise continued)
PO
MA
Thelesperma mullerii (Sherff) M in
Zaluzania sg Sch.-Bip. CRASSULACEA
Sedum sek ee McDonald (in prep) ep)
Sedum papillicaulum Nesom (in Sedum clausentt Nesom (in prep) Villadia cuculata Rose
Villadia misera (Lind].) R. Clausen Echeverta cf. simulans Rose CRUCIFERAE
Draba helleriana Greene
Erysimum capitatum Green
Pennelia longifolia (Benth. : Rollins Thlaspi mexicanum Stan 7UPRESSACEAE
Juniperus monticola Martinez CYPERACEAE
Carex bella Bailey
‘arex. orizabae Liebm. Carex schiedeana Kunze ERICACEAE eaeoe, pungens H.B.K. EUPHOR Euphorbia Her M.C. Johnston FAGACEAE Quercus greggli (A. DC.) Trel. Quercus spp.
FUMARIACAE G. ie eae Fedde GAR C
Ga ovata on var. ovata GENTIANACEAE
ae. amarella (L.) Borner Frasera speciosa Doug).
Halenia alleniana Standl. ex Wilbur GERANIACEAE
Geranium seemanit Pey So Beer Gien HL E. Moore GRA pines tricholepis (Torr.) Nash Brachypodinm a Scribn. ex Beal. pipe anomalus Calamagrostis ee, R. Br.
Deschampsia flexuosa (L.) Trin. Elymus trachycaulus (Link.) Gould
ex Shinners
>
~ mK MK
nm x
x x eK
~ s
~
bd ><
~
~*~ MM
~~ x
~ Kx
*
rs PS PS PS
re re
~
rs Ps
~ Xs
~ xs
~ x
26
(Floristic List continued)
PN
PO
MA
Festuca amplissima Rup Festuca hephaestophila hee ex Steud. Festuca pringler St.-Yves Festuca roser Piper Festuca rubra Festuca spose Vasey Festuca easels E. Pana Kohleria pyramida Mubhlenbergia rigens eee ) Hitch. Mubhlenbergia virescens Trin Muhlenbergia wolf (Vasey) Rydb. Phleum alpinum L Piptochaetium virescens (H.B.K.)
di
‘aro
Poa mulleri Swallen
Poa pratensis
Poa strictiramea A. tch.
Trisetum spicatum (L. . Richeer DROPHYLLACEAE
Nama whalenii Bacon (in prep)
Phacelia platycarpa Spreng. IRIDACEAE
Sisyrinchium schaffneri Wats. Sisyrinchium sp. nov. AE
Agastache palmers (B.L. ag ) Stand.
We macellaria Epl.
Salvia unicostata i
alvia sp McDonald (in prep) Scutellaria ae Brandeg sep pi Benth. LEGUMINOS AE Astrag ae purpusiit M.E. iia Trifolium schneidert Stand Vicia humilis H.B.K Vicia ludoviciana Nutt. Lupinus cacuminis Stand. LILIACEAE Calochortus marcellae Nesom Schoenocaulon sp. nov. Frame (in prep) Maianthemum stellatum (L. . Link set virescens (H.B.K.) MacBride LINACEA
Linum ae Pursh
re Ps OX OK ~~ XK
*
~ KK
~ x
mere eK
~ x
xX
~
ms
mK
Km
~~
re
~ x
re
rm
a
(Floristic List continued)
PN PO MA CO
2)
VI
ONAGRACEAE Epilobium angustifolium L. ssp. circumvagum Mosquin Oenothera priminervis A. Gray Ocenothera tetraptera Cav. APAVERACEAE Argemone subalpina McDonald (in prep) ORANTHACEAE Arceuthobium vaginatum (Willd.) Prest. SSp. vaginatum PINACEAE Pinus culminicola Andresen & Beaman Ae hartwegti Picea mexicana M. Martinez POLEMONIACEAE Polemonium pauciflorum Wats. POLOGONACEAE Ertogonum jamesti Benth.
Androsace ae ee is L ar. puberulenta (Rydb. ) Kunth
RANUNCULACEAE Aquilegia elegantula Greene Delphinium valens Stand. Ranunculus praemorsus H.B.K. ex DC. RHAMNACEAE Ceanothus buxifolius Willd. ex Schult. Ceanothus greggii Gray
OSACEAE Alchemilla procumbens Rose Fragaria californica Newberry Holodiscus dumosus (Nutt.) Heller Potentilla leonina Stand. Potentilla gers Rydb. Potentilla s v. Nesom (in prep) Rubus Bis L RUBIACEAE Galium uncinulatum DC Hedyotis ee (A. Gia) Fosberg SALICACEA Populus mais Michx. SAXIFRAGACEAE Heuchera mexicana eee Heuchera sanguin Philadelphus nae fai ) Hu Ribes neglectum Ros
Ribes microphyllus H. B.K.
> >< > ><
b >< b>
~ rr rs rm PK
~ XK
28
(Floristic List continued) PN PO MA CO VI
SCROPHULARIACEAE Castilleja bella Stand}. Castilleja a ae H.B.K. Penstemon barbatu
Penstemon es ie
SOIL E
xm mK rx Ke
Solanum verrucosum Schlecht. Solanum macropilosum Correll Physalis orizabae Dun. xX xX UMBELLIFERAE Arracacia schneideri Mathias & xX xX xX Constance Arracacia ternata Mathias & Constance x Arracacia tolucensis Hemsl. X xX Eryngiun ao > ean Constance &
x x
~~
Iter eg eats Coult. & Rose X XxX X X URTIC Urtica 7 a Blume xX Xx VALERIANACEAE Valeriana sorbifolia H.B.K. X var. oe
VERBENACEAE
Verbena an H.B.K. xX XX X VIOLACEAE
Viola Skee M.S. Baker Xx
Acknowledgments The following botanists assisted in the identification of the listed species: John Fay (Commelinaceae), Billie Turner and Guy Nesom
(Compositae), Nelly Diego (Cyperaceae), Manuel Gonzalez Ledesma,
Charlotte and John Reeder (Gramineae), John Bacon (Hydrophyllaceae),
.P. Ramamoorthy (Labiatae), Peter Hoch (Onagraceae), Daniel Nickrent
(Loranthaceae), Tom Duncan (Ranunculaceae), David Lorence
(Rubiaceae), Mike Nee (Solanaceae), David Boufford (Urticaceae) and Fred
Barrie (Valertanaceae). Their collaboration is gratefully acknowledged.
REFERENCES
BEAMAN, J.H. 1966. The vegetation, floristics and phytogeography of the summit of Cerro Potosi, México. Amer. Midl. Nat. 75:1— 33.
MCDONALD, J.A. (in press). Phytogeography of the alpine-subalpine flora of north- eastern México. in: T.P. Ramamoorthy, J. Fa, R. Bey & A. Lot (eds.), Biological Diver- sity of México: Origins and Disteibucions. Oxford Press, London
MULLER C.H. 1939. Relations of the vegetation and climatic types of Nuevo Leon, México. Amer. Midl. Nat. 21:687 — 729.
IDENTIFICATION OF THE PLANTS ILLUSTRATED AND DESCRIBED IN CATESBY’S NATURAL HISTORY OF THE CAROLINAS, FLORIDA
AND THE BAHAMAS
ROBERT L. WILBUR
ae of Botany, eae U eo urham, NC 27706, U.S.A.
Perhaps it will surprise some that after nearly 250 years botanists are still unable to identify several of the plants described and illustrated by Catesby (1730 — 1747) concerning a flora that surely must rank among the best known in this hemisphere. In addition a considerable number of Catesby’s plants can be identified only approximately or that, at the very least, legit- imate cause exists for debate over their identities. I believe that the explan- ation of this unsatisfactory state is that Catesby’s illustrations are very much lacking in those features that botanists depend upon in order to identify plants and that Catesby’s abilities verbally to describe the plants were if anything even less developed than his talents as a biological drafts- man. Each group of biologists, after noting the unsatisfactory rendition of the organisms in groups in which they are most expert, usually then in- dicates that Catesby’s greatest talents were ina group other than that which the investigator was most familiar. My conclusion is that the overall evalua- tion of Catesby’s biological depiction is not high as the details and even major features are often either not shown or are poorly depicted. The lack of detail and crudity in representation is indeed unfortunate since for many plants and animals Catesby was either the only one or a prime reference in those Linnaean publications that became the starting points in biological nomenclature. Ewan (1976, p. 89) noted that Linnaeus cited Catesby’s work ninety-five times in Species plantarum (1753), the starting point for most botanical nomenclature, and Linnaeus in later works or other authors later added to this number in the publication of additional new species based on Catesby’s Natural History. Howard and Staples (1983, p. 511) in their paper dealing only with plants concluded that “Catesby’s plates appear to be the types of twenty-five recognized taxa, of which twenty-one were described by Linnaeus and four by subsequent authors.” These plates were also found by them to be “the types of an additional twelve synony- mous names.” Clearly then the significance of Catesby’s work, artistically crude and almost completely devoid of significant botanical detail though
Sipa 14(1):29 — 48. 1990.
30
the plates may be, is undeniably great since these plates are in some cases considered to be the types upon which a given binomial rests.
More than three decades ago I began this study of the identities of the plants included in Catesby’s Natural History of the Carolinas. I soon en- countered obstacles that prevented me from completing the investigation in a timely manner. As might be expected some of the obstacles have in time been either directly solved by the publications of others or their work has enabled me to make progress when before I could not. Some of the obstacles that could not then be overcome by me have been solved by my increasing experience that time and greater familiarity with the plants in the field and the literature about them provides. To my chagrin Howard and Staples (1983) published a commentary on Catesby’s Natural History that largely fulfilled what I had only partly completed two decades before. They pointed out a prior and similar study to their own published by Ewan (1976) of which I was completely unaware. Since some of my conclusions differed significantly from either one or both of these two most recent studies, it seemed worthwhile to place on record my conclusions along with the reasons for my differences. The nature of such a study makes it certain that we can only hope to approach perfection incrementally. Hope- fully the future will judge that some progress in interpreting the identities of Catesby’s plants was made in this account. I would be remiss not to acknowledge the assistance and stimulation I obviously received from both Ewan’s and Howard and Staples’ earlier commentaries.
For those interested in learning about the life and accomplishments of Mark Catesby (1682 — 1749), the best source is Frick and Stearns (1961) “Mark Catesby, the Colonial Audubon.”
Some might consider that my criticism of the botanical draftsmanship and phytographic skills of this early colonial naturalist is too harsh. After all the various commentators have managed to identify the vast majority of the organisms depicted of both plants and animals. Perhaps, as a counter balance, Frick’s evaluation (1974) ought to be quoted: “The flaws of the natural History of Carolina are minor in comparison with its virtues . . . No other mainland area had so complete a natural history before the American Revolution as did South Carolina and eighteenth century Georgia, and certainly none so elegant. Mark Catesby’s achievement was unique.”
It might be meaningful to those who are very slightly statistically orien- ted to compare the differences between the three commentaries presented in the table. (I suggest though that these comparisons though are really not meaningfully subjected to statistical comparison, or, if so, not to the very unsophisticated comparisons made here where any change be it in autho- rity or in spelling was tallied as a change equally important as a change in
31
identity.) Be that as it may be, between Ewan and Howard and Staples there is a 24.5% difference, between Ewan and Wilbur there was a 28.5% difference, and between Howard and Staples and Wilbur a 10.2% change.
The identifications of the plants in Catesby’s Natural History made by me and the two most recent commentators are arranged in three parallel columns in the following comparative table. Where there are differences in identification, I have provided a brief explanation in the numbered foot-
notes referred to in the right-hand margin.
IDENTIFICATION OF CATESBY S
PLATES
Ewan (1974) Howard and Staples (1983)
Wilbur (1990)
Vol. I
Castanea pumila (L.) Marsh.
9. 9. Castanea pumila (L.) Miller 10. Columbrina reclinata (L Her.)
10. ellie eliptica (Sw.) Briz. Stern
IL. ie distichum (L.) Rich.
~~
: ce distichum (L.) Rich.
13. Myrica ae Loisel. 13. A pensylvanica Loisel. Ory L,
14. Oryza sativa L. 14. Oryza sativa 15. Smilax anf L. a S ee laurifolta L. 16. Quer tlos L 6. Quercus phellos L. es he 5a (L.) L. {sic!] . Quercus pie Miller 18. Quercus prinos L. {sic!] 18. Quercus pri a Quercus marilandica Muenchh. 19. Quercus arial Muenchh. 1 a. Quercus nigra 20a. Quercus nigra L. am alias ao L. 7 Mile nm L. 211. oe
211. Quercu.
r nae us aes
— c
i. os a Que la en 2. Quercus incana Bate 23. Quercus laevis Walte 24. Podophyllum ee L. 25. Chrysobalanus taco L
3. Quer 4 Pui tan L. rysob oe, Zanthoxylum a ercults L. 27. Cornus florida L.
alanus Waco : 26. Zanthoxylum ae hercults L. 27. Cornus florida L (Weston) oa. 28. Prunus virginiana L.. 29. Aristolochia serpentaria L. - Bursera simaruba (L.) Sarg.
bra rubra 28. Prunus virginiana L. 29. Aristolochia serpentaria L. 30. Elapbrium simaruba L.
ai Tex ¢ . Tex cassine L.
32. Uniola la > U niola a, L. 33. Hypoxis hirsuta (L. ver
34. Populus balsamifer
33. Hypox 34. Panidin ae 35 Ipomoea sagittata ca
Monotropa uniflora
9. Castanea pumila (L.) P Mill.
10. esis eliptica (Sw.) Briz. =|
11. Tesodam distichum (L.) L.C.
IB: Moriae ee Raf. *2
14. Oryza sativa L.
Quercus virginiana P. Mill. . Quercus michauxti Nutt 19. ee marilandica Musuehiy
ercHs sp 22, ae incana Barer. *5 23. Quercus laevis . *6 24. Podophyllum pan 1 25. Chrysobalanus i 26, Zanthoxylum ae peau L. 27. Cornus florida L
28. Prunus serotina Ehrh. *7
29. Aristolochia serpentarta L.
- Bursera simaruba (L.) Sarg. *8
. [lex cassine L.
2 Uniola paniulata L.
33. boxis sp.
34, is pees L. *10 35.
uniflora L. 36. os 36. 4 pe L. tia bahamensis (Northrop) 37. Tabebuia bahamensis (Northrop)37. Labebuia pe: (Northrop)
ritt. 38a. Carya tomentosa (Poir.) Nutt. b. Carya cordiformis (Wang.) K. Koch
38a. a alba (L.) K. Koch arya cordiformis (Wang.) Koch
38a. oe tomentosa (Poir.) Nutt. *11 b. € P. Mill.)
arya glabra (
sl 2
oP
(Identification of Catesby’s plates continued)
39. Magnolta virginiana L 39. Magnolia virginiana L. 39. Magnolia virginiana L.
40. Metopium toxiferum (L. ) Krug 40. die toxtferum (L.) Krug 40. ea toxiferum (L.) Krug Urban ban 41. Nyssa aquatica L. ae sylvatica Marsh. 41. Ni ‘pot Mest *13 42. Jacaranda caerulea (L.) Griseb.42. Jacaranda caerulea (L. )Griseb.42. J re ia 43. Gleditsia aquatica Marsh. 43. Gleditsia aquatica Marsh. 43. Gletitsia wie Mars 44. Gordonia lasianthus (L.) Ellis 44. Gordonia aes (L.) Ellis44. Gordonia lasianthus (L. : Ellis 45. Trillium catesbaei Ell. 45. Trillium catesbae 4S. Trillium catesbaet Ell. 46. Calycanthus lie Gy L. 46. Calycanthus te L. 46. Calycanthus floridus L. 47. Smilax herbacea L. 47. Smilax pumila Doan 47. Smilax pumila Walt. *14 48. Liriodendron tulipifera L. 48. Liriodendron tultpifera 48. Lirviodendron tulipfera L. 49. Catalpa bignontoides Walt. Be Catalpa bignonioides ae A Catalpa bignontoides Walt. 50. \ rillium sessile L . Trillium maculatum Raf. 7 ais amas Raf. *15 51 enispermum he Ey 5 . Cocculus carolinus (L.) DC. : : Coce (L.) DC. *1 52) —o bona-nox L. 52. Smilax tamnoides L. 52. Smilax an seen mix- SONS AT
pec 53. Gelsemium sempervirens (L.) Ait.53. Gelseminm sempervirens (L.) 53. Genin ce (L.) -Hi
4. Symplocos tinctoria (L.) LHer 54.
Aiton J. St.-Hil. Symplocos tinctoria (L.) LHer.54. Symplocos tinctoria (L.) LU Her.
Sassafras albidum (Nutt.) Nees55. Sassafras albidum (Nutt.) 55. Sassafras albidum (Nutt.) cn
- Platanus occidental ts 7. Rhododendron viscosum a ) Torr. 7
Nees var. mo//e (Raf.) Fern. Platanus occidentalis L. 56. Platanus occidentalis Rhododendron viscosum (L.) 57. Rhododendron viscosum a ) Torr. Torr. var. aemulans Rehder
58a. Cleistes divaricata (L.) Ames 58a. Cleistes divaricata (L.) Ames58a. Cleistes divaricata (L. : Ames b
- Echites eee aed. Echites umbellata Jacq. . Echites umbellata Jac
59.C€ pfolia Jacq.) Ur ae Casasta clustifolia Jacq.) Urban 59. Casasta ed (aca ) Urb. 60. Nyssa ite yssa aquatica L. 60. Nyssa aquatica 61. Osmanthus americanus (L.) Osmanthus americanus (L.) 61. Osmanthus americanus a )
Benth. & Hook xray Benth. & Hook.f.ex A. Gray 62. Acer rubrum L. 62. Acer rubrum L. 62. Acer rubrum L. 63. Persea ie - ) Sprengel 63. Persea borbonia (L.) Sprengel 63. Persea borbonia (L.) Sprengel 64. Halesia carolina 64. Halesia tetraptera Ellis 4. Halesia tetraptera Ellis *20 65. Campsis radicans a Seem. 65. Campsis radicans (L.) Seem. 65. si a fae e ) Seem. 66. Clethr ifolia L 66. Clethra sedis L. 56. Clethra 67. Juglans nigra L. 67. Juglans nigra L. 7. Juglans nigra 68. Chionanthus virginica L. 68. Chit ee L. ri Cia virginicus L. 69. Myrica cerifera L. 69. Myrica cerifera L yrica certfera L. 70. Gentiana catesbaer Walt. 70. Gentian cae Walter 70. Gentiana catesbaei Wal 71. Oxydendrum arboreum (L.) DC.71. (L.) DC.71. Oxydendrum arboreum ra )DC.
72. Salmea petrochiodes Griseb. Bales ed
77. Phymosia abutiloides (L.) a
- Scaevola plumierii - ) Vahl 79. . Fraxinus american a {sic!] ie 2 Orontium aqua 83. Peltandra es (Michx. ) . Morong
85. Avicennia nitida Jacq. 85. 86. unidentified 86.
Salmea pets Griseb. 72. Salmea petrobioides Griseb. Ri ate Urb. 75. Reynosia septentrionalis
Sl Ham.77. He ae arbuttlordes (L.
am. Scaevola plumiert (L. : Vahl = 79. Sen lama (L.) Vahl Praxin nus americana 80. Fraxinus caroliniana P Mill. *21
rontium aqua L. 82. ae pee L. Peltandea 5 ©) Schott 83. Peltandra virginica (L.) Schott & Engler [sic! & Endl. *22 Avicennia germinans (L.) L. 85. Avicennia germinans (L.) L. *23 Laguncularta racemosa (L.) 86. Laguncularia racemosa (L.) Gaertn Gaertn. *24
(Identification of Catesby’s plates continued)
92. Wedelia babamensis (Britt.) 92. Wedelia bahamensis (Britt.) Schulz 93. Borrichia arborescens (L.) DC.
98. Jacquinia keyensis Mez
ulz 93. Borrichia arborescens (L.) DC. 98. Jacquinia keyensis Mez
Vol. Il
24. Ecastophyllum brownei Pers. 24. el ecastophyllum
(L. 26. Xylophylla epiphyllanthus 26. Plan epiphyllanthus (L.) Brite
28a. Ocotea cortacea (Sw.) Britt. 28a. oe cortacea (Sw.) Britt. b. Galactia rudolphioides . Galactia rudolphioides (Griseb.) Hook. & Arn. (Griseb.) Benth. & Hook 30. Samolus ebracteatus H.B.K. (?)30. Unidentified 32. Picrodendron macrocarpum 32. Picrodendron baccatum
(A. Rich.) Brice. (L.) Krug & Urban 33a.Conocarpus erecta L. 33a.Conocarpus erectus L. b. Amyris elemifera b. Amyris elemifera I
38. Thallasia testudinum Konig 38. Thalassia testudinum Konig 421. Leucaena glauca (L.) Benth. 421. Lystloma latisiliquum Benth t.Banara reticulata Griseb. r gee cn ee (A. 43. ee racemosa (L.) Gray Willd.
h.) Sleumer iv Leucothoé racemosa ao . Unidentified legu
14. Acacia tortuosa (L.)
is Colocasia esculenta ) Schott 45. AJlocasia sp. or eine sp.
46. Croton eluteria (L.) Sw. 46. Croton eluterta (L.) 47. Callicarpa americana L. 47. Callicarpa americana a 48. Cissus tuberculata Jacq. 48. Cissus tuberculata Jacq
50. Canella winterana (L. ) Gaertn. 50. Canella winterana (L.) Gaertn 5 la.Caesalpinia oo Lam. b. Passiflora pallida L. $2. Decumaria are ara L. 53. Urechites lutea (L.) Britt.
Sla. Cjaline bahamensis Lam. . Pass Hikhoe suberosa L. 52. Unident 53. Urechites oe ae ) Britt.
54. Silene v ita 54. Silene vi 55. renee minuta (Aubl.) 55. Play concreta (Jacq.) Brite. ara
56. Lilium michauxit Poir. as Lilium ones | Ilex vomitoria L. [sic!] os Lilinm catesbaei Walt. 59. Echinacea purpurea (L.) 59. Echinacea purpurea (L.) oenc s (L.) Lam. Ipomoea batatas (L.) Lam. grandiflora L. 62 Cima virginica L. . Rhizophora sea L. 6A. Annona glabra
enc 60. ae batata. 61. Magnolia g
M agnolia
64. Annona glabra L.
65. Liquidambar styraciflua L.
66. Haemotoxylum campechianum L.66. Haematoxylon ¢ Hae
65. Liguidambar ‘noi L ; ampechianum the
matt i is hipinal spelling. ]
92. Wedelia bahamensis (Britt.) O.E. Schulz
93. Borrichia arborescens (L.) DC.
98. Jacquinia keyensis Mez
24. oe ae (L.) L.) T. 26. Phen iis *26
28r. Oo cortacea (Sw.) Britt. sl aaa j ae & Hook. 30. Un identified ae 32. Picrodendron baccatum ru, z 33a. Gumens erectus : b. Amyris elemifera 38. Thalassia ss Konig 421. Lysiloma latisiliquum Benth. *: r. oe aia (A. Sleumer *38 43. ee racemosa (L.) A. Gray
or Xanthosoma *31 eC roton ees
na 50. Canella winterana (L. ) aertn. Sla.Caesalpinia babamensis a b. Passtflora suberosa 32 52. Unidentified * 53. Urechites lutea (L. ) Britt. 54. Silene virginica L. 55. led concreta (Jacq.) weet *34 - Lilium aie L. *35 . Ulex vomitoria Ait ah Lilium catesbaet Walt. 59. Echinacea purpurea (L.) Moench 60. Ipomoea batatas (L.) Lam. 1. Magnolia grandiflora L. 3
gla : 65. Liquidambar styraciflua L. 66. Haematoxylum campechianum L.
34
(Identification of Catesby’s plates continued)
67. Annona cherimola Mill 67. Annona glabra L. 67. Annona glabra L. *3 68. Epidendrum nocturnum Jacq. 68. Epidendrum nocturnum Jacq. 68. aii nocturnum Jacq. 591 691 691. § nia minor Walt. *38 r. Sarracenia flava L. 1. Sarracenia X catesbaei f. ae flava L. ea ) Bell 70. Sarracenia purpurea 70. acenia purpurea 70. acenia purpur 71. Symplocarpus foetidus a ) Nutt.71. palosiens arse i. ) Nutt. : sme futids - ) Nutt. 2. Cypripedium calceolus L. 72. Cypripedium acaule Aiton . Cypripedium acaule Ait. *39
_ Cyp 73. Cypripedium calceolus vas. 73. Cypripedium pubescens Willd. es Cypripedium pubescens Willd. I sorrel
74. Epicladinm babiamam 74. Epidendrum boothianum 74. Encyclia boothianum (Lindl.) (Lindl.) Smal indley Dressler * 75. Sideroxylon ae Jacq. 75. pe ae foetidissimum 75. Mastichiodendron foetidissimum I ) Lam (Jacq.) Lam * 76. Diospyros virginina L. 76. Diospyros virginiana L. 76. Diospyros a L. 77. Catopsts berteroniana (Schultes)77. pi berteroniana (Schultes)77. Catopsis berteronian (Schultes) Mez (Schultes) Mez J.A. & J.H oes Mez 78. Spigelia martlandica L. 78. Spigelia marilandica (L.) L. 78. Spipelig marilandica (L.) L 79. Bourreria ovata Miers 79. Bourreria ovata Miers 79. Bourrerta ovata Miers 80. Magnolia macrophylla Michx. Magne ape ue )L. 80. Magnolia tripetala (L.) L. *42 8la. Swietenta mahogani Jacq. L.) Jacq.8la. Swietenia mahagoni (L.) Jacq. b. P, a rubrum (L.) ae Phoradendron Ed. (L.) b. Phoradendron rubrum (L.) Gri riseb. : sis lata (L.) Bur. 82. Bigno nonia a carea L. 82. Bignonia anc L. *43 83. Prelea trifoliata 83. Ptelea trifoliata L 83. Ptelea trifoliata 84a. peal om L. 84a. Philadelphus sido L. 84a. Philadelphus acer b. Smilax lanceolata L. b. Smilax lanceolata L. b. Smilax smallii Morong *44 85. Asimina ie (L.) Dunal 85. Asimina triloba (L.) Dunal 85. Asimina triloba (L.) Dunal 86. Annona reticulata L. 86. Annona reticulata L. 86. Annona reticulata L 87a. Sloanea emarginata L. 87a. Manilkara bahamensis Lam 87a. Gk kava babamensis Lam Meeuse Meeuse *4 b . Lpomoea microdactyla Griseb. b. a moea ila Griseb. 881. Epidendrum plicatum Lind). re Epidendrum plicatum 881. Encyclta plicat Lindley Britt. & Millsp t.Epidendrum cochleatum L. « Epidendram cochleatum L. r.Encyclia cochleata (L.) Lemee 89. Tillandsia fasciculata Sw. 89. Tillendsia balbis 89. Tillandsia balbisiana (Schultes) R & Schult Schultes f. *47 90. Thespesia dae (L.) Soland.90. Hubiscus tiliaceus L. 90. Hibiseus tiltaceus L. *48 Dla. Cordia se Dla. Cordia sebestena L. Dla. Cordia sebestena L. . Ipomoea es ra b. Ipomoea carolina \.. b. Ipomoea carolina L 92. Plumeria rubra 92. Plumeria rubra L. 92. Plumeria rubra L. 93a. Plumeria obtusa L. Da. Seis sei a 93a. Plumeria obtusa L. . Passiflora cupra b. Passiflor b. Passiflora cuprea L 94. Coccoloba piss Tea 94.C€ es liver cri Jacq. 94. Coccoloba diverstfolia Jacq. 95a.Hippomane mancinella L. 95a.Hippomane mancinella L. 95a.Hippomane mancinella L. Dendropemon ee (L.) b. san purpureum (L.) — b. selene purpureum (L.) rug & Urba Krug & Urban Kru Urban 96. Coccoloba uv fer ra (L.) Jacq. 96. Cocc slabs uvifera (L.) L. 96. Peis uvifera (L.) L. 97. Pithecolobium mucronatum 97. Pithecellobium seme 97. Pithecellobium bahamense Britt. ort Northrop *49 98. Kalmia an L; 98. Kalmic “latifolia L. 98. Kalmia letifolie L.
99. Clusia rosea Jac 99. Clusia rosea Jacq. 99. Clusia rosea Jacq.
(Identification of Catesby’s plates continued)
100. Raed Spinosa L. di
100. an Spinosa L.
100. a Spinosa L. Appe
endix endix 1. aaa eadia L 1. Dibba on mi 1. oe media L. Hamamelts virginiana L 2. Hamamelis virginiana i 2. Hamamelis virginiana L. 3. Cypripedium acau oe Gh eae acaule Ait. 3. ee =) ae *50 4. Rhus glabra 4.R 4. Rhus glabr. 5. Pancratium carolinianum L. 5. eis aie (L.) 5. Hele eiieiiehee (L.) Herbert Herbert *51 6. Theobroma cacao L. 6. Theobroma 6. Theobroma 7. Vanilla planifolia Andr. 7. Vanilla mexicana Miller 7. Vanilla ae Ande #52 8. Liltum philadelphicum L 8. Lilium philadelphicum L. 8. Lilium philadelphicum L. nacardium occidentale L 9. Anacardium occidentale L 9. Anacardium occidentale L 11. Lilium canadense L. Ll. Liltum canadens 11. Lilium canadense L. 12. fe atamasco (L.) 12. ie a atamasco (L.) 12. Zephyranthes atamasco (L.) erbert Her Herbert 13. panei malacodendron L. 13. wae malacodendron L. 13. Stewartia malacodendron L. 15. Magnolia acuminata ( L 15. Magnolia acuminata (L.) L. 15. nolia acuminata (L.) L 16. Panax quinquefolium L 16. Panax quinguefolius 7 16. Panax quinquefolius L. *53 171 lmia augustifolia L 171. Kalmia angustifolia 17] ngustifolia L 1. Rhododendron dia L dron maximum L. . ¢. Rhododendron maximum L. 18. Ficus brevifolia N 18. ee citrifolia Miller 18. Ficus citrifolia P. Mill. *54 20. Robinia hispida L 20. Robinia hispida L 20. Robinia hispida L
1) Johnston (1971), che most recent monographer of Colubrina (Rhamnaceae), included Colubrina reclinata (LHér.) Brongn. in the synonymy of Colubrina elliptica (Sw.) Brizicky & Stern
2) Although Catesby’s illustration is certainly not detailed enough alone to permit one to een species of Myrica, geographic distribution is of considerable assistance. It s been identified as Myrica pensylvanica Loisel. by Ewan and also by Howard and aie However, I believe it to be eae haenpyl Raf. as Myrica cnsaiea occurs no further south than northeastern Nort rolina while Myrica heterophylla mmon in the coastal plain from northern oo into southern New England nnn: of course coastal South Carolina, the site of Catesby’s most intensive work. Linnaeus (1753, p. 1024) cited this Catesby plate as the only element of the B {var.] of Myrica cerifera
3) The two eastern chestnut oaks were n rom each other by Linnaeus or by other botanists. Early in the nineteenth centuty W illdenow (1805, 4:440.) proposed Q. montana as the name for the mountain chestnut oak before Nuttall’s publication (1818, 2:215) of Q. michauxii for the swam ded under the binomial Q. prinus L. Hardin (1979) recommended that botanists discon- tinue using the binomial Q. prinus L. since the material in the Linnaean herbarium cannot be determined with certainty van the Linnaean binomial has been applied almost equally to either species. However most authors in recent decades have applied Quercus prinus L mountain or tock chestnut oak (= Q. montana Willd.) and Quercus michauxtt to the swa innaeus included a reference to Catesby’s account and plate in the synonymy of oan prinus but it is to be remembered that he included both species of chestnut oak under Q. prinus. Catesby's treatment was clearly that of the
36
es ee oak, ee michauxit Nutt., as his statements as to habitat and morphol- ate. Hardin’s suggested solution seems tempting since we have no way of bowing what is meant when Q. prinus is used alone in the literature without synonyms mon names or the mention of the other chestnut oak that had been originally ae with it. 4) Ewan did not make note of the ere ae rendition of the oak depicted on the right side of Catesby’s plate 1:1.2/ and I find both the illustration and brief description unidentifiable. Linnaeus (1753, p. 996) cited Catesby’s account of this taxon as a syn- onym of Q. rubra {var.} B. Howard and Staples indicate it to be Quercus rubra L. which would be difficult to prove — or disprove from Catesby’s ici aan. Linnaeus inclu- ded within his concept of Quercus se L., comprising both the typical element and the 8 variant, the very distinctive southern red or Spanish oak (Q. falcata), the turkey oak, Q. laevis.) as well as the red (or northern red) oak (Q. rubra). After a most rancorous series of papers dealing with the lectotypification of Q. rubra, extending through muc of the first half of the century we hopefully have settled the application of the name.
5) Ewan (1974, p. 92) no doubt carelessly identified this ee account as Quercus oo
laevis Walt., the turkey oak with pinnately lobed leaves. Linnaeus p. 994) based his Quercus ple {var.] y solely upon this citation of Catesby. The i description given by Catesby both confirm that Howard and Staples were correct in identifying the
plant as ac ae jack oak, Quercus incana Bartr. (=Q. cinerea Michx.), with its unlobed s
6) Although ne plate and account was included by Linnaeus in the synonymy of Quercus rubra, it should be remembered that Linnaeus included under that binomial several of the eastern of North hes red oaks: Quercus falcata Michx., Q. laevis Walt. and Q. rubra s.s. Catesby surely was dealing with the turkey oak, Q. /aevis, as noted by Howard He Staples and not with the northern red oak, Q. rubra, as sug ges- ted by Ewan.
7) rea like Linnaeus and most eighteenth century Sukie did not distinguish between Prunus virginiana L. and Prunus serotina Ehrh. The description and plate do not
wide the necessary details to enable us t ae what Catesby had. The scanty teins with its indication of Aaa large size and indication of abundance in the thick woods of Carolina make it certain that the plant Catesby knew from field experi- ence was Prunus serotina Ehrh. ee virginiana 1s unknown in South Carolina and very rare in the mountains of North Carolina and unknown elsewhere in that state.
8) The generic name Buarsera Jacq. ex L. (1762) is conserved over Elaphrinm Jacq. (1760).
9) Like Howard and Staples, I do not find that Catesby’s plate of what appears to be an Hypoxis can be identified to species. The description with its mentioned five perianth segments and 5 stamens instead of 6 is most unusual. Detailed information needed to make specific determinations is lacking.
10) I agree with Rouleau (1946, 106) and with Howard and Staples (1983, p. 536) that Catesby ee the common coastal plain, swamp poplar of the Carolinas, Populus heterophylla and neither P del/toides L. with its nan flattened petioles nor P balsamifera es each it has been synonymized in the
11) Constant juggling with the provisions of the International Code of Botanical Nomen- clature would seem to be a perfect prescription for instability in nomenclature. For over
37
four decades we have enjoyed relative stability in the scientific names of two of our commonest hickories but this stability seems threatened due to nomenclatural tinker- ing. Carya alba (L.) K. Koch had been abandoned at least since the mid- 1940s as an ambiguous name (see Rehder, 1945) since it was sometimes applied to the mockernut hickory (Carya tomentosa (Poir.) K. Koch) and sometimes to the shagbark hickory (Carya ovata (Mill.) K. Koch) as Linnaeus had included both tn ee alba. Originally no type was designaced ee sad alba, and hence it would appear Article 69 in its 1978 version of the ICBN could not be applied. The current ae of Art. 69 permitting the abandonment of names used in two or more senses not including the type hardly applies when no type was designated and the original concept proves to have been a mixture. Earlier versions of Article 69 rejected a name “if it is used in different senses and so has become a long-persistent source of error.” Howard & Staples argued that Juglans alba L. was typified by Crantz (1766, 1:157) since Crantz cited only Catesby in his brief account of Juglans alba This three-line account by Crantz consisted of the following:
2. IvGLaNns alba. IUGLANS foliis es wnee serratis, impari sessili. CATESB ys Yan 07
It would not seem that such action constitutes pert unless the author makes it clear that he intends to remove dissident elements from the protologue. No evidence exists chat Crantz was doing more than citing ae ek mentioned in the eee seen by him. Therefore, Carya tomentosa (Poir.) Nutt. is the correct binomial for th mockernut hickory. Just as is the case for Quercus prinus L. as suggested by Hardin, the best solution might well be to one Carya alba as a name used so often in such different senses that it would be better to exclude it from scientific This was proposed by Rehder (1945). Dr. James Luteyn of the New York erent Cas most kindly provided me with a copy of Crantz’s treatment.
12) Ewan (1974, p. 93) cis Donald E. Stone's identification of the separate, single n of Catesby’s 1:t.38 as Carya cordiformis (Wangenh.) K. Koch. Howard and oe (1983, p. 528) od this determination without comment. Ina genus as notoriously variable as is Carya, one surely must hesitate to determine the identity of a species based ona single nut especially when the artist is as careless as Catesby repeatedly demonstra- ted he was. Probably overly influenced by the most usual application of the common name, I had thought the sketch of the fruit and description referred to Carya glabra (P. Mill.) Sweet. Since the apparently sca globose fruit lacked a ridged husk, the identi- fication a at least possibly c t. Sargent state (1895, 7:167) that the “earliest authentic account of Hicoria lane ‘with an excellent figure of the nut, appeared in Gacy S Neel es of Caroli ” However it would be unwise to make much of a wager on the identity of a great many ne Catesby’s plates especially on one in which only a single fruit is ene
13) I agree with Eyde (1959 and 1964) and Howard and Staples (1983, p. 533) that 7 s plate and cemnaaes (1: 4.41) 1s Nyssa sylvatica Marsh. and not Nyssa aquatica
. as identified by E
14) The fruits of this species were illustrated and descibed by Catesby as “red of an oval f ees with Smilax pumila Walt. and is in conflict with the black, ane berries of § ae L. with which Ewan (1974, p. 93) identified it. Catesby (1:1. 47)
38
stated chat each berry has “a very hard pointed seed” which is true of S pumila Walt. (see Coker, 1944, p. 60), while the berry of S herbacea L. has “3 — 6 brownish seeds” accord- ing to Mangaly (1968, p. 250).
15) Although Linnaeus cited to Catesby 1: ¢.50 in the protologue of Trillium sessile L., Freeman (1975) demonstrated that the Linnaean species in the modern restricted sense does not occur in coastal South Carolina and is represented there instead by Trillium maculatum Raf.
16) The fruits of Menispermum canadense are black while those of Cocculus carolinus are red. Catesby’s description and plate are of red fruit and Catesby’s 1:1.5/ illustrates Cocculus.
17) The identity of Catesby’s plate is both crucial to nomenclatural stability and highly controversial. Fernald (1944, p. 38) stated that there “can be no question that the type of S. tamnoides L. was the Catesby ae Fernald concluded that Catesby’s plant was a perennia a woody, terete-stemmed vine. Howard and Staples (1983, p. 517), although accepting Fernald’s ere eons of Catesby’s plate, indicated that “a specimen obtained by Kalm (LINN 1132. 10) is preferable as leccotype” of S. tamnoides. Fernald had excluded Kalm’s ae from S. tamnoides as it was “a specimen of the herbaceous S. Pseudo-China.” Clausen (1951, p. 109) reached a very different conclusion as to the identity of Catesby’s plate and hence of the identity of Smilax tamnoides L. Clausen agreed that “Catesby's description and illustration are all important in the typification of 5S. tamnoides” but concluded with, I feel, convincing evidence that “Catesby’s illustra- tion and description were prepare ed from diverse materials” and “probably no species exists with the combination of characteristics as depicted.” Evidence was presented that two and more probably three species entered into Catesby’s description and illustration. Clausen concluded, since it was impossible to make a definite identification of what Catesby had, that the Linnaean name should be disregarded as “ambiguous.” It would seem to me impossible to identify Catesby’s plate and, as the specimen of the herbaceous element also included in the Linnaean protologue of S. tamnoides is of a herbaceous species and identifiable with S. psewdo-china L., it would seem for the present at least the woody species had best be known as Smilax Pispide Muhl. ex Torr
18) There is an obvious discrepancy in the authority of the combination of the binomial i acleagies (= Bignonta sempervirens L.) The combination is usually attributed n or Ait.f. (1811) and not to his father, W. Aiton (1789). Jaume Saint-
Hilaive (1805) apparently first made the combination Gelsemium sempervirens.
19) Eyde (1959, p. 212 and 1964, p. 130) stated that Catesby’s 1: .6/ and the accompany- ing description are of Nyssa aquatica L. The plate and description support this decision and argue against Ewan’s identification of it as Nyssa ogeche Bartr. ex Marsh.
20) The general confusion and misuse of the names applied to Ha/esia Ellis ex L. has been exhaustively dealt with by Reveal and Seldin (1976) and their clarifying conclusions are reflected by Howard and Staples (1983) and by me.
21) Fernald gi p. 390) pointed out that, although cited by Linnaeus in the protologue f Fraxinus americana L., Catesby’s plate and description clearly apply to the “southern Water-Ash which we call EF caroliniana P. Mill.”
22) Catesby’s plate (1: ¢.83) and description clearly is that of the green spathed, greenish berried Peltandra virginica (L.) Schott & Endl. and not the white spathed, red berried P sagittifolia (Michx.) Morong.
a,
23) As demonstrated by Compére (1963) among others, the correct name for the Afro- American Black Mangrove is Avicennia germinans (L.) L. and not Avicennta nitida Jacq.
24) In spite of the depiction of alternate leaves in 1:t.86. by Catesby, the plate surely is a crude representation of Laguncularia.
25) The generic name Dalbergia L.f. (1782) is conserved over the earlier Ecastaphyllum P. Br. (1756)
26) The aes Xylophylla L. was segregated from Phyllanthus L. based upon an erroneous description of the flower as pointed out by Webster (1956, 37:94). The segregate genus Xylophylla L. has been maintained by very few authors in recent decades.
27) Catesby’s 2:t.30 seems to be a badly garbled account and depiction of a most im- probable mixture. One can hardly trust the desc ription as it seemingly has internally contradictory statements e.g. the description of the fruit. Since it is said to be a shrub up to twelve feet high, Ewan’s suggestion that it is ae ebracteatus HBK. can be ruled out as a possibility. The flowers possibly suggest anaes in the Lauraceae like Litsea
aestivalis (L.) Fern. but the capsular fruit seems more suggestive of some member of the Andromedae like Lyonta or age This ee continues to resist all attempts at its identification.
28) Correll and Correll (1982, p. 410) place Pécrodendron macrocarpum (A. Rich.) Brite. in the synonymy of P. baccatum. C.D. Adams (1972, p. 216) is more uncertain for under P. baccatum he states “Probably endemic,” but P macrocarpum (A. Rich.) Britt., occurring
Bahamas, Cuba, Hispaniola and Grand Cayman is suggested as probably not really res As might be expected others take an intermediate position treating the el- ement occurring in the Bahamas as Picrodendron baccatum var. bahamense Krug & Urb.
29) Both Ewan and Britton and Millspaugh (1920, p. 162) identify Catesby’s 2: ¢ ee Soe sensu authors which has been shown by de Wit ae ) to be Lewcaena leucocephala ( .)de Wit. Catesby’s treatment describes a plan ry high” eee ‘large straight trunks some being three feet in diameter” and ‘ Whisee ee limbs.” pod was described as “an inch broad and almost five long.” The wood is said to be the i ae Bahamas afford and of the quality to be shipped to England. All of these features exclude Leucaena. The plant represented is probably Lysiloma latisiliquum (L.) Benth.
30) ae basionym of Banara minutiflora (A. Rich.) Sleumer (= Ilex minutiflora A. Rich., 845) has priority over Banara reticulata Griseb. (1860
31) The diagnostic details needed to distinguish between Xanthosoma and Alocasia are not made evident in Catesby’s generalized plate. Ca/ocasia can be ruled out as it has peltate leaves.
32) Although Linnaeus recognized three species of Passiflora in what is today treated as one variable species, oe exists as to which is the correct name. Dr. John McDougal ( an authority on the meso-American Passifloraceae, has looked into the problem and to dae has not : found any author earlier than Master (1872) who has unequivocally placed one name in the synonymy of the other. Master treated P. pallida L. as a variety of P. suberosa L. which would establish P. saberosa as the name to be maintained if the taxa were combined. MacDougal found that Robert Combs (1897, p. 424) appears to be the first author who unequivocally reduced one species to the synonymy of the other and he also chose to retain Passiflora suberosa L. This choice of binomials should settle the matter at least until someone finds an earlier publication that unequivocally made another hoice
40
33) Like Howard and Staples (1983, p. 540 — 542) Iam unable co accept Ewan's deter tion that the plant was Decumaria barbara L. The “certain discrepancies of habit, lowes color, and corolla shape are et coo ee to areeDe such an identification.” Like
them I am unable to suggest ndidate for the name. Decumaria is a woody
Oo Pp vine with opposite leaves which are much more ovate than the alternate, elliptical leaves of Catesby’s plate and description. The inflorescence of Decumaria is a cymose corymb while that of Catesby's plate is basically racemous. Catesby states the fruit to be 2- parted; Decumaria is 7 — 10-loculate.
34) Although its basionym is the first name applied to the species, the combination Polys- tachya minuta (Aubl.) Britt. (1903) is a later homonym of P minuta Rich. & Gal. (1845) and consequentially cannot be used.
35) The identity of 2:4.56 is somewhat controversial as the differences between Lilinm
michauxti Poit. and L. superbum L. are too subtle to be distinguished by either Catesby’s artistic skills or his er in phytography. Since ae L. superbum grows in Pennsylvania (Wherry, Fogg and Wahl. 1979: p. 103) that part of Catesby’s account can be assigned
with confidence. The at of the plate, although not based on the Pennsylvania plant, I Ne also identify it as L. superbum since its leaves seem more elliptical than spatulate.
the majority of the plate was ie from ee: Carolina material as seems more ee then Ewan’s identification as L. michawxii Poir. seems more understandable since that species is Senne in nt Carolina and L. superbum does not occur in South Carolina. However, the depicted leaves appear to fit L. superbum better than do those of L. michauxtt.
36) Both Ewan (1974, p. 97) and Howard and Staples (1983, p. 515) identified Catesby’s 2:t.02 as Commelina virginva L. but that Linnaean species has all blue petals while Catesby’s description indicates “two blue petals . . . and one very small white petal
’ Therefore it seems more probable ioe Cates by had Commelina erecta L. whose oven would at least match this description of the petal colors.
37) Ewan identified Catesby’s 2:1.67 as Annona cherimolia P. Mill. but that species has three large outer petals and three minute, scale-like inner petals while Catesby’s description calls for six fee sce P. cherimola is a montane species and is certainly not to be expected in the Bahamas and was not reported from those islands by either Britton and Millspaugh en or by the Corrells (1982). Catesby’s plate is almost certainly Annona glabra
38) Identification of the plants in this plate is difficult and the three i interpretations of it reflect our collective uncertainties. The plate is not carefully delineated and the colors are particularly unsatisfactory. Elliott (1824, 2:11) cites Catesby’s plate as part - the protologue of his Sarracenia catesbaei and How and Staples disposition of 2:1.69 reflects chis eeaios etation. The only suggestion of Catesby’s plate being Sarracenia cates- baer is that the venation of the flap-like hood is said to be 2 a Elliott's type of $ catesbaet is usually judged to be a hybrid between S. flava and S. purpurea and this is reflected in that the petals of the hybrid, instead of being cee yellow as they are in § flava or dark maroon as they are in S. purpurea are said by Bell (1952, p. 61) to be maroon externally and red-yellow internally. Catesby’s plate is no match for that description but it is equally a poor match for S. flava as its petals are depicted (at least in the copy I have seen) as a sickly greenish yellow. In spite of what is said above I feel that there is nothing in Catesby’s account or plate (the right-hand figures) that would exclude S. flava as the
41
most likely identification. The hood-like or cowl-topped leaf shown on the left side of the plate is in my opinion a crude effort to picture the distinctive leaf of S. minor Walt.
39) The difficulty in attempting to identify many of Catesby’s plates is demonstrated by Catesby’s rendition (2:1. 72) of this lady’s-slipper. The illustration is, like a large number in the two volumes, more of a crude caricature than a reasonable rendition of the botani- cal features upon which identification must rest. Ewan (1972, p. 94) identified the poor picture as C. calceolus, the yellow lady’s-slipper, and Howard and Staples (1983, p. 516) and Wilbur have identified it as C. acaule. The deeply fissured lip and the hint of red in the lip are about all there is to defend the latter choice. [lustrations indeed must border on being wretched if one has difficulty in distinguishing between two such dissimilar species.
40) The differences in our three identifications of Catesby’s 2: ¢. o merely reflect the three different commentators accepting different standards in the rapidly changing i dismemberment in such large orchid genera as the broadly once Epidendru
41) All are agreed as to the identity of Catesby’s 2: 4.75 but reflect the well-founded dismemberment of such broadly conceived genera as Sideroxylon L., now restricted to the Old World, by accepting the genus Mastichodendron Lam. as the American segregate.
42) Catesby’s description and plate are again not easy to reconcile with what exists in
pressed Linnaeus when he cited Catesby’s 2: ¢.80 in synonymy of Magnolia virginiana {var.} tripetala. This is in considerable conflict with the somewhat cordate or auriculate leaf base of M. macrophylla. No indication is evident on the plate or in the description that the leaves are other than green beneath while the lower surface of the leaves of M. macrophylla are strikingly white-glaucous. Catesby stated that the leaves of this species of Magnolia “are usually thirty inches in length” which greatly influenced Ewan in his identification of Catesby’s plate as M. macrophylla which has leaves reportedly up to 10 dm long. The leaves of M. macrophylla according to Fernald (1950, p. 676) are 3 —9 dm long while Radford, Ahles & Bell (1968, p. 476) state them to be up to one meter long. Comparable figures stated by these last authors for Magnolia see are 3 — 6 dm long and 1—4.5 dm long. In spite of the striking lack of agreement in leaf length by these authors, it would seem that Catesby’s stated size of the leaves nee fits M. macrophylla. The lack of detail in both illustration and description as to the pubescence on young twigs, buds and follicles prevents using these prime distinguishing features to separate the two species. On balance it seems to me that it is most likely that Catesby’s 2:1.80 represents Magnolia tripetala.
43) The discrepancy in the comparative table between Ewan and the other two commen- taries on the identity of che plant shown in 2:/.82 1s more apparent than real. There has been much eiscassion on the type of the Linnaean genus Bignonia over at least the past century and t have only recently been resolved by fiat of the International Botanical ee Something of the background can be gleaned from papers by Gentry (1972) and by Wilbur (1980). The result is that the International Code of Botanical Nomenclature (1988, p. 265) has listed Bignonia L. as conserved with Bignonia capreolata L. as its type. Consequently the current correct name is Bignonia capreolata L
44) Fernald (1944b) carefully analyzed the confused tangle into which this greenbrier had rown in the past two centuries and concluded that Smilax lanceolata L. was based upon
42
Virginian material and was nothing more than “the narrowest-leaved S$. /aurifolia” with
the expected black fruit. Catesby’s 2: ¢.84 is described as a non-spinous plant wit r
even scarlet berries. Catesby’s plant is Smlax smallii Morong which in Fernald day was unknown north of northern coastal North Carolina but is included in the recent Atlas of the Virginia Flora (see Harvill ef a/. 1986, p. 25). In decyphering the tangled history of Smilax se a but applying equally well to the history of a great many of the species liscussed in notes, Fernald (1944b) made ae following perceptive observation: “One sometimes ee the wisdom of starting our nomenclature of American plants with Linnaeus (1753). It is almost an exceptional North American species about which he was not hopelessly confused.”
45) Although S/loanea emarginata L. is the first binomial given to this species, the generic name is typified by a member of the Elaeocarpaceae and S. emarginata is a species of Manilkara (Sapotaceae). The Linnaean binomial cannot be transferred to Mani/kara as
re is an earlier Hawaiian species named Manilkara emarginata Lam (1925). Correll and Correll (1982, p. 1099), Long & Lakela (1971, p. 681) and Little (1979, p. 170) all treat this species as Manilkara bahamense (Baker) Lam & Meeuse. Cronquist (1945 and 1946) considers it to be but one of four subspecies which together comprise Manilkara jaimiqui (W right) Dubard. The south Bahaman and Cuban representative was treated as Manilkara jaimiqui ssp. emarginata (L.) Crong
46) The recent tendency among orchidologists has been to segregate distinctive groups of species from the formerly all-inclusive genus Epidendrum L. One of the most distinctive groups of approximately 150 species has been segregated as Encyclia Hook. and i characterized by its column being either free from the lip or at most partially adnate to it
while in Epidendrum the column is completely adnate to the lip (see Dressler 1961).
47) Smith (1938, p. 136 and 1977, p. 985) cites Catesby’s account and plate as illustrating Tillandsia balbisiana while Britton and Millspaugh (1920, p. 65) identify Catesby’s account with T. fasciculata Sw. | take the unscientific expedient of casting my vote with the more eminent authority on the Bromeliaceae. The differences between the two species strike me as too subtle to be discernible from either Catesby’s vague plate or description.
48) Linnaeus (1753, p. 694) cited Catesby 2:¢.90 with the treatment of Hibiscus populneus L.
Catesby’s description and plate both indicate the ges calycine teeth of Hibiscus
tiliacens which contrast greatly with the truncate calyx of Thespesia with which Ewan (1976, p. 99) equated it following Linnaeus. Britton 4 Millspaugh (1920, p. 273) correctly cited Catesby 2: ¢.90 with Parti eee (L.) St. Hil., a synonym of Hebiscus
tiliaceus L.
49) The difference sis the three commentaries ee Pithecellobinm are of little
consequence. Correll and Correll’s observation (1982, p. 678) has convinced them that
the alleged dificienees between P. mucronatum Britt. ex Coker and P. bahamense Northrop are of no taxonomic significance.
50) Although we are all agreed that Catesby’s ¢.9 of the eau must be Cypripedium acaule Ait., it should be pointed out that this plate well demonstrates the crudeness 0 y of Caeby: illustrations. The two leaves supposedly at basal in this species are eee as being borne about the midpoint of the stem and separated from each other by more than an inch of stem. It is by elimination that one determines the ae
of many of Catesby’s plates rather than by the faithfulness of the illustration
43
51) Again we are all agreed alist eis must be henley caroliniana (L.) Herb. or its basionym, but there is tion as to just what the name ae ‘a hope to resolve this uncertainty must await a badly ee revision of the gen
52) Until the much-needed revision of the genus Vani//a is undertaken and completed, one can scarcely be dogmatic as to the identity of Catesby’s plate or for that matter even o
e of most widely cultivated species of the genus. The protologues of the earliest
named species seem often to be mixtures and it seems impossible to straighten out the confusion until a modern revision is completed. Fawcett and Rendle (1963, a rearrangement of the 1910 edition, p. 118) indicated “that some of the old drawings peo V. inodora rather than V. pompona or V. planifloia, e.g. Catesby’s plate (Nat. Hist.
rol., App. t.7) which is quoted by Miller as his V. mexicana.
53) In spite of the fact that Linnaeus treated the genus Panax as neuter, the eo | is sculine in accordance with it classical treatment (see Flora N. America 28B:
54) General agreement exists that Ficus ss Nutt. (1846) is a synonym of Ficus citri-
folia P. Mill. (1768). A sampling of recent authors treating the two binomials in this
manner include Correll and Correll ae p. 419), Little (1979, p. 131), DeWolf (1960, p. 146) and eee (1988, p. 60).
APPENDIX: TAXA SYSTEMATICALLY ARRANGED
GYMNOSPERMS TAXODIACEAE Hy DROCHARITACEAE Taxodium distichum (L.) L. C. Rich. (1: Thalassia testudinum Konig (2: ¢.38) t.1l) LILIACEAE BeOS EER MS a caroliniana (L.) Herb. (2 App.: MONOCOTS 1.5)
Hypoxis sp. (1: £.33) Lilium pean L. (2 App.: 4.11) Lilium catesbaei Walt. (2: ¢.58)
AMARYLLIDACEAE (see Liliaceae)
ARACEAE om : : ; Lilium shiladeliphicin L. (2 App: 1.8) Orontium aquaticum L. (1: ¢.82) Pelcandra Lilium superbum L. (2: ¢ virginica (L.) Schott & Endl. (1: 1.83) Trillium catesbaei Ell. (1: ¢. ae Symphoricarpus foetidus (L.) Nutt. (2: Trillium maculatum Raf. (1: 7.5 71) ?Alocasia or Xanthosoma (2: 1.45) Zephyrancthes atamasco (L) Herb. e App.: 4.12) BROMELIACEAE ORCHIDACEAE Catopsis berteroniana (J.A. & J.H. Schultes) Cleistes divaricata (L.) Ames (1 £.58 above) z (2: t.77) Tillandsia balbistana Schule. Cc ypeipediam acaule Ait. (2: ¢.72 and 2 App.: f. (2: £.89) 3) Cypaipi pubescens Willd. (2: £.73) COMMELINACEAE (= C. calceolus var. ise (Willd.) Correll) Commelina erecta L. (2: t.62) Encyclia boothianu alae (2: t.74) Encyclia cochleata a 1S e (2: 1.88 right) GRAMINEAE Encyclia plicata (Lind1.) ye & Millsp. (2: t.88 left)
Oryza sativa L. (1: ¢./4)Uniola paniculata L. (1: cae : poWarclepenicuals§ Epidendrum nocturnum Jacq. (2: ¢.68)
44
Polystachya concreta Jacq.) Garay & Sweet. (2: #95)
Vanilla planifolia Andr. (2 App.: ¢.7)
SMILACACEAE
Smilax lanceolata L. (2: ¢.84 below) Smilax laurifolia L. (1: 4.15)
milax pumila Walt. (1: ¢.47)
Smilax spp. (a hopeless mixture) (1: 7.52)
ee)
DICOTS
ACERACEAE
Acer rubrum L. (1: ¢.62)
ANACARDIACEAE Anacardium occidentale L. (2 App.: ¢.9) Metopium toxiferum (L. so rug & ib. a 1.40) Rhus glabra L. (2 App.: ANNONACEAE Annona glabra L. (2.4.64 and 2 1:67) Annona reticulata L. (2: ¢.86) Asimina triloba (L.) Dunal (2: 7.85) APOCYNACEAE Echites umbellata Jacq. (1: 2.58 below) Plum obtusa L. (2: ¢. . above) a rubra L. (2: ¢.92 Urechites lutea (L.) Britt. o. 1.53) AQUIFOLIACEAE Ilex cassine L. (1: ¢.3/) Ilex vomitoria Ait. (2: ¢.57) ARALIACEAE Panax quinquefolius L. (2 App.: ¢./6) ARISTOLOCHIACEAE Aristolochia serpentaria L. (1: ¢.29) BERBERIDACEAE Podophyllum pelratum L. (1: ¢.24) BIGNONIACEAE
Bignonia capreolata L. (2: 4.82)
Tabebuia balemaencis (Norbiop) Bane (1: t.37)
BORAGINACEAE
Bourreria ovata Miers (2: Cordia sebastena L. (2: ¢. ie oui
BursERACEAE Bursera simaruba (L.) Sarg. (1: ¢.30) CALYCANTHACEAE Calycanthus floridus L. (1: 4.46)
CANELLACEAE Canella winterana (L.) Gaertn. (2: ¢.50)
CARYOPHYLLACEAE
Silene virginica L. (2: ¢.54)
CHRYSOBALANACEAE
Chrysobalanus icaco L. (1: ¢.25) CLETHRACEAE
Clethra alnifolia L. (1: ¢.66) COMBRETACEAE
Conocarpus erectus L. (2: ¢.33 above)
Languncularia racemosa (L.) Gaertn. (1: ¢.86) COMPOSIATAE
Borrichia arborescens (L.) DC. (1: ¢. ae
Wedelia bahamensis (Brite. ) O. E. Schuly (1: 1.92) CONVOLVULACEAE Ipomoea batatas (L.) Lam. (2: ¢.60) w)
Ipomoea microdactyla Griseb. (2: 1.87 below) Ipomoea sagittata Poir. (1: ¢
CoRNACEAE Cornus florida L. (1: f.27)
EBENACEAE Diospyros virginiana L. (2: 1.76)
Ericaceak (and see Monotropaceae)
Kalmia angustifolia L. (2 App.: ¢.17 left) Kalmia latifolia L. (2: 1.98)
Leucothoé racemosa (L.) A. Gray (2: ¢.43) Oxydendrum arboreum (L.) DC. (1: ¢.77)
Rhododendron maximum L. (2 App.: ¢.17 right Rhododendron viscosum (L.) Torr. (1: ¢.57)
EUPHORBIACEAE
Croton eluteria (L.) Sw. (2: ¢.4
Hippomane mancinella L. (2: ¢. 95 above)
Phyllanthus epiphyllancthus L. (2: ¢.26)
Picrodendron baccatum (L.) Krug & Urb. (2: 1.32)
FAGACEAE
Castanea pumila (L.) P Mill. t.9)
Q Quercus sp. (1: 4.21 right)
FLACOURTIACEAE
Banara minutiflora (A. Rich.) Sleumer (2: 4.42 ight
GENTIANACEAE
Gentiana catesbaei Walt. (1: ¢.70)
GooDENIACEAE Scaevola plumieri (L.) Wahl (1: ¢.79) GUTTIFERAE Clusea rosea Jacq. (2: ¢.99) HAMAMELIDACEAE Hamamelis virginiana L. (2 > t.2) Liquidambar styraciflua L. . 4 65) JUGLANDACEAE
Carya glabra (P. Mill.) Sweet (1: £.38) arya tomentosa ae Nutt. (1: £.38) Juglans nigra L. (1: ¢.67)
LAURACEAE
Ocotea coriacea (Sw.) Britt. (2: t.28 above) Persea borbonia (L.) Sprengel (1: 1.63) Sassafras albidum (Nutt.) Nees (1: 7.55)
45
LEGUMINOSAE a) Mimosoideae
Acacia tortuosa (L.) Willd. on
Lysiloma latisiliquum (L.) B on t.42 left)
Pithecellobium bahamense Rcehrop (2: 1.97)
b) Caesalpinoideae
Caesalpinia bahamensis Lam. (2: ¢,51 above) (1: 1.43)
editsia aquatica Marsh. t.
Haematoxylon campechianum L. (2: f.66)
c) Papilionoideae
Dalbergia ecastophyllum (L.) Taub. (2: ¢.24)
Erycthrina herbacea L. (2: 1.49
Galactia rudolphioides (Griseb.) Benth. & Hook. (2: t.28 below)
Robinia hispida L. (2 App.: ¢.20)
LOGANIACEAE
Gelsemium sempervirens (L.) J. St. Hil. (1: t.53)
Spigelia marilandica (L.) L. (2: 1.78) LORANTHACEAE (INCL. WISCACEAE) Dendropemon purpureum (L.) Krug & Urban
(2: t.95 below) Phoradendron rubrum (L.) Griseb. (2: ¢.81/ below) MAGNOLIACEAE
Liriodendron tulipifera Teele sf
Magnolia virginiana L. (1: ¢:39) MALVACEAE Hibiscus tiliaceus L. (2: ¢.90) Phymosia abutiloides (L.) Desv. ex Ham. (1: t.77) MELIACEAE Swietenia mahagoni (L.) Jacq. (2: ¢.81 above) MENISPERMACEAE Cocculus carolinus (L.) DC. (1: ¢.51) MONOTROPACEAE Monotropa uniflora L. (1: ¢.36) MORACEAE
Ficus citrifolia P Mill. (2 App: ¢./8)
46
MyricackAE RUTACEAE Myrica cerifera L. (1: 4.69) Amyris elemifera L. (2: ¢.33 below) Myrica heterophylla aL (1: #13) Prelea trifolia L. (2: 1.83)
Zanthoxylum clava-herculis L. (1: 4.26) NyssackAE
APOTACEAE Nyssa aquatica L. (1: ?
Nyssa sylvatica vee a t.41) Manilkara bahamensis Lam & Meeuse (2: ¢.87 above) eee Mastichodendron foetidissimum (Jacq.) Lam Chionanthus virginicus L. (1: £.68) Fraxinus caroliniana P Mill. (1: ¢.80) SARRACENIACEAE
Osmanthus americanus (L.) A. Gray (1: ¢.61) . ; Sarracenia flava L. (2: ¢.69 right) Sarracenia minor Walt. (2. 1.69 left) PASSIFLORACEAE Sarracenia purpurea L. (2: ¢.70) Passiflora cupraea L. (2: 1.93 below) Passiflora suberosa L. (2: ¢.5/ below) SAXIFRAGACEAE (INCL. HyDRANGEACEAE) Philadelphus inodorus L. (2: 1.84 above) PLATANACEAE . ; STERCULIACEAE Platanus occidentalis L. (1: ¢.56) Theobroma cacao L. (2 App.: £.6) POLYGONACEAE & ea TY RACACEAE Coccoloba diversifolia Jacq. (2: t.94) Halesia tetraptera Ellis (1: .64) Coccoloba uvifera (L.) L. (2: ¢.96) SYMPLOCACEAE
PRIMULACEAE
Dodecatheon meadia L. (2 App.: ¢.1)
Symplocus tinctoria (L.) LHér. (1: ¢.54) THEACEAE
RHAMNACEAE Gordonia lasianthus (L.) Ellis (1: 1.44) Stewartia malacodendron L. (2 App.: ¢.13)
Colubri lig (Sw.) Briz. ne t.10) Reynosia septentrionalis Urb. (1: ¢.75) Ti esnGn an vaene
REG AeGak Jacquinia keyensis Mez (1: 7.98)
Rhizophora mangle L. (2: #.63) VERBENACEAE
Avicennia germinans (L. es L. (1: 7.85) ROSACEAE Callicarpa americana L. 1.47)
Prunus serotina Ehrh. (1: ¢.28) VITACEAE
Cissus tuberculata Jacq. (2: 4.48)
RUBIACEAE Casasia clusiifolia (Jacq. , oe (1: £.59) UNDETERMINED PLATES Catesbaea spinosa L. (2: ¢. 100) (2: ¢.30) Mitchella repens L. (1: ¢. ss below) (2: t.52)
REFERENCES ADAMS, C.D. 1972. Flowering plants of Jamaica. Univ. of the West Indies. Mona,
Jamaica. AITON, WM. 1789. Hortus kewensis. London. 3 vols.
47
AITON, W.T. 1810— 1813. Hortus kewensis. London. BELL, C.R. 1952. ae hybrids in the genus joey . "Elisha Mitchell Sci. Soc. 8:55 — 80. pl. 107.
BRITTON, N.L. . C.E MILLSPAUGH. 1920. The Bahama flora. New York. i-viii, 1— 694
CATESBY, M. 1730— 1747. The natural history of Carolina, Florida and the Bahama Islands. . . 2 vols. folio. Londo
CLAUSEN, R. T. 1951. Smilax hispida versus S. tamnoides. Rhodora 53: 109 —
COKER, W.C. al = woody smilaxes of the United States. J. Elisha Hee a Soc. 60:27 —69. pl. 9
COMBS, R. 1897. ie eolieeved in the district of ea Province of Santa Clara, Cuba in 1895 —96. Trans. Acad. Sci. St. Louis 7:393—491. p/.30— 39.
COMPERE, P. 1963. The correct name of the Afro-American black mangrove. Taxon 12:150— :
CORRELL, D.S. and H.B. CORRELL. 1982. Flora of the Bahama Archipelago. J. Cramer. (50)+ 1692 pp.
CRANTZ, H.J.N. von 1766. Institutiones rei herbariae. Wien. 2 v
CRONQUIST, A. 1945. Studies in the SapOrace ac: IV. The North nen species of Manilkara. Bull. Torrey Bot. Club 72:550— 562.
1946. ie in the Saporeere VI. Miscellaneous notes. Bull. Torrey Bot. Club 73:465 — 47
DEWIT, H.C.D. 1961. neds and correct names of Acacia villosa Willd. and Leu- caena glauca (L.) Bth. Taxon 10:50—54.
DEWOLE G.P, Jr. 1960. Ficus in the flora of Panama. Ann. Missouri Bot. Gard. 47: 146— 165.
DRESSLER, Oy L. 1961. A reconsideration of Encyclia (Orchidaceae). Brittonia
ELLIOTT, - 1816— 1824. A sketch of the botany of South Carolina and Georgia. Charleston. 2 vols.
EWAN, J. 1974. Notes. pp. 89— 100 in the facsimile edition of The natural history of Carolina, Florida and the Bahama Islands by the late Mark Catesby. Beehive Press. Savannah, Georgia.
EYDE, R.H. 1959. ts discovery and naming of the genus Nyssa. Rhodora 61:209 — 218.
4. Typification of Nyssa aquatica L. Taxon 13:129— 132.
FAWCETT, W. and is B. RENDLE. 1910. Flora of Jamaica. 5 vols. incomplete. London. (vol. 1 was reprinted in Kingston, Jamaica in 1963).
FERNALD, M.L. 1944a. Smilax pseudo-china L. in Overlooked as transfers and novel- ties in the Flora of Eastern North America. Rhodora 46:32 — 39.
1944b. Notes on Smilax lanceolata. ee 46:39 — 42. 1946. Types of some American trees. J. Arnold Arbor. 27:386 — 394.
pl. 1 FREEMAN, J D. 1875. Revision of Tri//ium subgenus Phy/lantherum (Liliaceae). Brittonia 24k,
FRICK, G. : 1974. Introduction in the natural history of Carolina, Florida and the Bahama Islands ... by the late Mark Catesby. Beehive Press. Savannah, Georg
FRICK, G.E and R.P STEARNS. 1961. Mark Catesby, the colonial ees University of Illinois Press. x + 137 pp.
GENTRY, A.H. 1972. The type species of Bignonia L. Taxon 25:659 — 664.
HARDIN, J.W. 1979. Quercus prinus L. — nomen ambiguum. Taxon 28:355 — 357.
48
HARVILL, A.M. JR, T.R. BRADLEY, C.E. STEVENS, T.E WIEBOLDT, D.M.E. WARE, and D.W. OGLE. 1986. Atlas of the Virginia flora. 2nd. edition. Virginia
Boanal Associates. Farmville, VA. 135 p
HOWARD, R.A. 1988. Flora of the Lesser f aiiiles. Jamaica Plains. [Fzcvs in vol. 4:57 —64.]
HOWARD, R.A., and G.W. Staples. 1983. The modern names for Catesby’s plants. J. Arnold Arbor. a 511-546.
JOHNSTON, M.C. 1971. Revision of Colvbrina (Rhamnaceae). Brittonia 23:2 —53.
LINNAEUS, © 1753. Species plantarum. Stockholm. 2 vols.
LITTLE, E.L., Je. Checklist of United States trees (native and naturalized). U.S. Dept. Agric. Hiandh. 541. 375pp.
LONG, R.W. and O. LAKELA. 1971. A flora of tropical Florida. Univ. of Miami Press. XVil, 962 p
MANGALY, XK 1968. A cytotaxonomic study of = herbaceous species of Smilax: section Coprosmanthus. Rhodora 70:55 —82, 247 — 27
MASTERS, M.T. 1872. Passifloraceae in C. a von Martius’ Flora brasiliensis. 13:529 — 628.
NUTTALL, T. 1818. The genera of North American plants. Philadelphia. 2 vols.
REHDER, A. 1945. Carya alba proposed as nomen ambiguum. J. Arnold Arbor. 26:482 — 483.
ROULEAU, ie ie Populus balsamifera of Linnaeus not a nomen ambiguum. Rhodora 48:103 —
SARGENT, : . 1890 — 1902. The silva of North America. Boston and New York. 14 vols.
SMITH, L.B. — Bromeliaceae in the North American flora 19:61 — 228.
77. Tillandsioideae (Bromeliaceae). Fl. Neotropica 14(2):665 —
1492.
WEBSTER, G.L. 1956-1958. A ue study of the West Indian aus - Se es J. Arnold Arbor. 37:91 — is ce 1956; 38:5 ),
IDB, 709 = 315... 1997539: ie ee
ee E.T., J.M. FOGG, Jr. and H.W. WAHL. ee Atlas of the flora of Pennsyl- vania. Morris Arboretum, U. of Pennsylvania. 309 pp.
WILBUR, . “4 1980. ie lectotype of the generic name Bignonia — again. Taxon Bye Pe eam
THE CLEMATIS VIRGINIANA (RANUNCULACEAE) COMPLEX IN THE SOUTHEASTERN UNITED STATES
FREDERICK B. ESSIG Department of Biology University of South Florida Tampa, FL 33620, U.S.A.
ABSTRACT
The Clematis virginiana complex of eastern North America consists of two closely related and often confused species. The morphological, phenological, ecological and geographical characterization of these two species is clarified here, accompanied by notes on SL aii nomenclature and synonymy. Clematis apres Pursh is distinguished from the more widespread C. virginiana L. on the basis of leaves 5-foliolate co biternate as eset to 3-foliolate, and carpels 18— 35 as spoece to 40—60. Clematis catesbyana also flowers earlier in the season than C. virginiana and occupies drier habitats. Clematis catesbyana 1s restricted to several disjunct regions of the southeastern United States, while C. virginiana is widespread throughout eastern North America.
The Virgin's Bower of eastern North America consists of two closely rela- ted species that are often confused. Both are rampant vines that produce a profusion of small white flowers in the summer (fig. 1), followed by heads of long-tailed achenes in the fall (fig. 2). Clematis virginiana was described by Linnaeus in 1755 from a specimen probably collected in Pennsylvania (Essig & Jarvis 1989), and is common throughout much of eastern North America, from Quebec to Florida and westward to eastern Texas and Manitoba. Clematis catesbyana was described by Frederick Pursh in 1814, from a specimen collected by Mark Catesby in South Carolina in 1722 (fig. 3). According to Pursh, it differed from C. virginiana primarily in having biternate leaves with typically 3-lobed leaflets rather than ternate leaves with coarsely toothed leaflets (fig. 4). Specimens matching the description of C. catesbyana have subsequently been found in scattered areas throughout the southeastern U.S., from Virginia to Florida, and westward to Louisiana, Oklahoma and Kansas (fig. 5).
Since Pursh, authors have differed on whether C. catesbyana is truly dis- tinct from C. virginiana. The flowers of the two species are essentially identical in appearance, and it has not previously been clear whether the described vegetative differences correlated with geographically or ecologi-
ally distinct taxa, or were merely forms of one variable taxon. DeCandolle (1817, 1824) recognized both species, while Torrey and Gray
Siwa 14(1):49 — 68. 1990.
FIG. 1. Flowers of Clematis catesbyana
ii,
_
from central Florida.
OS
&
} 4 4 "yy % ¥ t
Cl &
FIG. 2. Clematis catesbyana from northwestern Florida, in fruit.
52
(1838 — 1840) treated C. catesbyana as a synonym of C. virginiana. Kuntze (1885), employing an extremely broad species concept, included both C. virginiana and C. catesbyana as subspecies under Clematis dioica (which was technically incorrect because the epithet virginiana has priority over dioica). Gray (1895) recognized both species, essentially on Pursh’s criteria. Small (1933) also recognized both species, and described an additional species in the complex, C. micrantha, which supposedly had smaller flowers.
Recent floristic authors have generally recognized one or the other species, without attempting to differentiate the two, implying usually that C. catesbyana represents only a morphological variant of C. virginiana. Steyermark (1963), Radford et al. (1968), and Wunderlin (1982) recognized only C. virginiana, with the latter two authors citing C. cates- byana as a synonym. Clewell (1985), on the other hand, recognized C. catesbyana as the species occurring in the Florida panhandle, but did not take into consideration specimens matching C. virginiana that occur there. He therefore did not deal with the differentiation of the two species. Keener (1975) and Keener & Dennis (1982), in the broader context of studies of the Ranunculaceae of the southeastern United States, recognized C. virginiana and C. catesbyana, but placed Small’s C. micrantha in syn- onymy under the latter. Keener’s (1975) study is the only recent work that attempts to differentiate between the two species, and provides some tenta- tive morphological criteria for separating them.
The present investigation, part of a long-term study of Clematis section Clematis (sensu Tamura 1968) worldwide, was undertaken to clarify the status of Clematis catesbyana relative to C. virginiana, and perhaps to achieve a better understanding of specific differences within the section as a whole. Throughout the section there are difficult complexes of species, and the differences between species seem at times to be minor and insignificant. Experience with this well-known complex from North America should therefore help illuminate other complexes.
MATERIALS AND METHODS
Specimens of the Clematis virginiana complex were borrowed from major herbaria throughout the eastern U.S. (A/GH, AUA, DUKE, FLAS, FSU, GA, KANU, LSU, MO, NCU, NO, NY, OKLA, PH, SMU, TENN, TEX, UARK, UNA, US, USCH, and USF). Data from herbarium sheets were entered into a computerized database using Asksam, a text-oriented database system that allows variable length fields. Label data, reproductive status, and various morphological characters were recorded. Specimens were initially sorted according to leaf character (leaflets 3 vs leaflets 5 or more), following Pursh and Keener. The database was then analyzed for
FIG. 3. Holotype (OXF) of Clematis catesbyana, collected by Mark Catesby in South Carolina.
54
correlations between morphological, ecological, phenological and geographical parameters. RESULTS
Comparative study of about 750 sheets of the Clematis virginiana complex in the southeastern U.S. revealed a strong correlation between the leaf characters described by Pursh and several previously unrecognized morphological, phenological, and ecological features, as well as with geo- graphic distribution.
Morphology: achene number
Plants with leaves 5-foliolate to biternate (C. catesbyana) consistently possess fewer than 35 carpels per flower (mostly 20 — 25), while plants with ternate leaves (C. virginiana) consistently have more than 40 carpels per flower (mostly 45—55). This is roughly twice as many carpels per flower in C. virginiana as in C. catesbyana, giving the achene heads of the former a fuller, more globose appearance than the heads of C. cateshyana (fig. 6)
A related character, achene color, was used by Keener (1975) to distin- guish between the two species. According to him, C. virginiana has achenes “light to dark-brown or greenish brown,” while C. catesbyana has achenes “reddish or purplish brown to dark blackish-purple.” This seems to be valid to a degree, but is not as clear-cut or reliable as achene number. Achene color in dried specimens varies considerably depending on ripeness and drying conditions. I found a number of specimens that could not be properly placed on the basis of this character.
Phenology
A measure of the flowering phenology of each species was obtained by treating individual specimens as data points. It was found that the two species respond differently to seasonal cues (fig. 7), with C. catesbyana flowering early in the season (early July to early August) and C. virginiana flowering later (early August to late September in the southeast). The data were plotted against latitude because, as one moves southward, flowering is progressively later in the season. Thus, within particular latitudinal belts there is little overlap in blooming period. In central Florida, for ex- ample, where natural populations and cultivated plants have been observed for several years, flowering periods of the two species are consistently separated by 2 — 3 weeks. Where there does appear to be some overlap, it is between specimens that are geographically remote from one another. Thus it appears that the opportunity for hybridization between the two species is
56
extremely limited, if it exists at all. The Atlantic coastal populations of Clematis catesbyana (see fig. 5) were not included in figure 6, because the latitudinal effect is offset, possibly because of the longer growing season along the coast. In North Carolina, for example, coastal populations flower
from late July to early September, a full month later than inland popula- tions at the same latitude, and even a little later than the Florida popula- tions. The question of hybridization with C. virginiana does not arise here, since these populations are geographically quite isolated.
Ecology and Geography
Both species are weedy, rampant vines inhabiting disturbed sites. Clematis virginiana, however, is confined to river margins and other habitats with damp to saturated soil, while Clematis catesbyana tends to occur on drier, well-drained, often calcareous sites. The latter has major populations on the Ozark Plateau, the Nashville Dome region of central Tennessee, loess bluffs along the Mississippi, Apalachicola, and Chattahoochee rivers, on shell mounds and sand dunes along the Atlantic coast, and in forested regions, often over exposed limestone, in west-central Florida (fig. 5). A few isolated populations in the Appalachians are associated with limestone outcrops. Both species are peculiarly lacking from the coastal plain of the Carolinas and Georgia, except for the narrow coastal population of C. cates- byana that extends from North Carolina to northeastern Florida. Alchough C. catesbyana is more often cited from calcareous habitats, habitat selection appears to be primarily for topography and drainage, rather than soil types or pH. Both taxa can sometimes be found over limestone substrates as well as on soils of more acid reaction, and thrive equally well when cultivated in rich, slightly acid soil.
A great many recent specimens of both species were collected along roadsides and other man-made habitats. Thus it is possible that some isola- ted populations have been spread beyond their natural range by humans in recent times. A large population of C. virginiana, for example, occurs in central Florida, in land disturbed by phosphate mining and along road sides. It most likely was introduced here recently, for it was not collected until 1976. This despite the fact that the population is traversed by State Highway 60, which had been travelled by a number of earlier botanists.
hen blooming and fruiting, the plants are very conspicuous along the road. Plants, apparently from this population, have now spread northward along Interstate 75, in low, wet roadside depressions into southern Pasco County. Clematis catesbyana, on the other hand, is apparently moving southward along the same highway in higher and drier spots, from natural populations in Hernando County into northern Pasco County.
) in the southeastern United
States.
DISCUSSION AND CONCLUSIONS
Extensive analysis of herbarium material of Clematis catesbyana and C. virginiana has shown that the two species differ significantly in morphology (carpel number, leaf dissection), in phenology, in geographical distribu- tion, and in habitat preference. Living populations, and populations represented by complete herbarium material, can be readily identified by the criteria presented here. No clear evidence of hybridization or true in- termediates has been seen. The combination of spatial and temporal separ- ation of known populations, moreover, strongly suggests that the opportu- nity for hybridization is rare if it exists at all. This isolation, despite broad- ly overlapping geographical ranges, indicates that the speciation process between Clematis catesbyana and C. virginiana is essentially complete. The disjunct distribution and greater variation of Clematis catesbyana suggests that it may be the older of the two species. These are well-defined, natural taxa and their recognition as species, as done recently by Keener (1975), is fully justified.
TAXONOMIC TREATMENT
General description (Clematis virginiana complex): Woody, deciduous to evergreen, dioecious vines, climbing by means of tendril-like petioles and petiolules. Leaves compound, thin, membranous, nearly glabrous above, coarsely toothed to entire, with sparse to dense short, simple, white hairs below; inflorescence of simple to compound, leafy to bracteate dichasia in the axils of leaves of current year's growth; axes hairy; flower buds ovoid (pistillate) or obovoid to pyriform (staminate), flowers white, sepals 6 — 14 mm long, 2—5 mm wide, linear-lanceolate to long-obovate; sparsely hairy above, more thickly so below, and densely hairy on margins, hairs fine, white; staminate flowers with ca. 30 to over 50 stamens, these somewhat shorter than the sepals, filaments flat, nearly as wide as the anthers, anthers ellipsoid, ca. 1 mm long, pistillodes rudimentary, inconspicuous, hidden in the thick hairs of the receptacle or often lacking altogether; pistillate flowers with numerous staminodes, similar to fertile stamens but shorter, sterile anthers rudimentary to nearly normal in appearance, lacking pollen; carpels numerous, ovary swollen, short-hairy, style elongate, nearly equal- ling the sepals, densely hirsute, stigma simple, curved-clavate; achenes lens-shaped, light to dark brown or reddish black, sometimes with a dis- tinct, thickened, lighter rim, sparsely short-hairy, persistent style 2.5—3.5 cm long, covered with long, white hairs.
These species are adapted to the mesic conditions of eastern North America, and are distinguished most readily from related western species by their large and membranous leaves. Clematis ligusticifolia Nuttall, for
FIG. 6. Achene heads of Clematis catesbyana (A) and C. virginiana (B).
60 example, differs in having slightly succulent or coriaceous, 5- to 7-pinnate leaves with stomata on both surfaces, and in other subtle characters (Keener
O73
DiaGnostic Key ro CLEMATIS CATESBYANA AND C. VIRGINIANA
Mature vegetative leaves 5-foliolate to biternate, leaflets usually 3-lobed, few-toothed, eae about 1/3 the length of the entire leaf, pistillate flowers with 18— 35 carpels; flowering e ak to mid-August; well- drained, ee ee sites, southeastern U.S. 2. ......0....... C. catesbyana Mature vegetative leaves ternate, leaflets dni coamely dentate, petiole 1/2 the length of the entire leaf or more; pistillate flowers with 40 — 60 carpels; flowering July to August in the northeastern U.S., early August to late September in the southeastern U.S.; low, moist areas, riverbanks,
—
roadside ditches, throughout eastern U.S. and Canada ............ C. virginiana
CLEMATIS © Sree Pursh, Fl. Amer. Sept. 2:736. 1814. — Tver: SOUTH CAROLINA, Catesby 1135 (HOLOTYPE: OXE photo!, see figure 3). Clematis divica a catesbyana var. variabilis Kuntze, Verh. Bot. Vereins Prov. Brandenburg 26:103. 1885, in ae Clematis virginiana vat. catesbyana (Pursh) Britton in Britton and Brown, HL ~N. US. 2:67. 1897.
In the este to this name, Pursh cited a Catesby specimen in the Sherard Herbarium at Oxford (“v.s. herb Sherard”). A single specimen attributable to this species (fig. 3) has been located in the Sherard Herbarium. The specimen, numbered 1135, was collected by Catesby in Carolina in 1722 and matches Pursh’s description well. It therefore can be considered the holotype.
CLEMATIS CORDATA Pursh, Fl. Amer. Sept. 2:384. 1814. — Tyee: WEST VIRGINIA. Summers Co.: on the ascent of Keeny’s Knob, ae the precipice called the Claypinch, Parsh s.n. 1806 Garner designated here: PH!). Clematis dioica subsp. cordata (Pursh) Kuntze, Verh. . Vereins ee 26103;
Clematis cordata was published at the same time as C. catesbyana, and has traditionally been treated as a synonym of C. virginiana. However, Pursh described the leaves as 5-foliolate, which suggests that it should properly be placed under C. catesbyana.Pursh indicated in the protologue only that he had seen living material of this species in the high mountains of Virginia. No type was designated. However, a Pursh specimen from the mountains of West Virginia (Keeny’s Knob, Summers Co.), matching the type description, and most likely collected in 1806 —07 (preceding the publication of the name in 1814), has been located at PH. It is designated here as the lectotype. Most of the leaflets have fallen off of the specimen but from the elongated leaf rachis and the presence of scars, it is clear that the specimen was at least 5- foliolate. Similar material has been recently
<s C): = 8 : 5 5 | O O | Q | ee e er i ‘ ‘2: O | GO @P qd <« S o Bo % 8 od 4 : Oo” Be O° km Qa Oo,0 <d<4 <4 _ @ Se a a ob i O° c om : 8 x < ; 3 o i < Te d< = < “ - <« q < a < 4 S a < 0) < P E qs < -_— P ile i 4 i 4 : 2 a ee ee | | (haere Coes oO ° 2 Se A
FIG. 7. Blooming time of Clematis catesbyana (solid triangles) and C. virginiana (circles).
collected from limestone outcrops in nearby Giles County, Virginia, which is clearly identifiable as C. cateshyana. There is no reason to consider C. cordata as a distinct taxon.
62
aac MIC kas see Man. S.E. Fl., p. 525. 1933. — Tver: ndo 5 Chooesahaitee Poa mee S of Brookesville, ela aa s Seen ao (LecroTYPE, designated here: NY!; isorype: GH In the appendix to his Manual (p. 1504), Small (1933) cites two specimens under this name as follows: “Type, Devil's Punchbowl, w. of Brooksville, Fla., Small, No. 11337; for fr. Choocochattee hammock, s. of Brooksville, Fla., Small, No. 10602, in herb N.Y.B.G”. Of these two syn- types, only number 10602 is annotated by Small as the type, and this is the specimen that contains the small flowers (not fruit) that figure prominently in his description. Sheet number 11337 in fact is a sterile specimen. There- fore, I designate Small et al. 10602 as lectotype of Clematis micrantha. Clematis micrantha was defined on the basis of its smaller flowers. Flowers on the type are indeed unusually small, with sepals only 5 — 6.5 mm long. In the newly collected material from the type locality (Brookesville area, Hernando County, Florida), however, sepals range from 6.5 to 10.5 mm long, well within the range of C. catesbyana as a whole. Also according to Small, plants are closely fine-pubescent in C. micrantha and minutely pub- escent or glabrate in C. catesbyana. I can see no difference when a wide range of material is examined. All specimens have fine, white hairs on stems and leaves that range from sparse to thick even on individual specimens. Small therefore based his species on a specimen that was evidently atypical of its population. The most distinctive feature of the Brookesville plants is the strong degree of leaf dissection. The leaves are biternate or even further divided into 11 or 15 small leaflets. This degree of dissection gradually diminishes northward, however, with leaves 5-foliolate to biternate in north Florida and mostly 5-foliolate in the Ozark region. Representative specimens examined — CLEMATIS CATESBYANA: (complete list of ex- siccate available from the author) ALABAMA. Clark Co.: borders of woods in rich soil, 1859, Denny 4 (UNA). KANSAS. Benton Co.: Ozark Plateau, Boston Mtns., generally wooded area near Bella Vista, 5 mi N of Bentonville, elev. 1000— 1200 ft, 22 Sep 1928, Demaree 5.n. (UARK). Carroll Co.: Ozark Plateau, Province of White River Hills, wooded NE facing slope along White River at Catron Bend, 5 mi NW of Eureka Springs, elev. 900 — 1000 fr, Rockwood, 27 — 30 Jul 1953, Leonard & May 6 (UARK). Cross Co.: Crowley's Ridge, Levesque, elev. 300 ft, 24 Jul 1939, ae 19587 (MO). Izard Co.: pe road from Croker to Guion near White River, 23 Jul 1970, Thomas 20192 (FLAS, TENN). Logan Co.: Mt. Magazine, plateau surface at ca 2800 ft, 17 Sep 1967, Tucker a (NCU). Newton Co.: bank of Clark Creek and park entrance, occasional, 14 Jul 1974, Thompson s.n. (UARK). Washington Co.: in Fayetteville on Dickson St., in a ditch adjacent to the street on a Sequoyah, 10 Jun 1975, Davis 891 (UARK FLORIDA. Alachua Co.: Paynes Prairie State Preserve, i of Gainesville, N side of Alachua anh twining up trees, 24 Oct 1981, Easterday 755 (FLAS). Citrus Co.: limestone outcrops, 3 mi SW of Pineola, 1 Aug 1948, Ford 2295a (TENN). Dixie Co.: S of Old Town, 11 Aug 1937, West & Arnold s.n. (FLAS); swamp at Suwanee, 5 Sep 1957, Godfrey
63
56042 (FSU). Duval Co.: Fort George Island, in delta of St. aan River, E of Jacksonville,
abundant in roadside vegetation along E side of island, growing in crushed shell, 26 Nov 1987, Essig 871126 — 1 (USF). Franklin Co.: Apalachi Chapman s.n. (MO). Gadsden Co.: near ground level, old wood stem 2.5 cm diam, at 6 m above ground 1.5 cm diam,
shrouding crown of willow tree, borders of floodplain ape vea Apalachicola R., by US 90 bridge, Chattahoochee, 21 Sep 1981, Godfrey 79145 (FLAS, FSU). Hernando Co.: abundant in old limestone quarry along CR 491, just N of jct. with US 98, 26 Sep 1986, eae 860926-1 (USF). Jackson Co.: climbing in trees along n-s paved rd. at Marianna ns State Park, 22 Jun 1960, Mitchell 447 (FSU). Lake Co.: vic. Eustis Lake, 16 — 25 re 1894, Nash 1731 (MO). Leon Co.: growing in roadside shrubbery along Hwy 90 at Sun Ray Rd., 1 Sep 1987, Essig & Hansen 870901-1 (USF). Levy Co.: on roadside vegeta- tion to 3 m high at Magnolia, 22 Sep 1959, Cooley et al. 7182 (FSU, NCU, USF). Liberty Co.: floodplain woodland, Apalachicola R., E of Sneads, 16 Aug 1982, Godfrey 79924 (FSU). Marion Co.: calcareous woodland near the Silver R., on Dupont property, 3 Oct 1984, Godfrey 81651 (FSU). Pasco Co.: on fence beside I-75, W side, ca 0.25 mi S of CR 41, 27 Aug 1987, Essig 870827-1 (USF). Polk Co.: at edge of swamp forest, dirt extension of Hinson AvE, near Lake Marion, E of Haines City, 4 Oct 1987, Essig 87 1004-1 (USE).
GEORGIA. Decatur Co.: on edges of mixed woodland by the office of Resource Mana- ger, Lake Seminole, 9 Sep 1979, Godfrey 77204 (FSU, GA). Early Co.: bank of Chatta- hochee R. at Sheffield’s Landing, 14 Aug 1901, Harper 1222 (MO). Liberty Co.: St. Catherine's Island, N end, edge of woods beside the housing compound, 24 Aug 1983, Jones, et al. Sa (GA). Seminole Co.: bank of Chattahoochee R. at Butler, 25 Jul 1947, Thorne 5663 (G
SAS. Br ike Co.: 6 mi E of Baxter Springs, near MO border, rocky wooded hillside, Ozark region, growing on thicket of Cornus asperifolia, 3 Jul 1948, McGregor 1937 (KANU).
KENTUCKY. Warren Co.: along roadsides and ditches on Jenkins Rd. ca 1 mi south of U.S. Lock & Dam £1 on Barren R., ca 5 mi NNW of Bowling Green, 20 Jul 1970, Nicely & Gough 3069 (NCU).
LOUISIANA. St. Helena Parish: abundant in open shrubby area ca 1 mi W of Chipola, assoc. with Ifex and Cornus, 30 Jul 1971, Allen 1296 (DUKE, LSU). West Feliciana Parish: ca | mi from post office of Plettenberg, trailing on trees along logging road, 22 Aug 1938, Correll & Correll enn (DUKE, LSU).
MISSOURI. Barry Co.: roadside, NE facing cherty slope, Hwy 112 east ame R27W, W1/2, sec. 27), 16 Jul 1979, aed 532 (UARK). Christian Co.: Low thickets along east fork of Bull Creek, 3 mi SW of Chadwick, 8 Jul 1937, pee ale a (MO); Rocky Hills, 28 Sep 1905, Bush 3479 (M 5) Douglas Co.: beside White R. and
O 14 ac Twin Bridges, 28 Jul 1969, Thomas 15817 (TENN). Greene Co.: 10 mi SE of Springfield, James River & MO Hwy 125, 25 Jul 1975, Stalker & Nelson Fe (NCU). Howell Co.: rocky wooded slope bordering Indian Creek, 3.5 mi W of willow Springs, 18
Jul 1982, Summers 1079 (MO). Shannon Co.: Montier, common, 29 Aug 1894, Bush 1 (MQ). Stone Co.: thickets along rocky creek, near Cle 25 Sep 1923, Palmer 23881 MO). Taney Co. mon in woods, Swan, 4 Jun 1899, Bush 99 (MO).
MISSISSIPPI Union Co.: State Hwy 346, roadisde 10 mi E of New Albany, 18 Jul 1966, Temple 3504 (GA, NCU). Warren Co.: edge of wooded se bluffs facing delea region, 2 mi N of Redwood, 12 Jul 1955, Ray 4910 (NCU). Wil on Co.: roadside, sandy soils, Smith eee ca 5.5 mi WNW of Woodville, 22 Jul ee & Clark 19800 (NCU). Yaz : 6 mi SE of Yazoo eee loessial soil along creek, common, high-climbing, 29 i 1959, McDaniel 1286 (U ORTH CAROLINA. Brunswick Co.: sane ae Summer 1925, Blomquist 3643
64
(DUKE). Carteret Co.: very abundant as liana in low dense live oak-juniper forest on Shackleford Banks, 7 Aug 1962, Anderson 386 (DUKE, FSU). Currituck Co.: thicket, 3 mi N of Waterlily, 31 Jul 1958, Ahbles G Duke 48210 (NCU). Dare Co.: depression between dunes of Duck, 28 Aug 1952, Radford 660 (NCU). Onslow Co.: Bear Island or Hammock Beach State Park at the western quarter of the island, abundant spreading spraw- ling vine forming dense cover on stabilized dunes at the western end of island, 25 Aug 1967, Wilbur 9590 (DUKE).
OKLAHOMA. Cherokee Co.: open woods of creek valley, 22.1 mi NE of Tahlequah on State 10, 29 Jul 1951, Wallis 860 (OKLA).
SOUTH CAROLINA. Beaufort Co.: very abundant in sunny disturbed areas and road- sides through abandoned fields, central Callawassie Island, 13 Oct 1981, Aulbach-Smith 2036 (USCH)
TENNESSEE. Cannon Co.: in limestone valley, fencerow on Rt. 145, 3 mi N of Wood- bury, 29 Jul 1958, Ellis 249-E (TENN). Carter Co.: on roadside in open place, toll rd. to Roan Met. at 3800 ft, 26 Jul 1934, Brown 100 (DUKE). Cheatham Co.: Ellis 196-E (TENN). Clay Co.: | mi N of Clay Co., line on Hwy 53, roadside, 7 Jul 1958, E//is 24736 (TENN). Coffee Co.: edge of woods, escarpment area, 28 Jun 1955, DeSe/m 593 (TENN Davidson Co.: on fence by Mountain View Rd., N of Murphreesboro Rd., toward Percy, Priest Lake, 14 Aug 1968, Kra/ 32340 (NCU). Giles Co.: NE of Pulaski, roadside on limestone, 13 Jul 1948, Sharp et al. 9796 (TENN). Grundy Co.: borders of hardwood forest on mountain slope, 2.3 mi N of Monteagle, ?4 Aug 1970, se 69759 (FSU, NCU). Jackson Co.: 0.25 mi ane Hwy 85 on Haydensville Rd., 7 Jul 1958, E//is ene (TENN). Macon Co.: bank of Long Cr., 9 Jul 1958, Ellis 24457 (TENN). Maury Co.: Green’s Mill Rd., ca 1 mi from US 31, 27 Jul 1957, Chappell 5.n. (TENN). Moore Co.: moist soil etore SE of Lynchburg, 7 Aug 1947, si é “i 5685 (TENN). Rutherford : growing over limestone on sides of road, betwee . 231 and Christiana, 31 Jul ner Ellis 326-E (TENN). Williamson Co.: in ener bottom, 1.5 mi SE of McDaniel, 6 Aug 1958, E//is 322-E (TENN). Wilson Co.: limestone — 0.5 mi N of 70 N, on old roadbed of Hwy 109, 23 Jul 1958, E//is 24771 (TEN
VIRGINIA. Giles Co.: 1.4 mi W of Mt. Lake Hotel on Va 613, su 1965, Scheer s.n. (NCU).
2
CLEMATIS VIRGINIANA L., Cent. I Pl., p. 15. 1755. — Tver: unnumbered specimen se TOTYPE: UPS, photo!), see Essig - Jarvis 1989. Clematis dioica subsp. virginiana (L.) Kuntze, Verh. Bot. Vereins Brandenburg 26:102. 1885. Clematis virginiana vat. genuina Kuntze, Rev. Gen. 1:2. 91, nom. aga
CLEMATIS CANADENSIS Miller, Gard. Dict. ed. 8, Clematis No. 5 1768. — Tyee: not designated.
2:
Miller described this species as having ternate leaves with cordate, toothed leaflets, which places it with C. virginiana. He cited ” Clematitis canadensis latifolia & triphylla. Sar.”, most likely referring to the French Canadian botanist Michel Sarrazin, who collected in southern Quebec between 1697 and 1734. There may be a specimen matching the descrip- tion at Paris.
65
ae HOLOSERICEA Pursh, Fl. Amer. Sept. 2:384. 1814. — Typr: . Herb. Walter.”; not seen.
Pursh described this species as ternate, and holosericeous-pubescent, with small white flowers. The vestiture of the foliage is of no taxonomic consequence, and the species clearly falls under C. virginiana. Pursh cites a specimen in the Walter Herbarium (BM) from Carolina. Walter's specimens are mounted several to a page. Two specimens on page 34 of this collection, seen in a photograph only, possibly match Pursh’s description. Most likely, one of these could be designated the lectotype, but I refrain from doing so until I have the opportunity to examine the specimens.
CLEMATIS MISSOURIENSIS pea: in Britton, Man. Fl. N. U.S. 1901,
part. — Tyee: Webber s.n. Sep 1886, Lincoln, Nebraska (LECTOTYPE, esi here: NY!). Clana virginiana var. missourtensis (Rydberg) Palmer & Steye Ann. Missouri Bot. Gard. 22:542. 1935. Clematis virginiana forma missouriensis
(Rydberg) eet Rhodora 39:309. 1937.
Rydberg mentioned only that the type came from Lincoln, Nebraska. The Webber specimen at NY is annotated as the type by an unknown hand. It is in fact the only known specimen, matching the description and com- ing from the type locality, that would have been available to Rydberg at New York at the time. Therefore, I designate this specimen as the lecto- type.
The species was distinguished on the basis of the undersides of leaflets being densely hirsute, and the achenes lacking the characteristic thickened rim of Clematis virginiana. The achene character, cited most recently by Gleason and Cronquist (1963), was discounted by both Fernald (1937) and Steyermark (1963). The distinction disappears when many specimens are examined. Specimens with densely hirsute leaves are particularly common from western Missouri northward into Minnesota and Ontario, but can be found sporadically throughout the northeastern United States. Also, many specimens with hairy leaves from southwestern Missouri, cited as Clematis virginiana forma missouriensis by Steyermark (1963) actually belong to Clematis catesbyana. Formal recognition of this taxon at any level is thus likely to lead to confusion and therefore should be avoided.
Representative specimens examined — Clematis virginiana: (complete list of exsiccate available ae the author).
ALA . Clarke Co.: Oak-Pine woodland, 6 mi S of Thomasville School, 5 Sep ren ee i 1 26 (NY). Monroe Co.: Haines Island, Rae limestone ridge and ravines along the Alabama River, 3 Sep 1985, Diamond 1742 (AUA).
ARKANSAS. Marion Co.: Buffalo Pt., ca 14 mi S of Yellville on Hwy 14 and off on Rd. 268, 6 Aug 1975, Smith 219 (UARK). Sine Co.: 2 mi W of Benton, on gravel shores of
66
river, 6 Sep 1942, Tolstead & Demaree (NEB). Sevier Co.: 1 mi W of Lorksburg in small ee 20 Oct 1932, Demaree ene (MO, NY, US).
FLORIDA. Escambia Co.: near Molino, along W side of L&N RR tracks betwee Pretty Branch and its south fork, 9 Sep 1980, Burkhalter 7190 (FLAS, FSU). Hillsborough Co.: Edward Medard State Park, Turkey Cr. Rd., ca 5 miS of Plant City, 25 Sep 1987, Essig 870925-5 (USF). Jefferson Co.: at and just niche borders of hardwood hammock at Wacissa Spring, 6 Sep 1982, Godfrey 79947 (FLAS, ay, Pasco Co.: in roadside vegeta-
on, low, mesic areas along I-75, ca 3 mi S of Hwy 54, 20 Oct 1987, Essig & Hansen 571000. 2 (USF). Polk Co.: growing in low moist area, ete the N side of FL 630, ca 1.5 mi W of Ft Meade, 15 Oct 1977, Shuey 1998 (USF).
GEORGIA. Harris Co.: low, sunny roadside, rocky soil <i GA 190 at Dowdell’s Knob on Pine MtN, 25 Aug 1971, Jones 21361 (GA, NCU). Morgan Co.: Hard Labor Creek State Park, swampy area beside main highway, 24 Aug 1978, Hill 668 (GA, NCU). Walker Co.: ne Mtn. Wildlife Management Area, NE of Harrisburg Gulf, 18 Aug
mo Coile et al. 2285 (GA).
AS. Douglas Co.: 2 mi E of Lecompton, Kansas ee Bluff, growing over << 20 Sep 1975, McG resu 28301 (KANU). Shawnee Co.: | mi E of Topeka, Kansas River Bluff, growing over thicket, 20 Sep 1975, McGregor 28306 (KANU).
KENTUCKY. Bracken Co.: rare, in weedy field at Meldahl Dam, ca 3 mi SE of Foster on Rt. 8, 24 Aug 1983, Buddell 2318 (NCU). Edmonson Co.: ca 1.5 mi NW of Pleasant Grove Church, Little Sally Branch, beside stream, 3 Aug 1968, Elmore 367 (NCU). McCreary Co.: Daniel ae ae oo Beaver Cr., 4.5 mi N of jet. Funston Ridge Planta- tion Rd. & Beaver Cr. Rd., 1972, Browne & Browne 72H28.2 (NCU). Todd Co.: roadside ee in area of Bees af 0.5 mi from Guthrie, 8 Oct 1949, Shanks et al. 14335 (TENN).
(coniae Caddo Parish: common in open woods off Ellerbe Rd, 9 mi S of Shreve- port, 17 Sep 1977, MacRoberts & MacRoberts 269 (NO). Oachita Parish: woods near Hogpen Rd, Lapine Rd., 3 mi S of LA 34, 10 Sep 1968, Holler 78 (NCU). Plaquemines Parish: margin of an Road C, eastern extension, 20 Sep 1978, Fleming 411 (NCU). West Feliciana Parish: roadside, 5 mi E of Tunica center on Rt. 66, 31 Oct 1976, Paulson sn. (LSU). Washington Parish: on LA 483, 2.1 mi W of Warnerton, 29 Nov 1958, Stone
! 0.
MISSOURI. Butler Co.: swamps, ° Oct 1905, Bush 3710 (MO). Dallas Co.: upper limestone thickets along Niangua R.., mi upstream from mouth of Douisenburg Cr., 5 Aug 1937, Steyermark 24261 (MO). os nt Co.: low thickets along N prong of Maramec R.., between Stone Hill and Indian Trail State Park, 4 Aug 1936, Steyermark 12472 (MO). Jackson Co.:, 5 Oct 1888, Bush 5.x. (MO). Marion Co.: Bear Cr. Bluffs, Oakwood, 10
MO).
MISSISSIPPI. Amite Co.: common along roadside and ditches, 15 mi N of Liberty, 20 Sep 1973, Westbrook s.n. (UARK). Forrest Co.: bottomland hardwoods along Leaf R. at McCallum, SE of feat 16 Sep 1966, Jones et al. ae (NCU). Jefferson Co.: weedy roadsides along US 61, 0.7 mi N of South Fork Cole , 3.3 mi N of Stanton, 7 Sep
1980, Pruski 1893 (TEX); ree disturbed me along on benweeii Reedy Cr. & Bogue Homo Cr., 1 Sep 1978, Morgan 1167 (FSU). Montgomery Co.: right-of-way and rich deciduous eee along LA Hwy 404, 0.9 mi W of Ava, 19 Aug 1967, Temple 6534 (GA). Tishomingo Co.: Hwy 72 at Yellow Cr., moist power line, 26 Sep 1974, Coleman 48426 (TENN). Wilkinson Co.: roadside, full sun, loess bluffs, ca 8 mi via air NW of Woodville on on Beaver Cr. Rd., between Lessley & Lanhart, 15 Sep 1970, Jones et al. 20143 (GA).
NORTH CAROLINA. Chatham Co.: seasonal stream bed, 1.2 mi E of NC 1008 on NC
67
1743, 31 Oct 1981, Jones K-8 (NCU). Cherokee Co.: Bog, near Grape Cr., 6 mi NW of Murphy, 31 Aug 1956, Radford 17625 (NCU). Granville Co.: creek bank, 3.2 mt S of Goshen, 28 Sep 1956, Ahles & Leisner 20118 (FLAS, NCU). Lincoln Co.: low pasture by Howard’s Cr., 0.6 mi E of Cat Square, 10 Sep 1958, Bell 15300 (NCU). Madison Co.: boggy pasture, 7.4 mi N of jet. with U.S 19 and 23 on 23, north of Mars Hill, 14 Jul 1958, Ables & Duke 46326 (NCU).
OKLAHOMA. Cherokee Co: open roadside of Illinois River Valley, 2.8 mi NE of Tahlequah on State 10, 30 Aug 1950, Wallis 162 (OKLA).
SOUTH CAROLINA. Barnwell Co.: King Cr., Savannah R. Plant, 5 Oct 1953, Kelley & Batson s.n. (USCH). Lancaster Co.: deciduous forest, NE side of Flat Cr., 0.1 —0.6 mi SE of CR 37, 2 Sep 1961, Walliamson & Ables 2001 (NCU). Richland Co.: rocky, open floodplain area along the Saluda River, under the I- 126 overpass, 2 Oct 1976, Aulbach 763 (USCH). Oconee Co.: N-facing wooded slopes of hill, 1.25 mi NW of Salem, 10.5 mi NE of Walhalla, 28 me 1950, Duncan 11892 (GA). McCormick Co.: near Troy, 20 Aug 1921, Davis s.n. (M
TENNESSEE. ae Co.: calcareous banks, borders of wooded slope, 2.5 mi S of Shelbyville, 30 Aug 1958, Godfrey 57528 (FSU). Carter Co.: deciduous forest, Round Bald Mtn., N slope, rocky spring below grass bald, 6 Aug 1956, Ramseur 1386 (NCU). Cheatham Co.: bank of Harpeth R. on Hwy 70, 15 Jul 1958, E//is 24366 (TENN). Clay Co.: jct. Hwy 53 & Dale Hollow Rd., N of Celina, 6 Jul 1958, Ellis 24730 (TENN). Coffee Co.: Morton Lake, growing on trees and shrubs, 27 Nov 1980, Terry 137 (AUA). Johnson Co.: edge of deciduous forest slope, bank of Watauga Lake on Hwy 67, S side of Butler, 3 Oct 1967, Mahler & Mahler 4590 (NCU). Marion Co.: spray-zone slope at Foster Falls, Fiery Gizzard Gorges, 24 Aug 1964, Clark 1201 (NCU). Obion Co.: Reelfoot (Lake) Wildlife Refuge Area, Walnut Log Division, Blue Basin, 18 Sep 1982, Utech et al. 82-480 (NCU). Robertson Co.: roadside 3 mi W of Greenbrier, 24 Jun 1958, Ellis 347-E (TENN). Polk Co.: along RR S of Hiwassee x at McFarlands in Cherokee NE 3 Sep 1970, Odenwelder & Bowers 45577 (TENN). Tipton Co.: beside gravel road at foot of Chickasaw Bluff No. 2, 1.3 mi N of jct. with Herring Hill & River Bluff RdS, 4 Oct 1968, Warrington 406.3 (NCU). Union Co.: edge of corn field by river, Island-E Norris Lake, 18 Sep 1934, eae s.n, (TENN).
. Harrison Co.: edge of Saag forest, ca 3.5 mi NW of Karnack, 15 Sep 1964,
a 30155 (LL, NY). Jasper Co.: 1.5 mi NW of jct. US 190/63, on 63 at cag crossing, SE side, 14 Oct 198x, Gites Sun. ES San Augustine Co.: clim trees and shrubs, edge of swamp woods, ca | mi S of San Augustine, 14 Sep 1968, Correll 36560 (FSU, LL).
oe Buchanan Co.: in Grundy, « on WA 83, 18 May 1968, James 9976 (NCU). Frederick Co.: 2.6 mi SE of jct. CR 615 & US 50, upland woods, 21 Jun 1968, James 10844 (NCU). Giles Co.: jct. VA 604 and 110, along margin of Sinking Cr, weedy roadside & creek margin, 20 Sep 1975, Rich 72 (GA). Henrico Co.: edge of marsh W of Elko Station, 3 Sep 1967, Harvill 17369 (NCU). Lee Co.: edge of old field near Wilderness Rd. campground, elev. 1300 ft, 28 Aug 1974, Hinckle 49612 (TENN). Prince William Co.: in shrubs on edge of swamp and on edge of open field of Marumsco Acres Lake Recrea- tion Area, 8 Aug 1981, Keyser 397 (FLAS).
ACKNOWLEDGEMENTS
I would like to thank the many people who assisted in this project, in- cluding S. K. Marner (OXF) and Roy Vickery (BM) for sending
68
photographs of type material, Richard Wunderlin, Bruce Hansen, Carl Keener, Nancy Moreno, and James Hardin for reading and commenting on the manuscript, and the curators of all the herbaria who loaned specimens for the study.
REFERENCES CANDOLLE, A. P. de. 1817. Regni vegetabilis systema naturale, Vol. ris. CANDOLLE, A. P. de. 1824. Prodromus systematis naturalis regni moe Vol. 1. Paris. CLEWELL, A. E 1985. Guide to the vascular plants of the Florida Panhandle. Gainesville: Florida State University Pre ESSIG, EB. and C. E. JARVIS 1989. Lectotypification of Clematis virginiana L. Taxon 38:
271277. FERNALD, M. L. 1937. Nomenclatural transfers and new varieties and forms.Rhodora
GRAY, A. 1895. Synoptical flora of North America 1(1):4. New York. KEENER, C.S. 1975. Studies in the Ranunculaceae of the southeastern United States. II. Clematis L. Sida 6(1):33 — 47.
KEENER, C.S. and W. M. DENNIS. 1982. The aa classification of Cee (Ranunculaceae) in temperate North America north of Mexico. Taxon 31:37 — 44. KUNTZE, O. see eu der gattung aie: Verh. Bot. Vereins Prov.
Sea 26:1
RADFOR _E. i . AHLES, and C. R. BELL. 1968. cea of the vascular flora of the en Chapel Hill: University of North Carolina SMALL, J. K. 1933. Manual of the southeastern flora. New The ig
STEYERMARK, J. 1963. Flora of Missouri. Ames: Iowa State Univers
TAMURA, M. 1968. Morphology, ecology and Piyplozeny of the eae VI. Sci. Rep. S. Coll. N. Coll. Osaka Univ. 16:2 2
ene and A. GRAY. 1838 — 1840. hee of North America. New York: Wiley and
ao ne R. P. 1982. Guide to the vascular plants of central Florida. Gainesville: University Presses of Florida.
CYPERUS ENTRERIANUS (CYPERACEAE), AN OVERLOOKED SPECIES IN TEMPERATE NORTH AMERICA
RICHARD CARTER Herbarium, Department of Biology ‘aldosta State College Valdosta, GA 31098, U.S.A
ABSTRACT Cyperus entrerianus Bockeler, an apparently recent introduction from South America or Mexico, is reported in temperate North America. It is presently known from 20 sites in five states in eee United States. Its distribution and ecology are discussed, and dif- ferences between it and closely related Cyperus /uzulae (L.) Retz. are detailed. A key 1s given to distinguish Cyperus entrerianus from other members of section Lazuloider, which occur in
North America.
RESUMEN Cyperus entrerianus Bockeler es registrada para la zona templada de America del Norte; esta especie aparentemente es de reciente introduccion de Mexico o America del Sur. Esta expecie se conoce actualmente de 20 localidades en cinco estados en el sureste de los Estados Unidos. Se ee aqui su distribucion eae y se detallan las diferencias entre esta y la especie mas afin Cyperus /uzulae (L.) Retz. Se incluye una clave taxonomica para distinguir Cyperus entrerianus de los otros ae de la seccion Luzuloider, que ocurren en America
del Norte.
INTRODUCTION
While conducting field work in southern Georgia during 1987, an un- familiar species of Cyperus Section Luzuloidei was located in Ware County. | tentatively identified it as Cyperus ochraceus Vahl var. excelstor Kikenthal? and sent a duplicate to Robert Kral (VDB) who identified it as C. surinamensis Rottb. Intensive field work during 1988 and 1989 revealed 12 additional sites from southern Georgia and western Florida to southeastern Texas. Additional specimens from Florida, Louisiana, and Texas have been located at FSU, IBE, and VDB.
This sedge has now been identified as Cyperus entrerianus Bockeler, which is primarily a temperate South American species (Kiikenthal 1936). The epithet “entrerianus” is derived from Entre Rios, the name of an historical- ly disputed area presently in Argentina and the type locality of C. en- trerianus (Bockeler 1878). Kikkenthal (1936) and Pedersen (1968) recognized C. entrerianus as a distinct species, and Kiikenthal (1936) placed it into section Luzuloidei of Cyperus. Also, this taxon was treated as a variety
Sipa 14(1):69—77. 1990.
70
of C. /uzulae by Barros (1938). Denton (1978), in a taxonomic treatment of the “Luzulae group” of Cyperus, placed the name into the synonymy of Cyperus luzulae(L.) Retz., but, in discussion, referred to “ ‘entrerianus’ and ‘luzulae’ modes of variation”. Furthermore, Denton (1978) cited specimens of C. /uzulae from Florida, Missouri, Oklahoma, and Texas, which I have not seen; it is unclear whether these are C, /uzulae or C. entrerianus. How- ever, C. /uzulae is primarily a tropical species, and it seems doubtful that it has or will become established in other than subtropical regions of the United States, such as extreme southern Texas or perhaps southern Florida.
TAXONOMY
Cyperus entrerianus is a stout rhizomatous perennial, which may be as much as 120 cm tall. It has bicarinate, dorsally grooved scales and florets characterized by a single stamen. It clearly belongs to section Luzuloidei as circumscribed by Kiikenthal (1936) and is closely related to Cyperus luzulae. | have observed Cyperus luzulae in the field in Peru [McDaniel 23640, Rimachi, Carter (IBE, VDB)} and in Dominican Republic {Carter 5201, 5220 (MO, VSC)}, and I believe it and Cyperus entrerianus are specifically distinct.
There are a number of differences between these two taxa. Cyperus en- trerzanus 1s a larger plant than C. /wzulae. The leaf bases of C. entrerianus are heavily black-pigmented, and their fibrous remains are persistent. Its base, unlike that of C. /uzulae, is deeply set in the substrate and has thick rhizomes with short internodes. Spikes of C. entrerianus are conspicuously compound and composed of one to several globose units, while those of C. luzulae generally are dense and conical and appear simple. The spikelets of C. entrertanus are more loosely arranged and have 16 — 32 pale greenish scales, while those of C. /vzulae are tighter and have 12 — 16 whitish to tawny scales.
Certain specimens (e.g., Carter 6296, 7319, 7447, 8040, 8093, and 8102) from southeastern United States exhibit sparsely scabrid culms and, thus, will key to Cyperus surinamensis with most conventional keys (e.g. Godfrey and Wooten 1979). The culm teeth are mostly retrorsely oriented as they are in C. surinamensis. This characteristic has not been previously noted in C. entrerianus in the literature. Also, the number of fertile scales per spikelet in C. entrerianus specimens from United States seems to be greater than in specimens from South America and falls within the range for C. surinamensis (see table 1). Furthermore, specimens of C. entrerianus from southeastern United States apparently are more robust than ones from South America, and in the United States Cyperus entrerianus appears to be an aggressive weed and is often locally abundant and dominates its disturbed
Tasre 1. Morphological comparison of Cyperus entrerianus with C. luzulae and C. surinamensis.
CHARACTERS
TAXA
C. entrerianus
C. luzulae
C. surinamensis
HEIGHT RHIZOME
LEAF BASE
SPIKE DENSITY SHAPE
SPIKELET
SCALE POSTURE
rhizomes with short
internodes blackish purple;
persistent & u
loose, thus individual spikelets distinct compound, units globose
16-32
1.8—2.1 mm
pale green
divergent ca 30°
cespitose, rhizomes
not at all evident
brown to reddish brown; neither persistent nor
fibrous
tight, thus individual spikelet
not distinct
appearing simple, nical
12—16 1.4—1.8 mm
whitish to tawny
divergent ca 30°
cespitose, rhizomes at all evident
brown to reddish brown, neither persistent nor fibrous
loose, thus individual spikelets distinct compound, somewhat flattened
18 — 48 1.6—2.3 mm
golden yellow to stramineous
divergent ca 45°
habitat. The
characteristics suggest introgression between C.,
aggressive nature and robust size of plants of C. entrerianus from southeastern United States might well be due to heterosis. These
entrerianus and C.
surinamensis, which frequently occur together in southeastern United States. A morphological comparison of Cyperus entrerianus, C. luzulae, and C. surinamensis is given in table |. Following is a key by which the North American species of section Luzuloider may be identified.
Kry to Cyperus SECTION LuzuLoiDE! IN NORTH AMERICA
1. Culm triquetrous, angles sharp, usually flattening when pressed and dried.
Achene abo ceeding inflorescence more than
ut 1/2— 2/3 as long as scale; inflorescence open; bracts ex- 7 Wane ae Aerts oon rca C. virens Michx.
Achene at least 3/4 as long as scale; inflorescence ce bracts ex-
ceeding inflorescence 4 or fewer
. drummondit T. & =C. robustus sae
1.
Culm obtusely trigonous to subterete, usually not flattening when pressed and dried. 3. Culm smooth. 4. Achene with conspicuous basal callosity 4. Achene without basal callosity. 5. Achene linear. ae ee) scales weak]
ie ae aise G2 ie Jas C. distinctus Steud.
times as long as wide; dorsal edge ¢
ened taste tas ; ee Steud. 5. Achene ae ea to oblong-ellipsoidal, 2 — 3 times
as long as wide; dorsal edge of scales merely curved into a
simple arc or angle, but not as above.
6. Bracts less than 3 mm wide, 3 or fewer bracts present, the longest of these usually strict and appearing as a continu- ation of the culm; culms slender, | — 2 mm wide at mid- 7. Achenes broadly ellipsoidal; scale cips oe
excurved; species common and wide-ranging in Oe wies- eatin see tat tebe C. acuminatns T. & H. 7. Achenes narrowly ellipsoidal to oblong; scale
more or less ee not conspicuously sea
species rare in U.S., stricted to Texas, Oklahoma,
Louisiana, and western "Flor > wielieaeas armas G; ie Vahl Widest bract usually more oon 4 mm broad, 4 or mc bracts exceeding the inflorescence, all bracts ascending to
spreading; culms mostly (2-) 3—5 mm wide at mid- culm
ON
8. Seikelee t at least 3.0 mm wide; scales with straight to
excurved tips, thus spikelet with toothed outline;
scales golden-yellow; in U.S., species restricted to
California and Oregon. ...............0.. C. eragrostis Lam. 8. Spikelet less than 3.0 mm wide; spikelet with an en-
tire outline; scales stramineous to brown, sometimes
yellow or red tinted, or pale green or whitish; species
of south-central and southeastern U. §
9. Achene broadly ellipsoidal, 2 —
3 times as long as wide,
wide; scales ovate, 0.6 mm wide as seen laterally, stramineous to yellow- brown; in the U.S., species restricted to Texas and
No
a eeu ehh har eae eee C. ochraceous Vahl
Achene oo) ellipsoidal, 3—4 times as long
as wide, 0.2 — mm wide; scales ee
0.3-0.4 mm wide as seen laterally, pale green or whitish.
10. Plants robust, usually more than 5 dm high, base blackish purple, leaf bases persistent and becoming fibrous with spikes con- spicuously compound (or rarely oa units globose; spikelets loosely arranged, with 16 — 32 scales; mature scales pale green
eee giis atte sh eek eh a eee Rae aun be C. entrerianus Bock.
10. Plants less than 5 (7) dm high, base eas ob ish; leaf bases not persistent; spi mpact, conical, and appearing simple, ae individual ee difficult to ae spikelets with 12— 16 scales; mature sc whitish to tawny. ............. ie (L.) Retz. a ae scabrid. Scale tips conspicuously excurved. ............ C. acuminatus T. & H. 7 Scale tps straight to slightly incurved. ants robust, mostly more than 5 dm high; leaf bases blackish purple; culms ae scabrid; scales pale green, di eee at about $0 desrecss 44.4ic4.404 9m Hacs C; entrerianus Bock. . Plants mostly less than 5 (8) ie high; leaf bases stram
—_ i)
us to bro wnish; culms densely scabrid; scales yellowi i freee at about 45 degrees............. C. surinamensis Rottb.
DISTRIBUTION AND ECOLOGY
As shown in figure 1, Cyperus entrerianus is primarily distributed in temperate South iaiceica and rarely in Mexico. Thus, it 1s not surprising that it has persisted and increased its range upon introduction into temperate North America, and it probably will continue to spread.
Cyperus entrerianus is a copious producer of achenes and is often locally abundant in its disturbed habitat. It has been observed growing on mucky sands in southeastern Georgia and northwestern Florida and sticky clays in southern Louisiana and southeastern Texas. Thus, soil texture seems not to be a major factor determining its distribution. However, it does apparently require disturbed sites with high-hydroperiod soils, such as ditches, depressions in flatwoods, pond margins, stream bottoms, and edges of salt marsh. Table 2 contains a composite list of species associated with C. en- trerianus 1n southeastern Georgia.
In addition to collections made by me from 1987— 1989, other specimens of C. entrerianus have been located at FSU, IBE, an DB, which had been identified variously as Cyperus pseudovegetus Steud., C. robustus Kunth, C. virens Michx., and C. virens var. drummondit (T. & H.) Kikenth. Distribution of C. entrerianus in the United States is shown in figure 2. Following is a complete list of Cyperus entrerianus specimens from United States, which I have seen.
Specimen citations. ALABAMA: Baldwin Co.: 1.1 mi E of Mobile, heavily disturbed fill area along and § of ae a at edge of Mobile Bay, 8 Aug 1989, Carter 8095 (MO, SMU, VDB, VSC). FLORIDA: Calhoun Co.: S$ of Blountstown, 23 Jul 1977, God- frey 75970 (FSU, IBE, VDB). Escambia Co.: Pensacola, on Scenic ek 24 Jul 1974, Godfrey 73755 (FSU, VDB); S ae of Ensley, wet ditch along hwy. 29, 8 Aug 1989, Carter 8093 (FSU, MO, VDB, VSC). Gulf Co.: 7.5 mi N of White . ity, by Fla. Ret. 71, 4 Aug 1981, Godfrey 79070 nan a M & K Tract, generally 6.5 air mit NW of Apalachi-
74
Taste 2. A composite list of species associated with Cyperus entrertanus in southeastern Georgia.
Carex festucacea Schk. Lipocarpha maculata (Michx.) Torr. Cuphea carthagensis Jacq.) Macbr. Ludwigia microcarpa Michx Cyperus croceus Vahl L. palustris (L.) Ell
C. haspan L. Lythrum alatum Pursh var. lanceolatum (Ell.) T.&G. C. odoratus L. Mitreola petiolata (Gmel.) T. & G C. pseudovegetus Steud. Murdannia nudiflora (L.) Brenau C. strigosus L. Phyllanthus urinaria L
C. surinamensis Rottb. Polygonum hydropiperoides Michx. C. virens Michx. Rhynchospora cephalantha Gray Eleocharis tuberculosa (Michx.) R. & S R. corntculata (Lam.) Gray. Hypericum mutilum UL. R. inexpansa (Michx.) Vahl Juncus marginatus Rostk. R. microcarpa Baldw. ex Gray
J. repens Michx. Verbena brasiliensis Vell.
cola, W of Sand Creek, T8S, R8W, S 1/2 Sec. 7, 8 Nov 1985, meng 9018 (FSU); 2 mi W of Daniels Road, ca 4 air mi NE of Overstreet, ee 15, T5S, RILW, 1 Jun 1989, Anderson 12034 (VSC), 20 Jul 1989, Anderson 12172 (MO, VSC); western edge of Wewahitchka, ditch by hwy. FL 22, 5 Aug 1989, Carter ae ‘(FSU MO, VDB, VSC). GEORGIA oe ee 0.8 mile W of Nahunta, 4 Jul 1988, Cane & Carter 6960 (FSU, GA, MO, SM B, VSC). Camden Co.: 2 mi NE of Waverly, along Hwy. US 17, 4 Jul 1988, Carter & a 6935 (FSU, GA, MO, SMU, VDB, VSC). Glynn Co.: ca 1.5 mi S of Brunswick near intersection of Hwys. US 84 and GA 50, oo of eels 26 Aug 1988, Carter & McCormick 7435 (FSU, GA, MO, SMU, VDB, VSC). Liberty Co.: just SE of Flemington city limits along Hwy. US 82, 26 Aug 1988, pe & McCormick 7447 (FSU, GA, MO, SMU, VDB, VSC). Ware Co.: western part of Waycross, N of Hwy. US 84, near corner of New Mexico and Virginia Avenues, 2 Sep 1987, Carter 6296 ae GA, MO, SMU, VDB, VSC); Waycross, creek bottom at corner of Blackshear and Riverside Streets, 18 Aug 1988, Carter 7319 (MO, SMU, VDB, VSC). LOUISIANA: Calcasieu Parish: 9.3 mi N of Hackberry, ditch along Hwy. LA 27, local in sticky clay, 10 Aug 1989, Carter 8130 (MO, SMU, VDB, VSC). Jefferson Davis Parish: E of Hwy. US 165 and 0.25 mi S of Hwy. I-10, 1.4 mi E of Hwy. LA 383 Iowa exit, locally common along mowed ditch and adjacent road shoulder in vicinity of rice fields, 10 Aug atiee Carter 8127 (MO, SMU,
VDB, VSC). St. Landry Parish: ca 3 mi W of Eunice alone Hw es 25 Jul 1975, Allen 6674 (VDB). TEXAS: Chambers Co.: 9.8 mi E of Wallisvi tand 11.9 mi E of Trinity River, sticky black clay at edge of rice field N of Hwy. I-10, common asd locally ae. along an cig ge 6 mile stretch of Hwy. I-10, 10 Aug 1989, Carter 8142 (M SMU, VDB, VSC). Fort Bend Co.: roadside clearing on Hwy. 59, 0.1 mi S of a Road, sandy soil, 29 Jul 1981, Kessler 4739 (VDB). Harris Co.: 0.5 mi E of Peek Road exit to Kary, ditch along Hwy. I-10 near rice fields, sticky black clay, 10 Aug 1989, Carter 8144 (MO, SMU, VDB, VSC).
DISCUSSION
The earliest collection of Cyperus entrerianus that I have seen from the United States is R. K. Godfrey 73755 (FSU) collected in 1974 in Escambia County, Florida. Thus, it would appear that C. entrerianus is a relatively recent introduction into southeastern United States. The largest popula-
] 1 . ircl kK 4 ?
FIG. 1. The distribution of Cyperus entrertanus. Closed circles b
on Kikenthal (1936) and Pedersen (1968).
tion (Carter 8142) was seen in Chambers County, Texas, where C. en- trerianus is common and locally abundant, in places forming almost pure stands, along an approximately 6 mile stretch of highway I-10.
Cyperus entrerianus is often locally abundant, and in eastern Texas and
Reis ‘a pa PD east re AS my Spy Secs:
[TY ns
if
mae ry = Ln]
Mes ywltra
any
rN Seccaulon ccalntes ag RES SY os
ON) eS
FIG. 2. The distribution by county of Cyperus entrertanus in the United States.
southern Louisiana it seems to be highly correlated with rice culture. Curiously, label data on a specimen (E. Lurvey 440, MO) of Cyperus en- trerianus indicates that it was collected in a rice paddy in Paraguay. Addi- tional information about sources of rice seed used in southeastern Texas and southern Louisiana might be helpful in determining the origin of C. en- trertanus in the United States. If the introduction of C. entrerianus is not an artifact of human activity, then it seems reasonable to assume that it might have been introduced by migrating water fowl. Electrophoretic analysis and comparison of proteins from individuals of different populations might show whether multiple introductions of C. entrerianus have occurred along the Gulf and Atlantic Coasts of southeastern United States or whether it has spread out from a single colony.
All of the southern Georgia populations were located in ditches along highways and two particularly extensive populations (Carter G Carter 6960 and Carter & McCormick 7447) were beside highways along which major construction had recently occurred. Thus, it ts likely that C. entrerzanus is being moved about along highways and perhaps in part by highway con- struction and maintenance equipment.
Cyperus entrerianus is to be expected in at least coastal regions of other southeastern states from North Carolina to Texas. Based upon its wide- spread occurrence in temperate South America far inland and at altitudes up to 410 m (P. Goetghebeur 4791, VSC), it is reasonable to expect that it
TF
will continue to spread from the outer coastal plain into interior physio- graphic regions. Additional life history and ecological studies should be done to determine the extent that C. entrerianus might become a harmful weed in North America.
ACKNOWLEDGEMENTS
I would like to thank Sidney McDaniel (IBE) for access to specimens and use of the library of the Institute for Botanical Exploration and, also, for sharing his knowledge of C. /wzu/ae in Peru. Paul Goreneerae (GENT) very kindly sent an excellent specimen (Goetghebeur 4791) of C. entrerianus from Argentina, which was most helpful. I am grateful to Robert Godfrey and Loran Anderson (FSU) for lending specimens, to Loran Anderson for sending recently collected material of C. entrerianus, and to curators and staff at MO for making specimens available for study. Loran Anderson, Charles Bryson (SWSL), Gerrit Davidse (MO), and Gordon Tucker (NYS) made helpful comments on the manuscript, Robert Kral (VDB) made specimens available for study, Blanca Leon provided the Spanish transla- tion of the abstract, and Sandra Howell (Louisiana Tech University), while she was a student intern at Missouri Botanical Garden, rendered able assistance by patiently trying out my key. The holotype of C. entrerianus was examined through the courtesy of the staff at B. Publication costs were met by a Valdosta State College Faculty Research Grant.
REFERENCES
BARROS, M. 1938. — Argentinas III. Anales Mus. Argent. Ci. Nat. “Bernardino en BY 253)=
BOCKELER, O. ere ‘er a thiels neuer, thiels ungenugend beschriebener Seca . Flora 61: 138—-
DENTON, M. : 1978. A taxonomic treatment of the Luzulae group of Cyperus. Contr. Univ. Michigan Herb. 11(4):197 — 271.
GODFRE and J. W. ee 1979. Aquatic and wetland plants or a, Ga States. Vol. 1. University of Georgia Press, Athens
KUKENTHAL, G. 1935—6. en eee -Scirpoideae-Cypereae. In: A. Baie (editor),
Das Pflanzenreich. IV. 20 (Heft 101):1—671.
PEDERSON, T. M. 1968. “Cyperaceae.” pp. 315 — 421. In: A. L. Cabrera (editor). Flora de la Provincia de Buenos Aires. Vol. 1. Coleccion Cientifica del I. N. T. A. Buenos Aires
CALL FOR APPLICATIONS FOR THE 1990 DELZIE DEMAREE TRAVEL AWARD
An endowment to underwrite an annual travel award ($250-$300) in memory of Dr. Delzie Demaree has been established. This award is given annually to a graduate student in systematics for travel to the Systematics Symposium sponsored each fall by the Missouri Botanical Garden in St. Louis. Such an award is a very appropriate way to honor Dr. Demaree because of the high esteem and long record of attendance (right up to the meeting of October 1986 prior to his death at age 97 the following July). In addition, this is a significant way to continue his legacy of assistance to students of botany.
The recipient of the 1989 travel award was Mr. Scott C. Zager, Univer- sity of Northern Iowa, Cedar Falls. Scott is studying 15 Iowa species of Carex, section Ovales. His major professor is Dr. Lawrence J. Eilers.
Letters of application for the 1990 travel award should be mailed to Donna M.E. Ware, Herbarium, Dept. of Biology, The College of William and Mary, Williamsburg, Virginia 23185. Applications should be postmarked by 15 Aug 1990. A complete application shall consist of a letter from the graduate student describing briefly their research and the benefits of symposium attendance, and a letter of recommendation from the student’s major professor.
Sipa _ 14(1):78. 1990.
EXTENSION OF NATIVE RANGE OF SABAL MEXICANA (PALMAE) IN TEXAS TO INCLUDE CENTRAL COAST
LANDON LOCKETT
210 Stevenson
Austin, TX 78703, U.S.A. ROBERT W. READ
Botanist Emeritus, National Museum of Natural ee Smithsonian Institution, Washington, D.C. 20560, U.S.A.
Authors report discovery of a wild population of Sabal mexicana Mart. in the Central Coast (Coastal Bend) region of Texas, present evidence that the species is indigenous to that region, and discuss conservation prospects.
RESUMEN
Los autores relatan el descubrimiento de una pean salvaje de Sabal mexicana Mart. en el litoral central de Texas, presentan evidencia de que esta region es parte de la distribucion nativa de la especie, y discuten las perspectivas a6 su conservacion
Although Sabal mexicana, the caulescent palm native to Texas, has under- gone several changes in its classification since first identified as a species distinct from Sabal palmetto, most botanists have described its native range in the U. S. as limited to the Lower Rio Grande Valley, at the extreme southern end of Texas. In identifying this palm as S. texana Small (1927), for example, described its distribution as “confined to a comparatively small area in the lower Rio Grande Valley.”
Orator E Cook, however, apparently believed otherwise. Cook (1908, p. 5n.a) stated that “Tall palmettos were seen in Jackson County as late as 1876 by Mr. J. D. Mitchell, of Victoria,” and in 1913 (p. 11) Cook noted that “Inodes texana,” as he called it, “seems to have extended much farther northward only a few decades ago, and specimens may still be found about Indianola or at other points along the Gulf coast.
Cook (1913) goes on to describe a new taxon — Inodes exul — in order to identify a population of sabal palms, of unknown origin, that for many years had been cultivated in Victoria, Texas. Beccari (1907) had by then rejected the genus Iodes, and a careful comparison by Davis (1942) elim- inated exw/ as a separate species by showing that the morphological features Cook considered distinctive for ex#/ fell within the range of variation of
Sipa 14(1):79— 85. 1990.
80
Sabal texana. Finally Moore (1971) reduced S. texana to synonymy of S. mexicana, a single species ranging from Texas to Central America.
But the basic mystery remained. Where had the Victoria palms come from? No Victoria resident, either in Cook’s time or today, seemed to know. Prompted by Cook's footnote, however, we suspected that Victoria's mystery palms were of local origin, and in August of 1989 initiated a search. We were soon rewarded by the discovery of a wild population of S. mexicana in the Central Coast, as well as of evidence that this species is native there.
After our inquiries in Victoria about wild palms, either past or present, resulted in an article in the THE VICTORIA ADVOCATE (Bowen 1989), four fishermen called to tell of palms, up to twenty feet tall, on nearby Garcitas Creek. Historian Brownson Malsch, of Edna, told us of a tall palm on the east bank of Garcitas Creek that used to be visible from the Highway 616 bridge. Rancher John M. Bennett took us to the fallen remains of this palm, which we found to have a trunk of at least 37 feet. Nearby stood a living specimen of S. mexicana with a 13-foot trunk, and scattered about the Bennett ranch, in the bottomland forest along Garcitas Creek, were other caulescent specimens, and seedlings. Bennett told us the tall palm had been standing there when his grandfather bought the ranch in 1890, and that there had been no prior settlement on the east side of the creek.
A boat trip up Garcitas Creek revealed a population of S. mexicana begin- ning approximately 300 yards north of the Highway 616 bridge and exten- ding for 2—3 miles upstream. Specimens ranging in size from seedlings to 20 — 25 feet stood along the bank and were scattered through the bottom- land forest, up to perhaps 50 yards from the creek. Some were on the edge of the water and looked as though they would soon be lost to erosion. Others stood on relatively high banks. All but two of the specimens we saw were on the east or Jackson County side of Garcitas Creek. The two on the west (Victoria County) side were both near the southern end of the popula- tion. Upstream from these two the only palms seen on the west side were an occasional specimen of S. minor. We saw no S. minor on the east side, or anywhere in association with S. mexicana. Specimens were sent to US (Lockett 101, Aug 1989) and SMU (Lockett 102, Sep 1989).
Sabal mexicana is readily distinguishable from S. mznor, the only other palm known to occur in the Central Coast. Aside from being caulescent, mature specimens show highly filiferous, strongly downcurved leaves hav- ing a costa measuring at least two feet, and dead petioles with split bases cling to their trunks. Even young specimens differ from S. minor by their relatively longer petioles and leaf segments, lighter color, and highly fili-
81
ferous, strongly costa-palmate leaves. The long, lax segments of the many seedlings give them a grass-like appearance.
Because S. mexicana is widely cultivated in South Texas it would be easy to suggest that the Garcitas Creek population has escaped cultivation. His- toric evidence, however, indicates that tall, rrunked palms have been found in the Central Coast since the earliest attempt at colonization. According to Weddle (1987, p. 11), in 1685 French explorer René-Robert Cavelier, Sieur de La Salle, established his short-lived Fort Saint-Louis colony on Garcitas Creek. The colony’s historian, Henri Joutel, devoted several pages of his account of the colony to a careful and impressively accurate descrip- tion of local flora and fauna. As related in Margry (1876 — 86, vol. 3, p. 212), this description includes the following passage: “I/ y a une autre espece d'arbres qui ont des branches longues d’environ trois ou quatre pieds, et leurs feuilles sont comme celles des lataniers, seulement plus grandes et plus larges. Ils portent un petit fruit, mais jen’en scaurois dire le goust, n'en ayant point mangé; mats un de nos gens me dit qu'il luy avoit semblé bon.”
Carlos E. Castaneda (1936 — 58, vol. 1, pp. 289 — 290) called him “the trusty Joutel, a man of sense and observation.” This is evident from Joutel’s account. Though knowing no names for many of the creatures and plants of the Texas coast, he describes recognizably everything from horned lizards to yucca — which, by the way, he clearly distinguishes from palms. There is no reason to suppose Joutel was imagining the plants he described in the quoted passage.
What were these plants? If palms, were they S. mexicana, or S. minor? Latanier is the vulgar name applied to Saba/ in Haiti. Bomhard (1935) remarks that “Latanier is the French form of the native name of tall, fan- leaved palms indigenous to certain islands belonging to France off the southeast coast of Africa.” She adds that in Louisiana this name, or “Jatania,” is commonly applied to S. minor. Perhaps Joutel had S. minor in mind when he used the word /atanier. If so, he clearly distinguished the “trees” (arbres) he saw on the Central Coast of Texas from S minor. The 3- or 4-foot petioles (branches) would be within normal range for S. mexicana (Davis, p. 94), but too long for most S. minor. The blades (fewz/les) are “plus grandes et plus larges’ than those of the /ataniers. Most important of all, Joutel describes the palms he saw as bearing an edible fruit. This clearly fits the fruit of S. mexicana (Tull 1987), but not the very small fruit of S. mznor.
In 1688 Karankawa Indians massacred the adult inhabitants of Fort Saint-Louis, but spared a few children, including Jean-Baptiste Talon. Al- most 10 years later Jean-Baptiste and his brother Pierre, whom La Salle had left with the Tejas Indians, had made their way to France, where they were
82
interrogated about their years spent with Indians in what is now Texas. In their deposition the Talons reported that there were “pa/miers” along all the rivers, including one — possibly the Guadalupe — to which the Karankawas had taken their women and children (including Jean-Baptiste) while they waged war against another tribe. (Weddle 1987, Part IV.)
During the 300 years since the time of La Salle’s colony most of the Central Coast palms appear to have been lost to wharf building. Teredo navalis, the shipworm, destroys wood immersed in warm salt water. Because Saba! palm trunks were immune to such destruction palm logs were in demand for wharf construction, and during the 19th century many were used as piles for the wharves at the port of Indianola, on Matagorda Bay (Malsch 1988). According to Malsch (pers. comm.), Francis E. Huck, whose father operated a lumber yard in Indianola before the city’s destruc- tion by a hurricane in 1886, reported to him that locally acquired palm logs were used for wharf pilings until the supply was exhausted, and that thereafter Huck’s father had had to import them.
This evidence from Malsch fits with J. D. Mitchell’s statement to O. EF Cook. If Mitchell saw tall palmettos in Jackson County “as late as 1876,” the implication is that something happened to them after that, but before the destruction of Indianola in 1886. Cook’s notes, salvaged by Dr. An- drew Archer of the Smithsonian Institution (where they are now housed) after they had been discarded by Cook’s employer, the United States Department of Agriculture, help complete the picture. In an item dated “Kingsville, [Texas] Aug. 3, 1906” Cook records Mitchell's tall-palmetto remark, then adds, “They were cut down for the sake of the wood. This may be taken to indicate that the native palmetto of Texas ranged at one time much farther North than at present, and that it will grow freely in the open air over the whole southern part of the state, as many casual plantings also prove.”
Mitchell, whom Burke (1978) called “the first native Texas naturalist,” was in contact with Cook because from 1904 almost until his death in 1922 Mitchell did field work for the U.S.D.A. Entomologist W. D. Hunter (1922), with whom Mitchell collaborated for years, called Mitchell “a fountain of accurate information.”
Palms are prized as ornamentals, and early settlers encountering them in the wild sometimes eliminated whole populations by digging them up for transplant. Brown (1976b, p. 152) documented cases where “two popula- tions [of S. palmetto} north of North Inlet [S. Carolina} have been either completely removed for landscaping purposes or otherwise destroyed since 1930. A small group of trees near North Litchfield Beach at Midway Inlet were removed in the early 1960's. At Murrells Inlet a much larger popula-
83
tion on Drunken Jack Island was removed beginning in the early 1930's. This extinct population represents the northernmost modern extension of the species on the South Carolina coast . . .”
Historian Malsch’s notes indicate that in 1925 Mrs. Alexander Lowe told him that in 1875 her husband bought two small palms from a wagon lot that pioneer nurseryman Gilbert Onderdonk brought from Jackson County for sale in Victoria. According to Malsch (pers. comm.) a few of these original transplants are still standing in Victoria, and many younger palms have been propagated from their seed. Cook (1913) identified the “type individual of the new species” (Inodes exu/) as the tree that “graces the lawn of Mrs. Martin O’Connor, of Victoria.” Mrs. O’Connor’s nephew, Mr. Dennis O’Connor (pers. comm.) reports that this palm, today over 40 feet tall, still stands at the junction of Liberty and Power streets. Although he has no record to prove it Malsch, who as a child was a neighbor of the Martin O’Connors, believes this palm is one of Onderdonk’s original trans- plants.
Even if historical evidence were lacking we believe it would be reason- able to conclude that the native range of S. mexicana includes the Central Coast. The existence of a wild, viable, reproducing population, only 200 miles from the known native range, is in itself evidence that the site of this population is part of the native range. Noting that sea currents on the southern Atlantic coast are northerly in late spring and summer, when the fruit of S. palmetto is dry and buoyant, Brown (1976a) suggested sea dis- persal as the mechanism that could have carried seed of that species from Florida up the Carolina coast to the northern extreme of its range. Although alongshore currents on the Texas coastal bend are normally southerly, approximately twice a year, usually in spring and early summer, this flow may be reversed. Likewise eddies off of the northerly loop current, in the central gulf, can cause a northerly alongshore current when they split upon arrival at the coast. (Kerry Whitledge, Senior Marine Scientist, University of Texas Marine Science Institute, Port Aransas, TX, pers. comm.) Brown also demonstrated that dry S. palmetto fruit remains buoyant several weeks. Since we have observed that dry S$. mexicana fruit floats readily, the dry skin forming an air chamber in which the seed rattles, we see no reason why S. mexicana could not follow the same dis- persal pattern as that suggested by Brown for S. palmetto. According to Davis (1942, p. 85) S. mexicana in the lower Rio Grande Valley may bloom almost any time of year, meaning dried fruit would be available year round. Floating fruit, washed down the Rio Grande, or rivers in Mexico, could be carried north to the Central Coast where tides and storms could push it into rivers and lowlands. Once mature trees were established on the Central
84
Coast their fruit could be spread by birds and small mammals. Coyotes, raccoons, chachalacas and rodents have been observed to eat S. mexicana fruit. (Rose Farmer, Manager, Sabal Palm Grove Sanctuary, Brownsville, TX, pers. comm
Obviously there are no cold stress restrictions on establishment of S. mexicana along the stretch of coast now determined to be populated with this species. As stated by Read (1974, p. 41) “. . . over a broad area of uniform edaphic and climatic conditions the same species of palm will likely be seen. On the other hand a change in the substrate is likely to support distinct species, 7
We thus believe it is safe to assume that in prehistoric times currents would have carried the seeds to all nearby suitable habitat. Putting it another way, it is questionable to speak of escape from cultivation when we find a population so well adapted to its environment, and showing evid- ence of having grown in that environment for many years, if the site of that population appears to be within the natural dispersal radius of the known native range.
Since we know of no age studies of S. mexicana we do not know the age of the larger Garcitas specimens. We suspect, however, that since their height is well below the approximately 50 feet the species can attain (Texas Forest Service 1989, p. 8), and since we found a dead specimen with a trunk of at least 37 feet, the Garcitas trees are younger generation, seeded by a few specimens that survived lumbering and transplantation. Further, we sus- pect that since the days of wharf building transplantation has continued to be a factor tending to limit the wild population. In Bennett Park, a small, unimproved and unpoliced county park on Garcitas Creek, we saw many young specimens of S. mexicana, but none that were caulescent, even though caulescent specimens were observable on adjacent private land.
Given the apparent reproductive vigor of the Garcitas population, we believe that conservation prospects for this and other possible stands in the Central Coast (or elsewhere) are promising, if habitat remains undisturbed and if, through education and protection, transplantation can be discoura- ged. Meanwhile we continue to look for other wild populations of S. mexicana, as well as further evidence of the species’ historic range. We urge all who discover specimens in the wild, or historic references to trunked palms (“tall palmettos,” etc.), whether in the Central Coast or beyond, to contact us.
ACKNOWLEDGMENTS
We thank Greg Bowen of THE VICTORIA ADVOCATE for publiciz- ing our search; Lon Drushel, Elbert Post, Victor Spiegalhower, and Rawley Koehl for telling us of the Garcitas palms; and Lon Drushel for taking us in
85
his boat to see them. We also thank John M. Bennett and Emily Dial for letting us search and take specimens on their ranches. We are grateful to Charles Spurlin, Robert Shook, Wayne McAlister and Mitzi Stewart, all of Victoria College, for information provided, and to the Victoria College Library for access to its J. D. Mitchell materials. We thank Sister Ann Linda Bell, Chairman of the Foreign Language Department of Our Lady of the Lake University, for her help in interpreting references to palms in the Joutel and Talon documents, and we are especially thankful to Carol Ramsay, James Stewart and George Stevenson for helping with searching and specimen collecting, and for their constant encouragement.
REFERENCES
BECCARI, O. 1907. Sabal texana. Webbia 2:78
BOMHARD, M.L. 1935. Sabal louisiana, the correct name for the polymorphic palmetto of Louisiana. J. Wash. Acad. Sci. 25:3
BOWEN, G. oe Wild search begins for ee Palm’. The Victoria Advocate, Aug 10, 1989, p
BROWN, K.E. nce aac studies of the cabbage palm, Sahal palmetto. 11. Dis- persal, predation, and escape of seeds. Principes, 20:49 — 56.
1976b. "Ecological studies of the cabbage palm, Sabal palmetto. IV. Ecology and geographical distribution. Principes 20: 148 —
BURKE, H.R. 1978. Contributions on is Mitchell to the early knowledge of Texas
mano.)
insects. Weleheimer Entomol. Ser. 24
CASTANEDA, C.E. en oe 58, Our ae heritage in Texas, 1519 — 1936. Von Boec-
mann-Jones Co., . 7 vols.
COOK, O.F 1908. ae : vegetation on the South Texas prairies. U.S.D.A. Bur. Pl.
Industr. Circ. No. 14. 1913. A new ornamental palmetto in southern Texas. U.S.D.A. Bur. Pl. Industr. Circ. No. 113:11— 14.
DAVIS, A.M.T. 1942. A study of Boscaje de la Palma in Cameron County, Texas, and of Sabal texana. _ Master of Arts thesis, University of Texas. 111 pp.
HUNTER, W.D. 1922. J. D. Mitchell [obituary]. Science, n. ser. 55: 469.
MALSCH, B. 8 eee the mother of western oo (Rev. Ed.) State House Press, Austin. 351 pp.
MARGRY, P. a (876— 86. Découvertes et établissements des francais dans l'ouest et dans le sud de Vikonpidu a (1614— 1754). Maisonneuve, Paris. 6
MOORE, H.E. 197 Perea and corrections to “An annotated ean of cultivated palms”. ee 15:102
READ, R.W. 1974. The sane of the palms. Principes 18:39 — 50.
SMALL, J.K. 1927. The palmectto-palm — Sabal texana. J. New York Bot. Gard.
26:132 > 143:
TEXAS FOREST SERVICE 1989. Texas big tree registry. Information and education, Texas Forest Service, Lufkin. 10 pp.
TULL, D. 1987. A practical guide to edible and useful plants. Texas Monthly Press,
5, 2 BPD: WEDDLE, RS. 1987. La Salle, the Mississippi, and the Gulf. Texas A&M University Press, College Station. 328 pp
BOOK REVIEWS
JOHNSTON, MARSHALL C. 1988. The Vascular Plants of Texas: A List, Up-dating the “Manual of the Vascular Plants of Texas.” Pub- lished by the author, 3905 Avenue G, Austin, TX 78751. Paper $11.00 + 8% sales tax in Texas; outside North America $15.00.
This is an update of the Correll & Johnston “Manual of the Vascular Plants of Texas,” published in 1970. This Manual is still available from the University of Texas at Dallas Book Store in Richardson, Texas. The new data is organized by page numbers corresponding to those of the Manual facilitating quick reference between the two. The literature citations follow the page numbers with the index last. Not only does this update the current knowledge but it was also to make some corrections in the original ene In doing so, I find in the reference section that my middle initial
” has been replaced with “J.” and “H.” Once a mistake is made, it is certain to be repeated not only by others but by myself as well. There will always be some aspect that one does not agree upon but this is an excellent attempt for this moment in time without reproducing the entire manual. WFEM.
JOHNSTON, MARSHALL C. 1990. The Vascular Plants of Texas. A List, up-dating the Manual of the Vascular Plants of Texas. Second edition. Published by the author, 3905 Avenue G, Austin, TX 78751. Paper $14.00 (US) prepaid shipment to North American addresses not requiring invoicing or billing. For shipments outside North America or shipments requring invoicing or billing send $17.00 (US). For deliveries to Texas addresses add 7.75% sales tax.
WOFFORD, B.E. 1989. Guide to the Vascular Plants of the Blue Ridge. 384 pp. University of Georgia Press, Athens, GA 30602. Paperback $15.00; Hardbound $35.00.
This ts an excellent manual for the identification of the vascular plants of the Blue Ridge Province. It has an illustrated glossary with the text con- sisting of dichotomous keys followed by indices to both common and scien- tific names. In identification manuals, the species description is a summa- tion of the taxonomic characteristics that are usually present in the keys that distinguish each taxon from the others in the treatment. Only in monographic treatments are the detailed species descriptions given in full. Thus, the lack of species descriptions does not diminish the effectiveness of this manual but actually enhances it.
Sipa 14(1):86. 1990.
SYNOPSIS OF CAREX SECTION LUPULINAE (CYPERACEAE) IN TEXAS
STANLEY D. JONES AND STEPHAN L. HATCH . Tracy eas Depar riment of Range Scie Texas AGM University, College Station, TX 77843, U.S.A.
ABSTRACT
Five species of Carex section Lupulinae occur in Texas; C. lupulina, C. lupuliformis, C. louisianica, C. intumescens and C. gigantea. Carex lupulina is the most common and wide- spread of the five. Carex lupuliformis, which is rare, grows mostly in calcareous sites in swampy woodlands. It has been recorded only in the northeast corner of the state in Bowie and Marion counties. Carex louisianica occurs infrequently in the eastern 1/3 of Texas grow- ing in acidic soils of swampy woods or bottomland hardwood forests. Carex intumescens 1s widespread across the distributional range of section Lupulinae in Texas but is infrequent within its diminishing habitat of acidic bottcomland hardwood forests. In Texas, C. gigantea is the rarest member of the section having not been collected for 46 years. The only two collections were made in Harris and Polk counties in acidic swampy woodlands. An artifi- cial dichotomous key, county distribution maps and comparable species descriptions are provided.
INTRODUCTION
Carex, with 31 sections represented by more than 80 species, is the largest genus of vascular plants in Texas. As is the case with most genera of the family Cyperaceae, Carex is difficult taxonomically. The section Lupulinae (J. Carey) Mackenzie is endemic to central and eastern North America (Reznicek and Ball 1974). It is restricted to the eastern 1/3 of Texas, being found westward to Hays County and southward to Nueces
ounty. However, the greatest concentration and diversity are found in the eastern 1/5 of the state. Section Lupulinae in the subgenus Carex has 3 stigmas, trigonous achenes, and unisexual spikes. Other characteristics of this section include perigynia 1 cm long or longer, coarsely nerved perigynia, a perigynium-body that is ovoid or globose-ovoid, and leaf lades that are strongly septate-nodulose.
Six species have been recognized in this section by Mackenzie (1935, 1940), Fernald (1950), Gleason (1952), Voss (1972), Reznicek and Ball (1974) and Menapace et al. (1986). Five occur in Texas: C. /upulina Willd. , C. lupuliformis Sartwell, C. louisianica Bailey, C. intumescens Rudge and C. gigantea Rudge. The remaining species C. grayi Carey is found im- mediately east and northeast of Texas and from the Gulf states north to southwestern Quebec. Waterfall (1979) listed C. grayz as occurring in
Sipa 14(1):87 —99. 1990.
88
Oklahoma. Rob Naczi (MICH; per. comm.) has recently collected it in McCurtain County, Oklahoma (Naczi 1890, MICH). Tony Reznicek (MICH; per. comm.) has collected C. grayi (Reznicek 8490, MICH) along the Little River in Sevier County, Arkansas. He stated, not only is it found immediately adjacent to Texas, but it quite likely occurs locally in river bottoms in extreme northeastern Texas, although it has not yet been collec- ted there.
Correll and Johnston (1970) recognized C. sntumescens Rudge, C. gigantea Rudge, and C. lupulina Muhlenb.. Carex /upuliformis Sartwell and C. louisianica Bailey were recognized as forms of C. /upulina Muhlenb.. They referred to Muhlenberg as the authority for C. /upulina as have other authors. However, Reznicek and Ball (1974) stated that Willdenow is the correct authority.
The primary objective of this paper is to examine the taxonomic status of C. lupuliformis and C. louisianica in the Texas flora. Other objectives are to provide distribution maps by county for each of the five species occurring in Texas and provide comparable diagnosis for each of the five taxa. The distribution maps were based on herbarium specimens.
METHODS AND PROCEDURES
This study was based on about 300 specimens examined from the follow- ing herbaria: (acronyms follow Holmgren et al. 1981) ASTC, MO, NLU, SHST, SMU, SWT, TAES, TAMU, TEX, UA and US. In addition, an isotype of C. /upuliformis was examined from PH. Field trips to east and southeast Texas were conducted throughout 1988 to supplement existing distribution and habitat records. Dorsal and ventral are used synonymously with abaxial and adaxial in this paper. Maturation dates are given as opposed to flowering dates because mature plants in fruit are used to es- tablish the diagnostic characters in all previously published artificial keys for Cyperaceae.
Micrographs were taken of representative achenes of each species using a JOEL-25s scanning electron microscope. Achenes were mounted on aluminum stubs via doubled sided tape and coated with 400 A of gold- palladium using a Hummer | sputter coater. Micrographs were taken at an accelerating voltage of 12.5 Kv’s. Photographs of the pistillate and stam- inate spikes were taken from herbarium sheets using a 35 mm Canon AE- 1 single lens reflex camera with Kodak MTAX film (100 ASA).
Species descriptions will be abbreviated to reflect characters that are diagnostic or which can be used in conjunction with other characters to distinguish between C. /upulina, C. lupuliformis and C. louisianica, or where new previously unrecorded information is provided. The chosen characters
89
will be given for all five Texas species. For a recent and complete species description of the section see Reznicek and Ball (1974)
RESULTS
KEY TO THE SPECIES
la. Pistillate spike outline tending to be globose (Fig. 1c); perigynia loosely arranged, spreading, drying dark olive-drab green. ............. 4.C. intumescens b. Pistillate spike outline oblong to cylindric (Figs. la,b,d,e); perigynia either wey arranged or not, drying stramineous, green or light olive- drab gre 2a. eae peduncles greatly exceeding the uppermost pistillate spike (Fig. 1d); perigynia loosely arranged, ascending-spreading ..... 3.C. loutsiantca 2b. Staminate peduncles shorter than to only slightly exceeding the upper- most pistillate spike (Figs la,b,c); perigynia either loosely eee or tightly arranged. 3a. Achenes distinctly wider than long (Fig. 2c), widest above the middle, subtruncate to truncate apically; perigynia loose to tightly arranged, usually spreading at right angles to the main AXIS! (BiG TE) sg anuwudensss i geen s a mea Pad ae Yaa . C. gigantea . Achenes as wide as long or longer, widest near the middle, not subtruncate or truncate apically, perigynia tightly arrange ascending or slightly spreading but usually not at right as to main axis
—
B)
ion
4a. Angles of achene smoothly curved (Fig. 2a), not a
faces flat co slightly concave. ..............0200.0. _ C. lupulina 4b. Angles of achene pointed (Fig. 2d), with ees ik cnobs, faces strongly concave ................. C. lupuliformis
1. CAREX LUPULINA Willd., “HOP-SEDGE”, aa Pl. 4:266. 1805. Tver: Willdenow 17210 ecm B, photo only TRIE
Blades flat, 1.5—6.4 dm long X 4—15 mm wide, long-attenuate, antrorsely scabrous distally on the adaxial and abaxial sides on the nerves, strongly antrorsely scabrous on margins of upper half, septate-nodulose. Bracts leaf-like, flat, 10-55 cm long X 2— 11mm wide, much exceed- ing the culm, antrorsely scabrous on the margins distally, the lower, at least strongly sheathing, septate-nodulose. Pistillate infructescence (Fig. la), below staminate spike, (1-)2 —5 per culm, not aggregated, 1.5 —6.5 cm long X 1.3—3 cm wide, oblong; peduncles 0.5 — 20 cm long, dis- tance between 2 lowest peduncles 1— 20 cm. Staminate inflorescence terminal, | or rarely 2 per clum, 1.5—8.5 cm long X 1—5 mm wide, narrowly linear; peduncles 0.5 —6 cm long, antrorsely scabrous, base of staminate spike shorter than or barely exceeding the top of the uppermost pistillate spike; anthers 2—4 mm long. Pistillate scales 6— 15 mm long 1—2.7 mm wide, lanceolate to lanceolate-ovate, | — 7-nerved, nar-
90
re
ss). la. Carex lupulina, \b. C. lupuliformis, \c.
.l.a—e. Pistillate spike (ps) and staminate spike C. intumescens, 1d. C. louisianica, le. C. gigantea.
I a—e. Achenes: 2a. Carex lupulina, 24% . 2b. C. loutsianica, 23 X . 2c. C. gigantea, 23 X . 2d. C. lupuliformis, 23 X . 2e. C. intumescens, 23 X.
92 rower and shorter than perigynia, white hyaline with green centers, acute to awned, awns to 6 mm long, antrorsely scabrous. Perigynia 11 — 19 mm long X 3—6mm wide, narrowly ovoid, glabrous, shiny, light to medium green to stramineous at maturity, wingless, not corky, inflated, stiffly erect to strongly spreading, sessile to + stipitate, (4-) 8 — 80 per spike, strongly 13 — 22-nerved; beak conic, 6— 10 mm long, bidentate. Achenes (Fig. 2a) 3—4(-4.5) mm long X 1.7—2.8 mm wide, rhomboid, trigonous, + stipitate, faces flat to concave, angles thickened internally. Distribution: Minnesota to Nova Scotia and south to Florida and Texas. Texas: by counties (Fig. 3b); regions 1,2,3 and 4 as defined by Gould (1975). Chromosome number 2”=56 (Reznicek and Ball 1974) »=30 (Wahl 1940). Since C. /upuliformis has a chromosome number of 2”=60, Reznicek has suggested that it is possible that Wahl may have had that species instead of C. /upulima. Maturation dates: April through October. Habitat: Open swamps, wet ditches, somewhat acidic-neutral to calcar- eous soils. Representative specimens: Angelina Co.: 1980, J. Ward & S. Hupp 459 (ASTC). Bowie Co.: 06 Aug 1983, E. Nixon, J. Ward & M. McCrary 12493 (ASTC). Brazos Co.: 11 May 1980, PB Fryxell 3181 (SMU). Cass oe 18 Jul 1967, R. Mitchell 3250 (TEX). Freestone Co.: 28 Oct 1983, E. Nixon & J. Ward 13170 (ASTC). Galveston Co.: 06 Ma 1976, FE Waller 3808 (TEX). Gonzales Co.: 04 Aug 1941, B. age ee (TAES). Grimes Co.: 10 Jul 1988, S. & G. Jones 1818 (ASTC). Hardin Co.: 21 May 1986, L. Brown age STC). Harris Co.: 21 May 1986, L. Brown tone Nene Harrison Co.: O09 Aug 1980, E. Nixon 10496 (ASTC). Hays Co.: Summer 1928, G.M.W. 5.2. (SWT). nce Co.: 08 May 1970, D. Correll & H. Correll 38642 (TEX). Hopkins Co.: 08 Jun 1953, L. Sima 15054 (SMU,TEX-LL). Houston Co.: 10 Jun 1970, D. Correll & H. Correll 38939 (TEX). Jasper Co.: 07 Jun 1981, J. Kessler 4527 (TAES). Jefferson Co.: 21 1948, J. Brenckle 48023 (SMU,TEX). Lamar Co.: 16 Jul 1968, D. Correll & H. Correll 35913 (TEX). Liberty Co.: 25 Apr 1941, R. Crockett 937 (TEX). Nacogdoches 5 Jul 1964, FE Waller, Jr. 183 (TAES). Newton Co.: 21 May 1967, J. Crutchfield 2585 (TEX). Orange Co.: 19 Jul 1946, D. Correll 13342 (TEX). Polk Co.: 11 May 1988, S. Jones & J. Wipff 1493 (ASTC,TAES). Red River Co.: 21 Jul 1969, D. Correll 37501 (TEX). Robertson Co.: 15 Aug 1982, T: Starbuck 2974 (TAMU). Sabine Co.: 19 May 1970, D. Correll & H. Correll 38705 (TEX). San Augustine Co.: 11 Apr 1987, E. Nixon 16194 (ASTC). Shelby Co.: 17 May 1988, S. & G. Jones G E. Nixon 1376 (TAES). Trinity Co.: 25 Apr 1988, E. Nixon 16411 (ASTC,TAES). Upshur Co.: 09 Aug 1950, V. Cory 57724 (SMU). Walker Co.: 15 Jun 1968, J. Bhatt 54 (TAMU). Wood Co.: 26 Aug 1985, E. Nixon 14967 (ASTC).
2.CAREX LUPULIFORMIS Sartwell, “HOP-LIKE SEDGE”, Carices Amer. ept. Exsiccatae, 2: No. 147. 1848 (wasionym: C. lupulina Willd. var. polystachia Schwein. & Torrey). Type: (HOLOTYPE: NY; isorypes: BM, PH!).C. lupulina Willd. var. polystachia Schwein. & Torrey, Ann. Lyceum Nat. Hist. New York 1:337. 1825. Carex lurida ers var. polystachia (Schwein. & Torrey) Bailey,
Proc. Amer. Acad. Arts 22:63.
Bracts leaf-like, flat, 20-70 cm one x 4—11 mm wide, much ex-
22
ceeding the culm, antrorsely scabrous on the margins, sheathing rarely absent, septate-nodulose. Pistillate infructescence (Fig. 1b), below stam- inate spike, occasionally with staminate above, 2 —6 per culm, not aggre- gated, 2—8cm long X 1.5—3 cm wide, the uppermost usually overlap- ping for most of their length, oblong or cylindric; peduncles 1— 13 cm long, smooth, distance between 2 lowest penduncles 2 — 17 cm. Staminate inflorescence terminal, occasionally below pistillate, 1 or 2 per culm, 2— 10cm long X 2—5 mm wide, narrowly linear, peduncles 1 — 12 cm long, smooth, base of staminate spike shorter than or barely exceeding the top of the uppermost pistillate spike; anthers 2.5 — 3 mm long (based on 2 specimens). Pistillate scales 6— 13 mm long X 1.8—3.2 mm wide, lanceolate, 3 — 7-nerved, narrower and usually shorter than the perigynia, brownish-hyaline with darker stramineous centers, tapering into an awn, awn to 5.5 mm long, antrorsely scabrous. Perigynia 12— 18 mm long X 3.8—6 mm wide, ovoid, glabrous, shiny, dull-green when immature to brownish-yellow at maturity, wingless, not corky, strongly inflated, ascending to slightly spreading, sessile, 8—75 per spike, strongly 17 — 25-nerved; beak conic, 6—9 mm long, bidentate. Achenes (Fig. 2d) 3—4.5 mm long X 2.4—3.4 mm wide, rhombic, trigonous, + stipi- tate, faces concave, angles thickened internally with prominent nipple- like knobs. Distribution: Northward to Quebec, as far south as Florida and westward to Texas: Texas: by counties (Fig. 3c); regions 1 and 3, known only from Bowie and Marion counties; rare. It is never common within its range. Chromosome number 2” = G60 (Reznicek and Ball 1974). Matura- tion dates: The only Texas dates are September and October. In conjunction with specimens examined from other states and Steyermark (1968) the maturation dates are June-October. Habitat: Swampy woodlands, mostly in calcareous sites.
Representative specimens: Bowie Co.: 28 Sep 1948, E. Whitehouse 20450 (SMU). Marion Co.: Jul 1962, D. Correll 26409 (TEX).
3. CAREX LOUISIANICA Bailey, “LOUISIANA SEDGE,” Bull. Torrey Bot. Club 20:428. 1893 (based on C. halei Carey). Type: (HOLOTYPE: K).
Blades flat, 1-40 cm long X 2—6 mm wide, long-attenuate, glabrous, upper half antrorsely scabrous on the margins, septate-nodulose. Bracts leaf-like, flat, 10 — 30cm long X 2—4 mm wide, much exceeding the culm, margins antrorsely scabrous, sheathing, septate-nodulose. Pis- tillate infructescence (Fig. 1d), below staminate spike, | — 4 per culm, not aggregated, 1.5—4.5 cm long X 1.5—2.5 cm wide, subcylindric to cylindric; peduncles 0.5 —5 cm long, smooth, distance between lowest 2 peduncles 2—10 cm. Staminate inflorescence terminal, | per culm,
94
0.5—7 cm long X 1.5—3 mm wide, narrowly linear; peduncles 3 — 10 cm long, with sparse antrorse scabrosity, base of staminate spike much ex- ceeding the top of the uppermost pistillate spike; anthers 2.5 — 3.3 mm long. Pistillate scales 4.5 —6.5 mm long X 1.5 —2 mm wide, lanceolate- ovate 3— 7-nerved, narrower and shorter than the perigynia, white hyal- ine with a green center stripe, long tapering apically. Perigynia 10— 14 mm long X 3.5—6 mm wide, ovoid, glabrous, shiny, dull green when immature to stramineous at maturity, wingless, not corky, strongly infla- ted, stiffly ascending to somewhat spreading, ae 10 — 30 per spike, strongly 15 —18-nerved; beak conic, 4.5—7 mm _ long, bidentate. Achenes (Fig. 2b) 2.5 = 95 mm long X 1.7 3 mm wide, rhomboid, trigonous, broadly stipitate, faces nearly flat, angles thickened internally. Distribution: Florida to Texas, northward to Indiana and east to the mountains of New Jersey. Texas: by counties (Fig. 3d); regions 1,2,3 and
Chromosome number unknown. Maturation dates: April through August. Habitat: Swampy woods, bottomland hardwood forests, acidic soils.
Representative specimens: Bowie Co.: 06 Aug 1983, E. Nixon, J. Ward & M. McCrary 12401 (ASTC). Fannin Co.: 27 Apr 1959, K. Rochart & B. Sinclair 70 (TEX). Gregg Co.: 14 Jul 1942, C. York 5.n. (TEX). Hardin Co.: 28 Mar 1982, J. Matos & D. Rudolf 344 (ASTC). Harris Co.: 08 May 1982, J. Kessler 5684 (TAES). Jefferson Co.: 20 May 1948, J. Brenckle 48024 (SMU), 29 Apr 1936, J. Steyermark 36120 (SMU). Liberty Co.: 13 Apr 1972, R. McFarlane 27 (ASTC). Newton Co.: 18 Apr 1958, A. Traverse 525 (SMU, TEX). Orange Co.: 22 May 1988, 8 & G. Jones 1640 (ASTC, TAES). Panola Co.: 17 May 1967, J. Crutchfield & E. Nixon 2737 (TEX). Polk Co.: 12 Apr 1941, E. Girvin s.n. oie Robertson Co.: 27 Apr 1982, T: Starbuck 1853 (TAES, TAMU). San Jacinto Co.: 14 Apr 1972, E. Nixon s.n. (ASTC). Trinity Co.: 17 Jul 1936, Goodrum s.n. (TEX). ei Co.: 10 May 1941, S. Warner 9 (TEX). Wood Co.: 18 May 1988, 8. & G. Jones & E. Nixon 1426 (TAES).
4. Carex INTUMESCENS Rudge, “BLADDER SEDGE”, Trans. Linn. Soc. London 8:97. 1804. Tyee: (HOLOTYPE: BM; IsoryPE: BM).
Blades flat, 9-30 cm long X 3—9 mm wide, long-attenuate, glabrous, upper half ancrorsely scabrous on the margins, septate-nodulose. Bracts leaf-like, flat, 5.5 — 22 cm long X 2—6 mm wide, much exceed- ing the culm, antrorsely scabrous on the margins, sheathless, rarely with short sheaths, septate-nodulose. Pistillate infructescence (Fig. 1c), below staminate spike, 1—4 per culm, aggregated, 1— 2.7 cm long X 1—2.8 cm wide, globose to subglobose; peduncles 0.3 — 1.5 cm long, antrorsely scabrous, distance between lowest 2 peduncles 0.2— 2.1 cm; staminate inflorescence terminal, | per culm, 1—5 cm long X 1—3 mm wide, nar- rowly linear; peduncles 0.5 — 4 cm long, antrorsely scabrous, base of stam- inate spike may or may not exceed the top of the uppermost pistillate spike;
“PAUPBIS “JC “Suasaunjur “ ‘ag “wotupistMo, * “pC *suusofzindny
"9g wunnduy xaapy gg capurnduT] UoIIdag “eC “sexay, UI sa1uno> Aq uonnqiuasiq ‘J—e “¢ “O]q
ch Tyre rane Lo
3b
te siti i ea
Z|
96
anthers 1.7 — 2.4 mm long. Pistillate scales 4—9.5 mm long X 2—3.8 mm wide, lanceolate-ovate to ovate,(1-) 3-nerved, narrower and shorter than the perigynia, white hyaline with green centers, obtuse to awned, usually strongly cuspidate, awn to 6.5 mm long, antrorsely scabrous, Perigynia 10— 17 mm long X 2.5—7.5 mm wide, broadly or narrowly ovoid, glabrous with a satiny luster, dark olive drab green, wingless, not corky, strongly inflated, usually spreading at all angles, sessile, (1-) 4— 15 per spike, strongly 13 —23-nerved; beak poorly defined, 2—4.2 mm long, bidentate. Achenes (Fig. 2e) 3.5 —5.7 mm long X(2.2-) 2.5—3.9 mm wide, ellipsoid to obovoid, trigonous, sessile, faces convex to nearly flat, angles not thickened. Distribution: Newfoundland to southeastern Manitoba, then southward to Texas and Florida. Texas: by counties (Fig. 3e); regions 1,2,3 and 4. Chromosome number x= 24 (Wahl 1940), 2
=48 (Reznicek and Ball 1974). Maturation dates: March through September. Habitat: Swampy woods, bottomland hardwood forests, acidic souls.
Representative specimens: Anderson Co.: 08 Sep 1971, E. Nixon & R. Sniffen 3359 (ASTC). Bowie Co.: 10 May 1984, E. Nixon 13752 (ASTC). Cass Co.: 17 May 1970, D. Correll & H. Correll 38690 (TEX). Galveston Co.: 08 Apr 1975, FE Waller & J. Baum 3506 cae Hardin Co.: 28 Mar 1982, J. Matos & D. Rudolf 347 (ASTC). Harris Co.: 23 Apr
), J. Kessler 3385 (SMU,TAES, TEX). Harrison Co.: 06 Aug 1977, E. Nixon & R. Aa 7702 (ASTC). Jasper Co.: 19 Apr 1951, E. Whitehouse 25036 (SMU). Jefferson Co. : O04 Apr 1946, R. Crockett 8295 (TEX). Nueces Co.: 18 May 1933 Parks & Cory 22615 (TAES). Orange Co.: 22 May 1988, 8 & G. Jones 1634 (ASTC,TAES). Polk Co.: 25 Apr 1937, C. York & B. Tharp 43319 (TEX). Red River Co.: 29 Apr 1969, D. Correll 37127 (SMU,TEX). Sabine Co.: 30 Mar 1971, E. Nixon 2247 (ASTC). San Augustine Co.: 15 Apr 1987, E. Nixon 15877 (ASTC,TAES). San Jacinto Co.: 11 Apr 1974, D. | 1308 (TAES). Shelby Co.: 05 Jul 1956, D. Correll 15350 (SMU,TEX). Trinity Co.: 10 Jul 1988, 8. & G. Jones 1802 (TAES). Tyler Co.: 22 May 1988, 8. & G. Jones 1677 (TAES). Upshur Co.: 14 Apr 1916, M. Young 113 (TEX). Wharton Co.: 07 Apr 1939, B. Tharp 43272 (TEX). Wood Co.: 18 May 1988, 8S. & G. Jones & E. Nixon 1440 (TAES).
5. CAREX GIGANTEA Rudge, “GIANT SEDGE”, Trans. Linn. Soc. London 8:99. 1804. Type: (HOLOTYPE: BM).
Blades flat, 2-6 dm long X 5-16 mm wide, long-attenuate, glabrous, upper half antrorsely scabrous on the margins, septate-nodulose. Bracts leaf-like, flat, 3-6 dm long X 6— 11mm wide, much exceeding the culm, antrorsely scabrous on the margins, sheathing, septate- nodulose. Pistillate infructescence (Fig. le), below staminate spike, 2—5 per culm, not aggregated or only very little, 3—8 cm long X 2—3 cm wide, oblong to cylindric; peduncles nearly sessile (4) cm long, smooth, distance between lowest 2 peduncles 5 — 20 cm; staminate inflorescence terminal, 1—5 per culm, 2—8 cm long X 2—4 mm wide, narrowly
97
linear; peduncles 2—8 cm long, smooth, base of staminate spike shorter than or not much exceeding top of uppermost pistillate spike; anthers 2.5—3.3 mm long (based on 5 specimens). Pistillate scales 4.5 — 10.5 mm long X 1.5—2 wide, lanceolate to lanceolate-ovate, 3 —5-nerved, narrower and shorter than perigynia, white-stramineous hyaline with green centers, acuminate to awned, awns to 2.5 mm long, entire to slight- ly antrorsely scabrous. Perigynia 11—18 mm long X 4—6 mm wide, narrowly ovoid, glabrous, shiny, yellowish green to dark green, wingless, not corky, inflated, frequently spreading at right angles to main axis to slightly ascending, 20—75 per spike, strongly 17 —22-nerved; beak conic, 6—9 mm long, bidentate. Achenes (Fig. 2c) 2.2 — 2.6 mm long 2.7 —3.3 mm wide, obconic with subtruncate to truncate summit, trigo- nous, broadly stipatate, faces concave, angles thickened internally. Dis- tribution: Florida to Texas, northward in the Mississippi Valley to Ken- tucky, Missouri and Indiana, east and northward to Delaware. Texas: by counties (Fig. 3f); regions 1 and 2 found only in Polk and Harris counties, rare. Chromosome number unknown. Maturation dates: May through September. Habitat: Swampy woodlands, acidic soils.
Representative specimens: Harris Co.: 16 Jul 1943, E. Boon 224 (TEX). Polk Co.: 14 May 1942, E. Brinklen 42 — 160 (TEX).
DISCUSSIONS
Reznicek and Ball (1974) stated that the series is clearly divided into two groups based on external morphology of the achenes. Carex lupulina, C. lupuliformis, C. loutsianica and C. gigantea are in one group, and C, zn- tumescens and C, grayi are in the other. Menapace et al. (1986) assessed the phenetic affiliation of species in section Lupulinae by examining achene epidermal micromorphology using scanning electron microscopy. Using silica platforms with or without central bodies in conjunction with macro- morphological features, they supported the division of section Lupulinae into subsection Lapulinae (J. Carey) Kukenth., (C. lupulina, C. lupult- formis, C. louisiana and C. gigantea) ane subsection Intumescentes Menapace, Wujek and Reznicek (C. smtumescens and C. gray?).
Based on our examination of herbarium specimens C. /owisianica of subsection Lupulinae is frequently confused with C. sntumescens. Bot species grow in the same habitat and have the same basic habit. However, C. louisianica (Fig. 1d) has subcylindric to cylindric pistillate spikes (ps), perigynia ascending to slightly spreading, perigynia drying to a light olive-drab green or stramineous brown in color, and the peduncle of the staminate spike (ss) greatly exceeds the uppermost pistillate spike. Carex intumescens (Fig. 1c) has subglobose to globose pistillate spikes (ps),
98
perigynia spreading at all angles, perigynia drying to dark olive-drab green, and the staminate spike (ss) moderately surpassing the uppermost pistillate spike. Carex louisianica (Fig. 1d) can be easily separated from C. lupulina (Fig. la) by the peduncle of the staminate spike (ss) of C. lowisi- anita greatly exceeding the uppermost pistillate spike (ps). In C. /upulina the staminate spike rarely or slightly exceeds the uppermost pistillate spike.
Carex lupulina, a common species, is frequently confused with the rare C. lupuliformis. Carex lupuliformis has only been collected twice in Texas. The most recent collection was made in October of 1962. Morphologically they are similar and difficult to differentiate in the field. However, C. lupulina is most frequently found in open acidic swampy sites, swampy acidic forest edges and acidic roadside ditches associated with adjacent swamps, whereas C. /upuliformis is most frequently found in basic or calcar- eous swamps. Carex lupulina, variable in stature, can grow as large as C. lupuliformis under favorable conditions. However, C. /apuliformis is consist- ently the largest Carex of the section and one of the largest species of the genus in North America. Reznicek and Ball (1974) stated that when grown in favorable conditions, C. /wpuliformis is certainly one of the largest and most stately of Carex in Canada and also one of the rarest. The achenes of C. lupuliformis (Fig. 2d) are the best diagnostic character. They have pointed angles with nipple-like knobs and deeply concave faces. Achenes of C. lupulina (Fig. 2a) have neither pointed angles nor nipple-like knobs and the faces are flat to slightly concave. We concur with Reznicek and Ball (op. cit.) that Willdenow is the correct authority of C. /upulina.
Carex gigantea, like C. lupuliformis, has only been collected twice in Texas. The most recent collection was made in July 1943. Site locations listed on the herbarium labels for both collections are unclear making original locations impossible to find. If these two species are found to be extant in Texas, then serious consideration should be made by the Texas Organization for Endangered Species to list both species as “state endange- red species” as defined by Beaty and Mahler (1987).
ACKNOWLEDGMENTS
We thank Gretchen D. Jones for help in the overall preparation of the plates and for editorial comments. Editorial comments from Hugh D. Wilson, J.K. Wipff and David Castaner are also appreciated. We thank the curators from all herbaria from which loans were made; making this research possible. We acknowledge Texas A&M Electron Microscopy Cen- ter for the use of their equipment. This is Technical Bulletin TA 24828, Texas Agricultural Experiment Station.
a0 REFERENCES
BEATY, H.E. and WM. E MAHLER, revisors 1987. Endangered, threatened, & watch lists of plants of Texas. 2nd rev. Texas Organization for Endangered Species, Austin,
Texas.
CORRELL, D.S. and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas
FERNALD, M.L. 1950. Gray’s manual of ee 8th ed. Van Nostrand Reimhold Se ae York.
GLEASON, 1952. Illustrated flora of the northeastern United States and adjacent Canada. a Publishing Company, New York.
GOULD, EW. 1975. Texas eae checklist and ecological summary. Tex. Agric. Exp. Sta. Bull. MP-585.
HOLMGREN, PK., W. KEUKEN and E.K. SCHOFIELD, compilers 1981. Index herbariorum | E Bd. 7. W. Junk B. V., The Hague.
MACKENZIE, K.K. 1935. Cyperaceae. Cariceae. North Am. flora, 18:393 —478.
----- . 1940. North American Cariceae. New York Bot. Gard. Vol. 2. New York.
MENAPACE, EJ., D.E. WUJEK and A.A. REZNICEK. 1986. A systematic revision of the ns Carex rene an respect to the section Lupulinae. Can. J. Bot. 64:2785 — 2788.
aeeICne A.A. and PW. BALL. 1974. The taxonomy of Carex series Lupulinae in Canada. Can. J. Bot. 52:2387 — 2399.
Rg es J.A. 1968. Flora of Missouri. The Iowa State Univ. Press. Ames, lowa. SS, = 7 ee Michigan flora. Part 1. Gymnosperms and monocots. Cranbrook Inst. Sci.
WAHL, a " oe Chromosome numbers and meiosis in the genus Carex. Amer. J. Bot.
:458 — 570. WATERFALL, U.T. 1979. Keys to the flora of Oklahoma. 6th ed. Published privately, Stillwater, Oklahoma.
BOOK REVIEWS
BEAL, ERNEST O. and JOHN W. THIERET. 1986. Aquatic and Wet- land plants of Kentucky. Kentucky Nature Preserves Commission Scientific and Technical Series Number 5. Kentucky Nature Preserves Commission, 407 Broadway, Frankfort, KY 40601. Paper $20.00 + $1.50 per book for postage and handling. 315 pp. Illust- rations by Sara Fish Brown.
This is an excellent identification manual that contains keys and diagrammatic illustrations, more often than not, emphasizing and illust- rating the key taxonomic characters. The arrangement of the taxa and illustrations is simplifed by being alphabetical. It is recommended as a supplement to anyone’s library for its wide ranging application. WFM.
GRAINGE, MICHAEL and SALEEM AHMED. 1988. Handbook of Plants with Pest-Control Properties. John Wiley & Sons. The authors are with the Resource Systems Institute of the East-West Center in Honolulu, Hawait.
The volume is a computerized database with data taken from the litera- ture. “Information in this document is presented in three sections. Section I catalogues about 2,300 plants