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 having pest-control properties; Section II is a listing of about 800 pests and the plants that reportedly control them; and Section HI lists another 1,000 plants that are either poisonous in na- ture or reportedly control diseases and nematodes of humans and animals. The latter are candidate plants for screening for activity against crop pests.” This is an excellent basic reference for any comtemplated study of this subdiscipline. WFM. SCHULTES, R.E. 1988. Where the Gods Reign, Plants and Peoples of the Colombian Amazon. 308 pp. Synergetic Press, Inc., P O. Box 689, Oracle, AZ 85623. This volume is a collection of annotated black and white photographs depicting the vegetation and life of the Colombian Amazon. In addition to the Preface and Foreward, there is a very informative chapter on “Amazonia” covering the history, topography, climate, people, etc. Sipa 14(1):100. 1990. X YLOTHAMIA (ASTERACEAE: ASTEREAE), A NEW GENUS RELATED TO EUTHAMIA GUY L. NESOM, YOUNGBAE SUH, DAVID R. MORGAN, and BERYL B. SIMPSON cals of Botany, esd y Texas Austin, TX 78713 U.S.A ABSTRACT e 27 species of Ericameria sensu stricto occur primarily in the western United States and paises: Mexico. Eight species traditionally pie with Ericameria are primarily endemics of the Chihuahuan Desert region and comprise a well-defined natural group separated from the others. Although the last are Te similar to Ericameria in habit and morphology of the leaves and irae. they are strongly divergent in other aspects, particularly their zygomorphic (vs. regular) disc corollas with long (vs. short) lobes and phyllaries with an apical glandular patch but without a prominent midline (vs. no apical patch but a resinous midline). Studies of patterns of restriction site variation in chloroplast DNA corroborate the observations that these two groups are widely divergent phylogenetically and place Ericameria sensu stricto nearest Chrysothamnus and the Chihu- ahuan species closest to Exthamia. The latter Eas are ae ated as a new genus, Xylo- thamia Nesom , Morgan, & Simpson, and the following new eombinations are proposed: X. diffusa (Bench, ) Nesom, X. ee Gray) Nesom, X. parrasana (S. F Blake) Nesom, X. pseudobaccharis (S. FE Blake) Nesom, X. purpusii eee Nesom, X. riskindii (B. Turner & Langford) Nesom, and X. tiantha S, E Blake) Nesom. One new species is described: X. johnstonii Nesom. A key to the species is provided, as well as a summary of a nilicseons morphological description, a distribution map for each one. KEY WORDS: Xylothamia, Ericameria, Haplopappus, Asteraceae, Astereae, Mexico. RESUMEN Las 27 especies de Ericameria sensu stricto se encuentran distribuidas principalmente en el oeste de los Estados Unido y el noroeste de Mexico. Ocho especies que anteriormente han sido incluidas en Ericameria comprenden un grupo natural bien definido y separado de las demas especies, y son principalmente especies endémicas del desierto Chihuahuense. Aunque estas ocho especies son aaa a Ericameria en cuanto al habito y la morfologia de las hojas y la capitulescencias, se destacan fuertemente en otros aspectos, especialmente en cuanto a las corolas zigomorficas (vs. Senne ae con lébulos largos (vs. cortos) de las flores del disco, y los filarios con una mancha apical oe pero sin una linea medial prominente (vs. sin mancha apical pero con linea medial resinosa promiente). Los estudios de los patrones de variacién de los sitios de restriccion del ADN de los cloroplastos apoyan a las observaciones que estos dos grupos son fuertemente divergences filogeneticamente e indican que ae sensu stricto es mas semejante a Chrysothamnus mientras que las ocho especies del desierto Chihuahense se asemejan mas a Euthamia. Estas ultimas especies se incluyen en el género nuevo Xylothamia Nesom, Suh, Morgan, and Simpson, con las siguientes pe ee nuevas: X. diffusa (Benit. ) Nesom, X. palmeri (A. Gray) Sipa 14(1):101— 116. 1990. 102 Nesom, X. parrasana (S. E Blake) Nesom, X. ontario (S. E Blake) Nesom, X. purpusii (Brandegee) Nesom, X. riskindii (B. Turner & Langford) Nesom, and X triantha (S. E Blake) Nesom. Una especie nueva se ae X. johnstonii Nesom. Se incluyen una clave para la identificacion de las especies, asi como un resumen de la tip- ificacion, una descripci6n morfolégica, y un mapa de la distribuci6n para cada una. Hall (1928) treated as Haplopappus sect. Ericameria a group of species with a subshrubby habit, punctate-resinous, mostly narrow, entire leaves, and heads arranged in relatively compact, flat-topped capitulescences. He divided sect. Ericameria into two groups, (1) those with a paniculate or racemose-paniculate capitulescence or with solitary heads and (2) those with a regularly corymboid capitulescence. With the exception of E. dif- fusa, however, all of the species that he treated belong to Ericameria sensu stricto in the sense of the arrangement proposed in the present paper. Various authors have followed the early lead of Nuttall (1841) in recognizing Ertcameria as distinct, but most have provided little or no com- ment on their concept of the genus (e.g., Bentham 1844; Wiggins 1933; Shinners 1950). In a study that preceded his treatment of Haplopappus, Hall (1907) himself considered Ericameria distinct. Urbatsch (1976, 1978) has recently published several taxonomic studies of species groups of Ericameria as a genus, and with Wussow (1979) he transferred Haplopappus linearifolius DC. of Hall’s Haplopappus sect. Stenotopsis into Ericameria. \n his contribution to the North American checklist by Kartesz and Kartesz (1980), Urbatsch treated the genus as distinct and included some of the species of Haplopappus sect. Asiris. Jepson (1925) treated Ericameria as a separate genus, but since Hall’s monograph (1928), the only major floristic treatment to segregate it from Haplopappus has been that of Johnston (1970) for Texas. Finally, in concert with the exclusion of the group of species discussed in the present paper, Nesom (1990) has formally broad- ened Ericameria to 27 species by including those of Haplopappus sects. Steno- topsis, Asiris, and Macronema, creating a taxon coordinate in rank and vari- ability with the closely related genus Chrysothamnus. Urbatsch (1978, p. 298) noted that the Chihuahuan Desert species of Ericameria “stand apart from one another and from their California relatives in that each has a unique flavonoid complement and one or more extraordi- nary morphological features (Urbatsch, ined.). However, two characters, their zygomorphic disk corollas and their relatively long, thick style branches, give the Chihuahuan Desert species unity.” Only five species were treated by Urbatsch, and he observed that one of them, E. daricifolia, is most Closely related to the Californian species E. pinifolia and E. brachy- lepis rather than to any Chihuahuan Desert species. Johnston (1967) transferred Aster palmeri to Ericameria (as the nomen 103 novum E. austrotexana) and perceptively noted that its closest relatives 1n- cluded E. triantha, E. diffusa, E. parrasana, and E. pseadobaccharis. In his treatment of Ericameria for the flora of Texas (Johnston 1970), he made the even more remarkable observation that “There is a superficial and perhaps more than superficial resemblance of [Euthamia pulverulenta E. Greene} to Ericameria austrotexana.” In this study, we corroborate and extend Johnston's hypothesis of in- terspecific relationships and Urbatsch’s observation that the eastern (“Chihuahuan”) and western (“Californian”) species groups of Ericameria sensu lato are distinct from one another. Further, we find evidence to sup- port Johnston's speculation regarding the relationships of E. austrotexana, as discussed below. In the following discussion, the phrase “Californian” species refers to those of sect. Ericameria (Nesom 1990), of which the “Chihuahuan” Species (the group of 8 treated in this paper) have been con- sidered a part. The “Chihuahuan” species include one that is endemic to western Mexico but that is clearly related to those from the east. COMPARISON OF THE CHIHUAHUAN AND CALIFORNIAN SPECIES OF ERICAMERIA The the obvious similarities in their woody habit, narrow, resinous leaves, flat-topped capitulescences, and their base chromosome number of x=9, the Chihuahuan and Californian groups of Ericameria are sharply separated by the contrasts in the following couplet. 1. Phyllaries not basally indurated, with a discrete, orange-glandular midrib Oo ip, sometimes prominently broadened distally but without a apical glandular patch; disc Sie regular, with lobes cut 1/4-1/2 i length of the throat, all lobes of equal or nearly equal length, erect or sometimes recurved; species primarily of the Sonoran and Mohave Decent. 25.2 ots, 0 dk mes Naas a nme se ee ee Ve een he eee Californian Ericamerta Phyllaries white-indurated on the lower half with the midrib not percept- ible or else very thin, and with a prominent glandular or herbaceous- glandular patch on the distal hale. disc corollas strongly zygomorphic, with lobes deeply incised, usually nearly to the base of the throat, and strongly unequal in length, reflexing or coiling; species primarily of the Chihuahuan Desert: (Bie. Wink sitet ta ae bp bios ee eee eee ee Chihuahuan Ericameria —_— The difference in phyllary morphology is consistent and easily observable and in itself is strongly suggestive that two phylads are represented. As pointed out to us by Loran Anderson (pers. comm.), the presence or abs- ence of an apical glandular patch is not constant within Chrysothamnus, but among the species considered here, it appears to be diagnostic. The zygomorphic corollas of the Chihuahuan species are even more remarkable, because, to our knowledge, they do not occur in any other North American 104 cae aes are a aa 7 " ] Ps | | | | 4 i: | ; eS ec eceeenere diffusa LSA | HA rs \ » a triantha t— aN ; F | a ay O X. purpusii & X. pseudobaccharis @ X. parrasana & X. riskindii @ X. palmeri QO X. johnstonii FIG. 1. Geographic distribution of the species of Xylothamia. 105 Astereae. Typically, two of the sinuses are cut nearly to the base of the throat, one is very shallow, and the other two are intermediate in depth. The two lobes on either side of the shallow sinus are erect, but the other three are sharply reflexed to coiling. COMPARISON OF THE CHIHUAHUAN SPECIES WITH EUTHAMIA Euthamia is a very sharply defined genus of about eight species (Sieren 1981), although the putative species are so similar to one another that dis- agreement still exists with regard to species limits. It has been considered a part of So/zdago in the past, but recent workers (Kapoor and Beaudry 1966; Anderson and Creech 1975; Cronquist 1980) have recognized it as a dis- tinct genus. All species of Evthamia are primarily herbaceous, viscid peren- nials with a branching system of fibrous-rooted rhizomes and numerous heads usually in a compact corymboid capitulescence, although the heads may be more loosely arranged in some species (e.g., E. occidentalis Nutt.). The leaves are narrow, often 3-nerved, and punctate-resinous. The phyllaries are strongly graduated in several series, narrowly oblong- lanceolate with white, indurated bases, and have an apical herbaceous patch that is strongly viscid-glandular. The disc corollas are generally, though not always, fewer than the rays, and they are deeply lobed but regular in symmetry. As noted by Kapoor and Beaudry (1966), the anther filaments are joined to the corolla at the tube-throat junction, in contrast to most other genera with which it has been compared. The base chromosome number is x=9, and diploids, tetraploids, and hexaploids are known (Sieren 1981). One of the few floristicians yet to include species of both groups in a single treatment has noted that the the resemblance between Euthamia and the Chihuahuan species of Ericameria might be more than superficial Johnston 1970). Plants of both groups have narrow, resinous-punctate leaves, very similar phyllaries, the disc corollas are relatively deeply lobed with coiling-reflexing lobes and similar insertion of the staminal fila- ments, and all have at least a tendency to produce flat-topped capitulescences. Of the other genera that appear to be closely related on the basis of molecular evidence, Gutierrezia, Gymnosperma, Amphiachyris, and Bigelowia (Suh 1989), only the last has a chromosome number of » =9. In its herbaceous habit with leaves primarily basally disposed and its turbi- nate-cylindric heads in a densely compact corymb, it appears far less similar than Eathamia to the species of Chihuahuan Ericameria. The species of Chihuahuan Ericameria are separated from Euthamia by the differences in the following couplet. 106 _ . Plants woody subshrubs from woody roots, not rhizomatous, with stems and leaves minutely papillate in all but one species; leaves 1-nerved; heads solitary or in a loosely to compactly corymboid capitulescence; disc corollas SHONGIY 2YROMOIDUIGS ce ia cee tated ui olaseee ages Chihuahuan Ericameria 1. Plants primarily herbaceous perennials from a system of a rooted rhizomes, with stems and leaves glabrous to sparsely hairy, never papillate; leaves often 3-nerved; heads 7 a compact aie disc corollas POOWIAE, og cae eg ee Geeta se bh wee Ree ena aaS Euthamta RELATIONSHIPS ACCORDING TO MOLECULAR EvIDENCE Studies of molecular variation show that the Chihuahuan species of Ericameria are only distantly related to those of the Californian group. Rec- ently completed comparisons of North American Astereae using data from restriction site variation in chloroplast DNA by Suh (1989) and Morgan (in prep.) have each included three species of Ericameria, including the Chihu- ahuan group. In both studies, Ericameria ericoides (Californian) is most closely related to E. [sect. Macronema] discoidea and Chrysothamnus. Suh found that Ericameria austrotexana (Chihuahuan) 1s most closely similar to Euthamia, and Morgan, whose study did not include Euthamia or its close relatives according to Suh’s analysis, found Ericameria triantha (Chihu- ahuan) equally similar to Solidago, Aster, Machaeranthera, and Heterotheca. Figure 2 shows an abbreviated summary of the combined results of Su and Morgan, each of whom is preparing more detailed analyses of his data for publication. Each of the lineages shown is named as a “group” for one of the major genera that occurs within it and each group has some representat- ive members listed. Each of the six groups represented in the terminal polytomy is strongly defined, but hypotheses of relationships among them are weakly supported and an unequivocal resolution is not possible. Never- theless, the complete separation of the Chihuahuan and Californian species of Ericameria is clearly shown. Ericameria ericoides (Californian) is related to Chrysothamnus, and E. austrotexana and E. triantha (Chihuahuan) are most closely related to Euthamia of the Gutierrezia group. In summary, the hihuahuan species of Ericameria are sharply distinct morphologically from the Californian species. The former are shown by molecular data to be much more closely related to Eathamia, to which they are similar in fea- tures of capitular and leaf morphology. To account for their unique position according to both morphological and molecular evidence, we segregate this group of 7 species as a new genus. Xylothamia Nesom, Suh, Morgan, & Simpson, gen. nov. Type species: Xylothamia (Aplopappus) triantha. Aspectu Ericameriae Nutt. similis sed differt phyllariis in dimidio inferno albi-induratis nervo medio non perceptibili in dimidio superno area prominenti glandulosa vel herbacei- 107 MACHAERANTHERA group Xa ea Xylo feta ASTER group Trip HETEROTHECA group Croptilon Bradburia GUTIERREZIA group Gym i iad utha Teiothanie CHRYSOTHAMNUS group Ericameria BACCHARIS ERIGERON group Conyza FIG. 2. Phylogenetic relationships of selected sane of North American Astereae, based on ay from restriction site analyses of chloroplast DNA combined from the studies by Y. Suh and D. Morgan. Ericameria and Xylothamia are members i, different clades. landulosa et corollis disci zygomorphis lobis profunde incisis in longitudine inaequalibus Euthamiae Nutt. affinis sed habitu ligneo non rhizomato et corollis disci zygomorphis [obs profunde incisis in longitudine inaequalibus differc. Woody, evergreen subshrubs 0.2—3.0 m tall, from woody, non- rhizomatous roots; stems and leaves minutely papillate in all except X. diffusa. Stems often minutely ridged. Leaves linear-oblong to linear- lanceolate or obovate, 1-nerved, entire, flat to involute-terete, strongly to weakly or not at all punctate-resinous. Heads campanulate to turbinate- campanulate, 3 — 6 (-8) mm wide, more or less solitary and loosely aggre- 108 gated to densely aggregated in a compact, corymbiform capitulescence; phyllaries strongly graduated, narrowly oblong-lanceolate with white- indurated, enervate bases, with an apical, strongly viscid-glandular herb- aceous patch, the margins hyaline; receptacles deeply alveolate, with the alveoli margins broad to deeply dissected and linear, nearly naked in X. triantha. Ray flowers 0 — 8 (-13), fertile, the corollas yellow to white, with ligules barely extending past the phyllaries. Disc flowers 4— 22 (-50), more numerous than the ray, perfect, fertile, the corollas yellow, sometimes drying purplish, 3.0 — 5.0 (-5.5) mm long, strongly zygomor- hic, with two of the sinuses cut nearly to the base of the throat, one very shallow (1/4 — 1/3 as deep as the former), and the other two intermediate in depth, the two short lobes erect, the others reflexed-coiling; staminal fila- ments inserted at the tube-throat junction; collecting appendages of the style branches ovate-lanceolate to linear-triangular. Achenes subcylindric to turbinate, 1.5 —2.4 mm long, with 5 —8 barely discernible nerves, moderately to densely sericeous; pappus bristles persistent, in a single series. Base chromosome number, x = 9. A chromosome number of 7 = 9 pairs has been reported for X. diffusa (Pinkava and Keil 1977), X. palmeri (Urbatsch 1975), X. triantha (Anderson et al. 1974; Urbatsch 1975; Powell and Powell 1977), and X. purpusiz (Urbatsch 1975). The name of the genus is intended as a reference to its close relationship to Exthamia as well as to emphasize the relative woodiness of the plants. KEY TO THE SPECIES OF X YLOTHAMIA — Leaves spatulate or lanceolate-triangular; heads solitary. (2) . Leaves mostly linear; heads in loose to compact, cymose clusters. (3) Ye — 2. Leaves obovate-spatulate, minutely Pap pillate; heads mm wide, radiate; southeastern Coahuila, Nuevo Leon .............000.00000. X. riskindit Leaves narrowly lanceolate-triangular, Reeve hirtellous-hispidulous; heads 4—5 mm wide, eradiate; Chihuahua, Durango, Coahuila ....... X, purpusit 3. Leaves involute, appearing terete; heads eradiate or with |—3 tiny rays hidden within the involucre; disc flowers 3 — 7. (4) 3. Leaves narrow but evidently flattened; heads edie disc flowers 7 — 22. (5) 4. Stems minutely papillate-scabrous, ray flowers absent; Chihuahua, Durango, Coahuila, Nuevo Leon, and southwest Texas............... X. triantha 4. Stems glabrous; ray flowers 0 — 3; coastal and near coastal Baja California Sur, Baja California Norte, Sonora, SUMAlOA ea) itd a eee be eee hes X. diffusa 5. Leaves widely spaced and 1 ee heads in short racemes; disc flowers 7 — 14; south-central Coahuila.............0...0..00... X. pseudobaccharis 5. Leaves sare crowded and conspicuous, heads in loose corymbs; disc flowers 9 — 22. (6) 6. Stems minutely papillate-scabrous; heads in a phyllaries weakly graduated; southern Coahuila and adjacent Zacatecas .............. X. parrasana 109 oN —~ . Stems glabrous; heads in loose panicles; phyllaries strongly graduated. (7) . Leaf margins smooth; ray flowers 5 — 11; disc flowers 9 — 13; corollas white to cream; flowering August-October (-February); southeastern Texas, northern Nuevo: Peon and lamaulipas «y.evcdsesey eet kee ere sels X. palmeri Leaf margins utely scabrous-ciliate; ray flowers 12— 15; disc flowers ls = 20; eis nae flowering May-June; San Luis Potosi........ X. jobnstonti ~— ~ lL. XyLorHamia diffusa (Benth.) re comb. nov. — Ericameria diffusa Benth., Bot. Voy. Sulphur 2:23. 1844. : MEXICO. Baja CALIFORNIA SuR. Mocdatens Bay, 1839, R. B. Hinds s.n. — Alp ia DC., 1836. Solidago diffusa ae ) A. Gray, Proc. Amer. Acad. 33199-1861: Bon se (Benth.) A. Gray, Proc. Amer. Acad. Arts 8: or ie. Chrysoma diffusa (Bent Greene, ie 3:10. 1895. Linosyris sonoriensis A. Gray, Proc. Amer. Acad. Arts 8:291. 1870. Type: MEXICO. Sonora. District of the Yaqui River, 1869, E. Palmer s.n. (HOLOTYPE: GH!). Aster sonortensis (A. Gray) Kuntze, Rev. 317. 1891. Aplopappus sonortensis (A. ee E Blake, Contr. U.S. Natl. Herb. 23: 1490. 1926. —_~ Subshrubs 3 — 15 (-20) dm tall, glabrous, resinous, punctate. Leaves 2— 10(-25) mm long, involute, linear and more or less terete, spreading to ascending, sometimes upcurved or downcurved, with an apiculate, slight- ly falcate apex. Heads sessile to short-pedicellate in compact cymes, turbi- nate, 2.5—3.5 mm wide; phyllaries strongly graduated, the inner 3.0—4.5 mm long. Ray flowers 0 — 3, the corollas |— 3 mm long when present, hidden within the involucre. Disc flowers 4 — 5, the corollas 3 — 4 mm long, sometimes drying purplish. Achenes sparsely to moderately sericeous, surface not obscured. Chromosome number, 7 =9 pairs. Baja California Norte, Baja California Sur, Sonora; coastal and near coas- tal sites, sandy and gravelly plains, bottomland alluvium, dunes, in coastal scrub, salt flats, Yucca-Larrea-Pachycereus, Prosopis-Larrea, 0 — 90 (-450) m; Oct-Dec (-Jan, Apr). Distinguished by its discoid heads and terete, usually upturned leaves, which are variable in length but tend to be very short. We have not seen the type of this species, but Bentham’s description of the disc corollas as “sub- bilabiate” leaves no doubt as to its identity. There 1s some variation in the relative depth to which the deepest lobes are cut, and rare plants produce flowers with lobes of nearly equal length. Even in these, however, the lobes are much deeper than in species of Ericameria sensu stricto, the distinctive morphology of the phyllaries is apparent, and the plants have rarely been misidentified as to species. The existence of a close relationship between Xylothamia diffusa and Chrysothamnus paniculatus hypothesized by Hall and Clements (1923) was based on similarities in leaf and phyllary morphology. The latter species, along with C. seretzfolius, is unusual in Chrysothamnus in its punctate leaves 110 and phyllaries with an apical, herbaceous-glandular patch, and the two have been recognized as a separate section within the genus (Anderson 1984). Both species, however, have narrowly cylindrical heads and phyllaries in vertical files, features that ally them with Chrysothamnus. The flavonoids of Xylothamia diffusa have been studied (Urbatsch ef a/. 1976), but hypotheses of relationship among species of Ericameria (as previ- ously understood) based on flavonoid data have been undocumented (Urbatsch 1978; Clark et al. 1980) or have included only a few species (Urbatsch and Wussow 1979) 2. XYLOTHAMIA johnstonii Nesom, sp. nov. Xylothamia palmeri (A. Gray) Nesom similis sed foliis majoribus marginibus scabri- ciliatis, flosculis radii et disci numerosioribus, corollis luteis, et florescentia vernali differt. Shrubs up to 0.7 m tall, with slender, woody branches, glabrous, resi- nous, not punctate or papillate. Leaves linear to narrowly oblanceolate, (10-) 15-40 mm long, 1—2 (-45) mm wide, the margins minutely scabrous-ciliate. Heads broadly turbinate, 6-7 mm wide, on bracteate peduncles, in loose panicles; phyllaries strongly graduated, the innermost —6 mm long, with thin-hyaline margins. Ray flowers 12—15, the corollas yellow, commonly drying purplish, 4—6 mm long, with ligules 2—4 mm long. Disc flowers 16 — 20, the corollas yellow, 4—5 mm long. Achenes ca. 1.5 mm long, densely strigose-sericeous. Endemic to central San Luis Potosi; ca. 1200-1700 m; May-Jun. Type: MEXICO. 7 Luis Poros. Bagre, Minas de San Rafael, May 1911, C. A. Parpus 5021 (HOLOTYPE: GH US!). Additional ia a MEXICO. San Luis Potosi. Santa Maria del Rio, Microondas Hill, steep slope, 3 Aug 1988 (almost completely past flower and fruit), Boldt 29643 (TEX); region of San Luis Potosi, 1878, Parry and Palmer 383 (GH); 15 km NE of Guadalcazar, 22 Jun 1955, Rzedowski 6028 (US). Xylothamia johnstonii is similar to X. palmeri in its flat, linear, non- punctate leaves, strongly graduated phyllaries with thin-hyaline margins, radiate heads in loose panicles. The new species differs in its larger leaves with minutely scabrous-ciliate margins, greater number of disc and ray and disc flowers, yellow corollas (commonly drying purplish), and spring flowering. It is named for Dr. Marshall C. Johnston, who first recognized its distinctness (Johnston 1967). 3, gala ee (A. Gray) Nesom, comb. — Aster palmeri A. Gray, Amer. Acad. Arts 17:209. 1882. ae a aan 1967): UNITED STATES. Texas. [Maverick Co.:} Eagle Pass on the Rio Grande, Sep-Oct 1879, E. Palmer 516 (GH!; tsoLecrorypres: PH, US). Ieocoma ee (A. Gray) Shinners, Field & Lab. 18:27. 1950. Ericameria austrotexcana M. C. Johnston, nom. Lit uthw. Nat. 12:106. 1967; not Ericameria palmert (A. Gray) H. M. Hall. Johnston (1967) selected Palmer 516 from among several syntypes. He referred to the GH sheet as the “holotype,” abe his intention clearly was the selection of a lectotype. Bushy shrubs 0.5—3.0 m tall, the stems, leaves, and phyllaries glabrous, resinous but not punctate. Leaves linear-elliptic to narrowly oblanceolate, 5— 15 mm long, 0.8— 1.5 mm wide, the margins smooth. Heads turbinate-campanulate, 4— 5 mm wide, immediately subtended by reduced cauline leaves, solitary but loosely clustered in cymose panicles; ae strongly graduated, the innermost 4—5 mm long. Ray flowers 1, the corollas white, 4—5 mm long with ligules 2—3 mm long. ai flowers 8— 15, the corollas white to cream, 3.8—5.0 mm long. Achenes 1.5— 1.8 mm long, densely strigose-sericeous. Northern Nuevo Leon and Tamaulipas, southern Texas; 10-600 m; brushy vegetation, saline flats, coastal dunes; Aug-Oct (-Feb). Distinguished from the other species of Xylothamia in its leaf surfaces that are not evidently punctate and its white ray and disc corollas. Xylothamia palmeri, X. jobnstonit, X. parrasana, and X. pseudobaccharis have narrow, flattened leaves and appear to be closely related among themselves. Xylothamia riskindii also probably belongs with this cluster of species, but its spatulate leaves, large heads, and relatively shallower (though unequal) corolla lobes are unusual. One Mexican collection (Tamaulipas, Buena Vista “Hda” [Hidalgo?}, 21 Jun 1919, Wooton s.n., US) is anomalous and appears to show some of the features of Xylothamia johnstonii. These plants appear to belong with X. palmeri in their relatively few-flowered (8 pistillate, 8 hermaphroditic) heads and in their geographic location. Like X. johnstoniz, however, they have leaves with minutely scabrous margins, yellow corollas, and they are early flowering. 4. XyLOTHAMiA parrasana (S. E Blake) Nesom, comb. nov. — Ericameria parrasana §. E Blake, Contr. Gray Herb. 52:26. 1917. Type: MEXICO. es Sierra de Parras, rocky slopes, Mar 1905, Purpus 1005 (HOLOTYPE: GH!). lopappus parrasanus (S. FE Blake) S. E Blake, Contr. U. S. Natl. Herb. ses 192 Subshrubs 1.5—2.0 dm tall, punctate-resinous. Stems minutely scabrous with thick, translucent, short, papillose projections. Leaves flat, mostly linear-lanceolate with a slightly falcate apex, 5— 10 mm long, glabrous to papillate like the stems, with sunken glands. Heads campanu- late, 5 — 6 mm wide, short-pedicellate in a distinctly corymboid capitules- cence; phyllaries weakly graduated, the inner 3.0—3.5 mm long. Ray flowers 5 — 11, the corollas 5 mm long, the ligules 3.5 mm long, 1.0 mm 112 wide. Disc flowers 15 — 22, the corollas 3.54.0 mm long. Achenes densely sericeous. Sierra de Parras in s. Coahuila and adjacent Zacatecas; rocky slopes; (Mar-) Jul-Aug. Recognized by its flat, linear, punctate leaves, campanulate, radiate heads, weakly graduated phyllaries, and numerous disc flowers. 5. XYLOTHAMIA pseudobaccharis (S. E Blake) Nesom, comb. ee pseudobaccharis §. F Blake, J. Washington Acad. Sci. 40:47. oo Type: ME . CoaAnutta: arid limestone hills of Sierra Paila, Valle Seco, General Cepeda, a m, 4 Jul 1944, J. C. Hinton (G. B. Hinton et al. 16546) (Hototyre: US!). Ericameria psendobaccharis (S. E Blake) Urbatsch, Sida 7:299. 1978 Subshrubs 2— 10 dm tall, glabrous to minutely papillate. Leaves resi- nous but not evidently punctate, flat, linear, 2— 14 mm long, 0.5 — 1.0 mm wide. Heads solitary to sessile or short-pedicellate in short, loose racemes, Campanulate-turbinate, 3.0 — 3.5 mm wide; phyllaries strongly graduated, the inner 3—5 mm long. Ray flowers 3—6, with ligules 2.5 —3.0 mm long, 0.5 — 0.8 mm wide. Disc flowers 7 — 14, the corollas 4.0—4.5 mm long, often drying purplish. Achenes moderately sericeous. South-central Coahuila, rare; limestone or gypsum slopes; izotal; 1200 — 1500 m; Jul-Sep. Recognized by it short, widely spaced, inconspicuous leaves, the plants appearing primarily as a mass of erect, intricately branched stems. 6. XYLOTHAMIA purpusii (Brandegee) Nesom, comb. nov. — Ericameria parpusi: Brandegee, Univ. California Publ. Bor. 4:191. cn ae MEXICO. CoanHulILa: Cerro de Macho, Parpus 4479 (HoLoTyPE: UC; tsoryee: GH!). A plopappus purpusti (Brandegee) S.F Blake, Contr. U.S. Natl. Herb. 23:1491. 1926. Subshrubs 15 — 30 cm tall, the stems and leaves minutely and densely hirtellous-hispidulous; axillary fascicles of leaves often prominent on older stems. Leaves stiffly erect, thick with the thick midrib 1/4 — 1/3 as broad as the blade, narrowly lanceolate-triangular, mostly 2—5 mm long, appearing prominently short-decurrent, not punctate or resinous. Heads solitary, campanulate, 4—5 mm wide; phyllaries graduated, the inner 5 —6 mm long. Ray flowers absent. Disc flowers 8 — 10, the corollas hairy, 4.2—5.0 mm long. Achenes densely sericeous. Chihuahua, Durango, Coahuila; rocky hills of gypsum, sometimes mixed with limestone; 1100— 1200 m; May, Aug-Oct. Xylothamia purpusii, with its stiffened, lanceolate-triangular, non- punctate leaves, dense and minutely hirtellous-hispidulous vestiture, and solitary, eradiate heads is morphologically isolated within the genus. 113 7. XytorHamia riskindii (B. Turner & Langford) Nesom, comb. — Ericameria riskindii B. Turner & Langford, Madrono 29:234. ss TYPE: oo CoaHulILa: ca. 24 km E of Saltillo, S side of Sierra La Viga, ca. 6.5 km E of Jame along wood cutter's road, 10,000 ft, 15 May 1977, een et - 161566 (HOLOTYPE: TEX!; tsorypes: MEXU, RSA). Low, rounded subshrubs ca. 8— 15 cm tall, with thick, translucent, short, papillose projections, punctate-resinous. Leaves obovate-spatulate, 8— 10 mm long, with a falcate apiculum. Heads solitary, sessile to subse- ssile, broadly turbinate, 7 —8 mm wide; phyllaries obovate to ovate or ovate-lanceolate, the innermost 6.5 —7.0 mm long. Ray flowers ca. 13. Disc flowers 30 —50, the corollas 4.5 —5.5 mm long, the lobes uneven but not so strongly as in the other species. Achenes moderately strigose, the surface not obscured. Southeastern Coahuila, Nuevo Leon; limestone and gypsum areas, pine- fir-oak woodland; 2100 — 3000 m; Apr-May 8. XYLOTHAMIA tata (S. E Blake) Nesom, comb. nov. — Aplopappus trianthus 8. E ke, J. Washington Acad. Sci. 28:485. 1938. Type: UNITED STATES... Texas. es ster Co.: Chisos Mountains area, along road from Study Butte to oe 31 Aug 1937, B. H. Warnock 1126 (noLotryeE: US!). Ericameria triantha (S. E Blake) Shinners, Field & Lab. 19:133. 1951. Subshrubs 2— 10 dm tall, with minute, translucent papillae, resinous but not or only weakly punctate-glandular. Leaves 5 — 15 (-20) mm long, 0.3 —0.5 mm wide, involuted, more or less terete. Heads turbinate, 3 — 4 mm wide, short-pedicellate to sessile, in loose cymes. Phyllaries strongly graduated, sometimes in subvertical files, the inner 4—5 mm long. Ray flowers absent. Disc flowers 3 — 7, the corollas glabrous, 4—5 mm long. Achenes densely sericeous. Chihuahua, Durango, Coahuila, Nuevo Leon, and sw. Texas; gypseous, calcareous, igneous, or saline habitats, slopes or commonly desert flats with gravelly to fine alluvial soils, matorral, mesquite-creosote bush, or Suaeda-Atriplex, 700-1500 m; Jul-Oct, continuing sporadically with rain. Recognized by its reduced, terete leaves, and turbinate, eradiate, few- flowered heads in clusters. Xylothamia triantha and X. diffusa are similar in their highly reduced number of disc flowers as well as their involuted, terete leaves. The latter is the only species in the genus with a distribution primarily in western Mexico. The former is also unusual in its wide geographic range, which reaches Chihuahua and Durango. Blake (1938), in his description of Aplopappus trianthus, was the first to point out its similarity to X. diffusa. 114 Urbatsch (1978) later noted that the flavonoid complement of X. triantha is most similar to that of X. diffusa. In contrast to the notable similarity among species of Euthamia, those of Xylothamia display an extreme degree of differentiation among themselves. The species of Xylothamia with long, flat leaves as well as clustered heads, X. palmert, X. johnstonii, and X. parrasana, are perhaps the most closely related to Euthamia. Xylothamia triantha and X. diffusa, with terete leaves appear to be more distantly related, although these two species have the most Exthamia-like capitulescences of the species of Xylothamia. In Xylothamia, the ovate style appendages of the disc flowers with a minutely papillate vestiture in several species (X. diffusa, X. palmeri, X. parrasana, and X. riskindi?) contrast with those in X. triantha, which are linear-triangular with long, relatively sparse, and widely divergent collect- ing hairs. The remaining two species have appendages that are more or less intermediate in morphology between those two extremes. Xylothamia parrasana, which was included in Haplopappus sect. Stenotopsis by Hall (1928) on account of its silvery-white pappus bristles and ovate style appendages, was correctly excluded from that group by Urbatsch and Wussow (1979). Xylothamia palmeri also has white pappus and even more strongly ovate style appendages. It ts clear, however, that these similarities between Xylothamia and Ericameria cannot be considered to be strictly homologous. In all the species of Xylothamia except X. diffusa, which is completely glabrous, the stems and leaves have a vestiture of minute, translucent papillae. The papillae are pronounced and conspicuous in X. parrasana, X. triantha, and X. riskindit but poorly developed in X. pa/meri and X. pseudo- baccharis. In X. purpusii the tips of the papillae are drawn out into fine, acicular tips and, densely arranged, give the plants a_hirtellous- hispidulous appearance. Ericameria juarezensis (R. Moran) Urbatsch and E, martivensts Wiggins have short, translucent-vitreous hairs that are similar to the papillae of Xylothamia. The only species of Xy/othamia that is not resinous is X. purpusii. There is variability, however, in the occurrence of the recessed glandular- punctations that are characteristic of Ericameria. Most taxa are punctate- resinous but there are no punctations in X. purpusii, X. palmeri, or X. johnstonii. In X. diffusa, the punctations are evident on some plants but not on others. Although Hall (1928) heavily emphasized the occurrence of such “resin pockets” in his classification of sections of Haplopappus, vari- ation in their occurrence is known in other natural groups. For example, Hall himself noted that some species of Isocoma lack them, whereas they are present in others. 115 ACKNOWLEDGEMENTS We greatly appreciate the review and comments of Dr. Billie Turner and of the journal reviewers Drs. Meredith Lane and Loran Anderson. We also thank GH and US for loans of specimens and Dr. David Keil for sending fresh material of Ericameria ericoides to Y. Suh for analysis. REFERENCES ANDERSON, L. C., D. W. KYHOS, T. MOSQUIN, A. M. POWELL, and P. H. RAVEN. 1974. Chromosome numbers in Compositae. IX. Haplopappus and other Astereae. Amer. J. Bot. 61:665 — 671. ANDERSON, L. C. and J. B. CREECH. 1975. aaa leaf anatomy of Solidago and related Asteraceae. Amer. J. Bot. 62:486— 493. ANDERSON, L. C. 1984. An overview of the genus Chrysothamnus (Asteraceae). Pp 29 — 45 in Proceedings — Symposium on the biology of Artemisia and Chrysothamnus. Provo, Utah. BENTHAM, G. 1844. The botany of the voyage of H. M. S. Sulphur. London BLAKE, S. E 1938. Eleven new American Asteraceae. J. Washington Ac: a Sci. 28:478 — 492. CRONQUIST, A. 1980. ee ‘Vol. 1, Flora of the southeastern United States. Univ. North mae Press, Cha HALL 1907. C ot Gece) HALL, H. aE and E E. CLEMENTS. 1923. The phylogenetic method in taxonomy: The North American species of Artemisia, Chrysothamnus, and Atriplex. Carnegie Inst. Publ. 320: 135 HALL, H. M. 1928. Sect. Ericameria, in The genus eee ee study in the Compositae. Carnegie Inst. Washington, Publ. JEPSON, W. L. 1925. Manual of the flowering plants of nee cae California Press, Berkeley. JOHNSTON, M. C. 1967. Ericameria austrotexana M. C. Johnston (Compositae), nomen novum. Southw. Nat. 12:106— 109. 1970. Ericameria. Pp. 1577 — 1578, in Correll, D. S. and M. C. Johnston. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, 5 ompositae oe southern California. Univ. California Publ. Bot. Texas. KAPOOR, B. M. and J. R. BEAUDRY. 1966. Studies on Solidago. VU. The taxonomic status of the taxa Brachychaeta, Brintonia, C alge Enea, Oligonenron and Petradoria in relation to Solidago. Canad. J. Genet. Cytol. 8:422 — KARTESZ, J. T. and R. KARTESZ. 1980. Cn of the vascular plants of North ~ America. Univ. North Carolina Press, Chapel Hill. MORAN, R. 1969. Five new taxa of hea ad gee mia from Baja California, Mexico. Trans. San Diego Soc. Nat. Hist. 15:149 — 164. MUNZ, P A. 1974. Sa al Pp. 174— 181 in aie of southern California, Univ. of California Press, Ber NESOM, G. L. 1990. a summary of Ericameria (Asteraceae: Astereae), with the inclusion of Haplopappus sects. Macronema and Asiris. Phytologia 68: 144 — 155. NUTTALL, T. 1841. Descriptions of new species and genera of plane) in the natural order of the Compositae. Trans. Amer. Philos. Soc. ser. 2. 7:283— 453. 116 PINKAVA, D. J. and D. J. KEIL. 1977. ea counts of Compositae from the United States and Mexico. Amer. J. Bot. 64:680 — POWELL, A. M. and S. A. POWELL. 1977. ee numbers in Asteraceae. Madrono 25:160 — 169. SHINNERS, L. H. 1950. Notes on Texas Compositae — IV. Field & Lab. 18:25 — 32. SIEREN, D. J. 1981. The taxonomy of the genus Euthamta. Rhodora 83:551—579. SUH, Y. 1989. Phylogenetic studies of North American Astereae (Asteraceae) based on chloroplasts DNA. Ph.D. dissertation, Univ. Texas, Austin. URBATSCH, L. E. 1975. First chromosome number reports for some Compositae. Southw. Nat. 19:283 — 285. 1976. aes of the Ericameria cuneata complex (Compositae, Astereae). Madrono 23:338 — 345. , I. J. MABRY, M. MIYAKADO, N. OHNO, and H. YOSHIOKA. 1976. Flavonol peri ethers om Ericameria diffusa. Phytochem. 15:440— 441. URBATSCH, L. E. 1978. The Chihuahuan Desert species of Ericameria (Compositae: Aecrae: Sida 7: ome — 303. and J. linearifolius Tee — sistance: Brittonia 31:2 GINS, I. L. 1933. New plants from Baja California. Contr. Dudley Herb. lt6f=176.1 7 WUSSOW, 1979. The taxonomic affinities of Haplopappus at 5 980. Haplopappus. Pp. 266 — 270 in Flora of Baja California. Stanford Univ. Press, see California. THE HERBACEOUS FLORA OF THREE WECHES FORMATION OUTCROPS IN EASTERN TEXAS ROBERT J. GEORGE and ELRAY S. NIXON Department 2 Biology, Stephen FE. Austin State University Nacogdoches, TX 75962, U.S.A. ABSTRACT The Weches Geologic Formation, which was formed during the Eocene Epoch, supports a herbaceous flora in a region where forest is the general climax vegetation. Soils are shallow and basic in contrast to the deeper acid soils of eastern Texas and the sites are usually waterlogged during spring. Species with the highest importance values in the Weches plant communities are Sedum ee bellum, Satureja arkansana, ee ee Arenaria patula, Valerianella 7 and Trifolium dubium. Lesquerella pallida a eavenworthia lexana are endemic to these outcrops. Disjunct species include Chie pial ie Liatris mucronata, Pa virginica and Petalostemon Pees sile Species richness sa from 19 to 81 species for the sites studied, while species diversity ranged from 3.23 to 4.56. Although the Weches sites ees many species in common with ane ney plant communities in the southeastern United States, overall similarity was generally low. RESUMEN La formacion geolégica del Weches, formada durante la época Eocena, sostiene una flora herbacea en una regi6n donde el bosque es el climax vegetal. Los terrenos son poco profundos y basicos en contraste con los terrenos mas profundos y acidos del Este de Tejas; los sitios He lo ponets al estan an cies durante la primavera. — ores mas valiosas e impo en las c nidades de plantas del Weches son Sed: Sa tarka po ! Sand, ‘ ne vaginiflorus, rae ee weed as y Tm iin. E eoanes pallida y Leavenworthia texana son endémicas al Weches. Las es dislocadas inc Calylophus drummondianus, Liatris mucronata, Se nee y Petalostemon seca rimum. La riqueza en especies varié a 49 a 81 especies en los sitios a ecu ma : ntras la diversidad de especies varid de 3.23 a 4.56. Aunque los sitios del uchas especies en comun con la comunidad de cedros en la regi6n sudeste de los Estados Unidos, la semejanza en general no es muy grande. a < < =) INTRODUCTION The geological deposits of eastern Texas are quite interesting because they result from activities of Gulf of Mexico waters and continental rivers (Sellards et al. 1932). Marine deposits were laid down when oceanic waters advanced over the land. When these waters receded, rivers deposited sedi- ment seaward. Thus, there are several layers of marine deposits alternating with terrestrial deposits. The Weches Geologic Formation is a marine deposit formed during the Eocene Epoch. It extends from Sabine County near the Louisiana border to Atascosa and Frio counties in south central Sipa 14(1):117 — 127. 1990. 118 Texas in a line generally paralleling the coast. The formation is composed of glauconitic clays, marl and rich fossiliferous deposits. Mud stone often 1s associated with the Weches. Fresh exposures of the glauconitic stratum have an olive green cast, but they turn reddish-brown with age. These ex- posures usually occur naturally on the slopes of hills as a result of erosional activity. Generally, outcrops are about five to 20 m wide and occur in 1sola- ted or segmented strips, usually not more than 100 m in length. Conditions associated with these outcrops are unique for East Texas. Because of the presence of mudstone, the soils are generally rocky and shallow, precluding the growth of woody vegetation. Hence, these sites are conducive to the growth of herbaceous heliophytes. In contrast, the predo- minant vegetation in East Texas is generally a pine-hardwood forest. Weches soils are basic and calcareous as a result of the rich marine fossil component (Sellards et al. 1932). Because of the glauconitic clay stratum, outcrops are often very wet, especially in early spring. Downward percolat- ing water from overlying soils moves laterally over the impermeable clay of the Weches until it exits on hillsides where outcrops occur. On the other hand, soils frequently become very dry in summer and fall due mainly to their shallowness. These characteristics are in contrast to the generally acid, deep, well drained sandy loam soils of East Texas. Interest in the floristics of Weches outcrops began in 1981 when E. S. Nixon and J. R. Ward rediscovered a population of a white flowered mustard, Lesquerella pallida, on an outcrop near San Augustine, Texas (Nixon et al. 1983). This species, which is endemic to outcrops of the Weches Formation, had not been seen since its initial collection in the 1830's. Since little was known concerning the plants that grow on the Weches Formation, we analyzed the herbaceous vegetation and compiled a list of plants of three naturally occurring outcrops. STUDY SITES The three study sites, located in San Augustine County in eastern Texas, are within Gould’s (1975a) Pineywoods Vegetational Area and Braun's (1950) Oak-hickory Forest Region. The topography of the area 1s characterized by gently rolling hills. Average precipitation, which is fairly evenly distributed throughout the year, is about 122 cm and average annual minimum and maximum temperatures are about 12° to 25° C respectively (Larkin and Bomar 1983). January, February and March preci- pitation was below normal during the study year, 1986. This caused a reduced seepage flow over the Weches in early spring and thus affected the number of individual plants present. In contrast, May and June experien- ced above average rainfall. Lg? The three study sites are located within pastures on gently sloping hills. Mudstone is exposed at all three sites. The Weches outcrops studied are about 60 m long and up to 20 m wide. Soils over mudstone generally are less than 15 cm deep, whereas those associated with the glauconitic clays are deeper. Some woody vines, shrubs and small trees occur on deeper soil but the sites are generally open. Some of the more common woody species are Gleditsia triacanthos, Liquidambar styraciflua, Rosa bracteata, Juniperus virginiana, Forestiera ligustrina, Rhamnus lanceolata, Cornus drummondit and the vine Czssus incisa. The pastures have been mowed and grazed by cattle for many years. These factors undoubtedly have influenced the flora of the Weches outcrops. Topography and lack of woody plants generally hindered or did not require the mowing of the study sites. Pastures are usually mowed to impede woody plant invasion. METHODS AND PROCEDURES The herbaceous flora was analyzed using 50 x 100 cm quadrats. Two transects were established at each of the three study sites. The transects were parallel to the length of the Weches outcrops; and sample plots were placed every two meters along the transects. Thirty quadrats systematically were placed at each site in January of 1986. During the growing season (March through October) of 1986, quadrats were visited approximately every two weeks. During each visit, all plants were identified, and those in plots counted. Because Valerianella radiata and Valerianella stenocarpa can be distinguished in the field only by their small fruits (about 2 mm long), we lumped the two species. Based on collecting information, it is likely that V. radiata is the most abundant of the two species. For each species, frequency and density data were used to calculate relative frequency and relative density, which then were summed to give an importance value. Voucher specimens of all species on the three sites are on deposit in the Stephen E Austin State University Herbarium (ASTC). Nomenclature follows Correll and Johnston (1970), Gould (1975b) and Kartesz and Kartesz (1980). Sorenson’s index (IS = 2C/A + B) was used to compare floristic similarity of the vegetation of the three Weches sites and of the Weches vegetation with the vegetation of cedar glades in the Southeastern United States. In this index, C is the number of species common to the two communities being compared, A is the total number of species in one community and B the total number of species in the other community. Species diversity for the three Weches sites also was computed using the Shannon-Weiner diversity index (Shannon and Weaver 1949): i 2D pi 120 log> pi, where pi is the decimal fraction of the individuals belonging to the i" species. Soil samples were collected from the upper 15 cm of soil at the three study sites and pH, phosphorous, potassium, calcium, magnesium, and texture were determined by personnel at the Stephen E Austin State University soil testing laboratory. RESULTS Soils Soil pH at the three sites ranged from 7.6 to 8.2 and averaged almost 8. Levels of calcium (> 2500 ppm), potassium (>250 ppm), and magnesium (>250 ppm) also were high. Available phosphorus ranged from 10 to 12 ppm. Soil textural class ranged from sandy loams to sandy clay loams. Herbaceous Flora Plants began flowering on the Weches outcrops during March, with the spring flora consisting primarily of Satureja arkansana, Sedum pulchellum, Valerianella spp., Arenaria patula and the introduced clover Trifolium dubium (Table 1). These taxa made up 62 percent of the density and 45 percent of the importance value of the spring flora. From March through May, 59 taxa flowered and 12,734 plants were recorded in the quadrats. These plants averaged 284 individuals per m°. The spring flora of the three Weches sites varied in species richness rang- ing from 45 taxa at site | co 35 taxaat site 3. Although site 3 had the fewest taxa, it had the highest number of plants (4,655), and thus the highest average number of plants per m* (314). Valerianella spp. and Satureja arkansana were the most important species at site 1; Plantago virginica, Trifolium dubium and Melilotus indicus were of secondary importance. Trifo- lium dubium was prevalent at site 2, along with Arenaria patula, Plantago avistata, Bromus japonicus and Cerastium glomeratum. Sedum pulchellum, Satureja arkansana and Arenaria patula dominated site 3. Fifty-three per- cent of the species on Weches outcrops flowered in the spring. Species richness and density were lower in summer (June through August) than in spring. Species richness declined to 29 species and density to 23 plants per m* (Table 1). Only 1,021 plants were recorded in quadrats. During the summer, the Weches flora was composed primarily of Croton monanthogynous in association with Euphorbia nutans, Cynodon dactylon, Helenium amarum and Euphorbia maculata (Table 1). Croton monanthogynous was important at all three sites. Other species with high importance values at site | were Euphorbia nutans, Cynodon 121 dactylon, Petalostemon pulcherrimum and Palafoxia rosea. At site 2, Helenium amarum, Cynodon dactylon and Paspalum notatum were dominants where as the more important species at site 3 were Leucospora multifida, Euphorbia maculata and Croton capitatus. Grasses dominated the fall (September through November) flora of Weches outcrops, with Sporobolus vaginiflorus being the principal species (Table 1). Lespedeza striata was the only non-grass species among the top five dominants. These five dominants comprised 71 percent of the import- ance value and 73 percent of the density. Species present averaged 39 plants per m” , nearly double the number present during summer (Table 1). There were 1,729 plants recorded in the 90 quadrats representing 24 species. Sporobolus vaginiflorus was the most important species at site 1. Associated species were Digitaria ciliaris, D. ischaemum, Aristida oligantha and A. dichotoma. These same species dominated sites 2 and 3 along with Panicum hallii at site 2 and Lespedeza striata at site 3. Sporobolus vaginiflorus had an importance value of 143 at site 3. In summary, 112 herbaceous taxa were recorded in quadrats at the three study sites; site 1 had 81 taxa, site 2 76 and site 3 49. The 90 quadrats contained, at one time or another during the growing season, 15,484 plants. Dominants at the Weches sites included weedy introduced species such as Trifolium dubium, Cynodon dactylon, D. ischaemum, Lespedeza striata, Bromus japonicus and Cerastium glomeratum, the widespread rock outcrop plant Sedum pulchellum, plants that are indicative of wet sites like Satureja arkansana, and plants such as Valertanella radiata, Sporobolus vaginiflorus and Croton monanthogynous which grow on more mesic to dry sites. Most 0 the dominant taxa are rather small plants. Forty-five herbaceous species were found on the Weches study sites in addition to the 112 recorded in quadrats (Table 2). Of these 157 taxa, 81 (52%) were annuals or biennials and 76 (48%) were perennials. These 157 taxa represent 39 plant families. The three largest families are the Poaceae (38 taxa), Asteraceae (16), and Fabaceae (14). These three families account- ed for 43% of the Weches outcrop species. Of the species recorded in quadrats, 53% flower in the spring, 27% in summer and 20% flower in fall Indices of Similarity and Species Diversity The index of similarity was 0.62 between sites | and 2, 0.63 between sites 2 and 3, and 0.52 between sites | and 3. Twenty eight taxa, most of which are weedy species, occurred at all three sites. The species diversity index was 4.52 at site 1, 4.56 at site 2 and 3.23 at site 3. b22 Tasce 1. Frequency, density and importance value data for during spring, summer, and fall. } } f three Weches outcrops Relative Species Frequency Frequency Density Relative Imp. % % No/M? onsi Value! Spring anne oa May) Satureja ark 62.2 6.19 44.28 16.26 22.45 Sedum pu hellum 44.4 4.42 41.54 15.25 19.67 Trifolium dubium 73.3 7.30 31.38 11.52 18.82 Valerianella spp 52.2 5.20 26.80 9.84 15.04 Arenaria pat 42.2 4.20 26.42 9.70 13.90 Others? 72.64 114.16 37.44 110.08 Totals 99.95 284.58 100.01 199.96 Summer (June through August) Croton monanthogynous 54.4 21.68 5.94 26.15 47.83 Euphorbia s 36.7 14.60 1.72 7.54 22.14 Cynodon dactylon 16.7 6.64 2.92 12.83 19.47 Helenium amarum 12.2 4.87 2.88 12.73 7.60 Euphorbia maculata 18.9 7.52 1.16 5.09 12.61 Others 44.66 8.08 35.69 80.35 Totals 99.97 22.70 100.03 200.00 Fall (September ea November) Sporobolus va, or 75.6 22.74 14.92 38.81 61.55 Digitaria ciliaris 44.4 13.38 5.46 14.23 27.61 Digitaria ischaemum 41.1 12.37 4.24 11.05 23.42 Lespedeza striata 25.6 7.69 2.66 6.94 14.63 Aristida oligantha 26.7 8.03 2.34 6.07 14.10 Others? 35.75 9.84 22.92 58.67 Totals 99.96 39.46 100.02 199.98 'Sum of relative frequency and relative density. Other species recorded in plots at the study sites: Acalypha virginica Allium canadense ta Astranthium integrifolium oerhaavia erecta Bothriochloa saccharoides Bouteloua curtipendula Briza minor Bromus japonicus Calylophus drummondianus Lesquerella pallida Leuco Palafoxia rosea Panicum anceps Panicum flexile Panicum hallit 123 (TasLe | cont.) Carex muhlenbergii Paronychia virginica Cassia fasciculata Paspalum dilatatum Cenchrus incertus Paspalum notatum Cerastium glomeratum Paspalum setaceum Chaerophyllum tainturieri es pulcherrimum onyza Canadensis Phalaris caroliniana Croton capitatus Physalis viscosa Croton glandulosus Plantago aristata Cuphea viscosissima Plantago virginica Cyperus flavescens ‘oa annua yperus ovularis Polanisia dodecandra Dichanthelium laxiflorum Portulaca oleracea Diodia teres Pyrrhopappus multicaulis Dracopis amplexicaulis Ranunculus parviflorus Eleocharis compressa Sabatia campestris Eragrostis hirsuta Salvia lyrata Eragrostis intermedia Setaria geniculata Erig strigosus Sherardia arvensis Euphorbia dentata Solanum carolinense Euphorbia spathulata Sphenopholis obtusata Galactia volubilis Sporobolus asper alium virgatum Stachys crenata Geranium carolinianum Stipa leucotricha Geranium dissectum Tridens flavus Hedeoma hispidum Trifolium repens Hedyotis crassifolia Triodanis perfoliata Hedyotis nigricans Trisetum interruptum Heliotropium tenellum Verbena brasiliensis Hordeum pusillum Verbena halei Hypericum drummondii Veronica arvensis Krigia occidentalis Vicia angustifolia Leavenworthia texana Vulpia octoflora Lepidium virginicum Endemic, Disjunct and Rare Species Only two of the 157 taxa on Weches sites in East Texas are endemic to Weches outcrops. One, the white flowered mustard, Lesquerella a listed as endangered by the U. S. Fish and Wildlife Service (U.S. Dept. of the Interior 1987). Only five populations of this species are known to exist, all in San Augustine County, Texas. With the designation of the Texas populations of Leavenworthia aurea as L. texana by Mahler (1987), this new taxon 1s now endemic to East Texas Weches outcrops. Although Sedum pulchellum is found only on Weches outcrops in eastern Texas, it grows on rock outcrops elsewhere in the southeastern United States (Clausen 1975). Weches disjuncts include Calylophus drummondianus, Liatris mucronata, and Paronychia virginica, which are disjunct from the Edwards Plateau 124 (about 380 km to the southwest) and north central Texas (about 328 km to the northwest) (Correll and Johnston 1970). Another disjunct, Petalostemum pulcherrimum, is disjunct from central Texas about 225 km to the west (Correll and Johnston 1970). Other species considered rare in eastern Texas are Heliotropium tenellum, Eleocharis compressa and Cuphea visco- sisstma (Correll and Johnston 1970). DISCUSSION In contrast to the generally acid soils of eastern Texas bottomlands (pH 4.4 to 4.8, Nixon 1986), mesic uplands (pH 4.4 to 4.6, Nixon et al. 1980), and dry sandy uplands (pH 4.6 to 6.2, Ward 1984), Weches soils are basic with pH ranging from 7.1 (Nixon et al. 1983) to 8.2 (this study). Calcium content, likely a result of the fossilized shells of marine organisms, is much higher than that of the surrounding forest soils (Nixon et al. 1980, Marietta and Nixon 1983, Nixon et al. 1987). In addition, Weches outcrops are partly characterized by shallow soils over mudstone. The shallow depth and dry conditions of summer generally preclude woody plant establishment. Some trees, shrubs and woody vines are present on the outcrops, but only in pockets or areas where deeper soils occur. Shallow soils also are characteristic of cedar glades in the southeastern United States, where they form over limestone and dolomite (Baskin et al. 1968, Baskin and Baskin 1988). Thus both the Weches and cedar glade communities, which are dominated by herbaceous species, are examples of edaphically controlled plant communities (Baskin and Baskin 988) Because Weches Formation outcrops in eastern Texas usually contain mudstone, communities growing on them can be classified as rock outcrop communities. These types of communities have received much attention in the southeastern United States (e.g. Baskin and Baskin 1985a, Baskin and Baskin 1988). Where limestone or dolomite is at or near the surface they are called cedar glades (Baskin and Baskin 1985a). Since eastern Texas 1s within the Eastern Deciduous Forest (Braun 1950), comparisons were made of Weches and cedar glade communities. Comparisons indicate some floristic similarity. All of the Weches dominants (Table 1), with the excep- tion of Trifolium dubium, Euphorbia nutans, Lespedeza striata, Digitaria ciliaris and D. ischaemum are present in cedar glade communities (Baskin et al. 1968, Baskin and Baskin 1975a, Somers et al. 1986, Bridges and Orzell 1986). Quarterman (1986) noted that the thinner soils of Tennessee glades are dominated in the spring by Leavenworthia spp., Arenaria patula and Sedum pulchellum, and that Sporobolus vaginiflorus is a dominant grass on these soils during the summer. 125 Taste 2. Herbaceous species recorded from outside the plots at the study sites. Alophia drummondii Andropogon glomeratus berosa m manthium sessiliflorum yeas strigosus Delphinium vimineum Desmodium marilandicum Dichanthelium angustifolium arpa Geum canadense Ipomopsis rubra i odium occidentale 2 2 a Petalostemon multiflorum Phlox p Physalie ei Prunella vulgaris Ranunculus fascicularis a Sisyrinchium langloisu Sisyrinchium Sali Spiranthes ce Verbesina virginica Viola pratincola Viola rafinesquii Lists of species found on southeastern glades also were compared with our combined Weches list using Sorensen’s index of similarity. Indices of similarity between cedar glade communities in middle Tennessee and the Weches were .26 (Bridges and Orzell 1986) and .25 (Baskin et al. 1968, Baskin and Baskin 1975a). Glades in Kentucky were less similar with in- dices of .17 (Baskin and Baskin 1975b) and .16 (Baskin and Baskin 1985b). Plant families most representative of the herbaceous vegetation of Weches outcrops are the Poaceae, Asteraceae, Fabaceae and Euphorbiaceae. These same families are principal components of cedar glade communities in middle Tennessee (Somerset al. 1986). About one-half of the Weches outcrop species are perennials, whereas 70% percent of the 414 taxa of cedar glade communities in the southeastern United States are perennials (Baskin and Baskin 1985a). Fourteen percent of the 157 Weches site taxa are introduced whereas 20% of the cedar glade taxa are introduced (Baskin and Baskin 1985a). Of over 400 taxa of vascular plants growing on cedar glades in the southeastern United States, 29 are endemic to those sites (Baskin and Bas- kin 1985). Only two of the 157 Weches taxa in eastern Texas are endemic 126 to Weches sites (Nixon et al. 1983). Baskin and Baskin (1985a) found that | all of the endemic annuals were winter annuals and flowered in the spring. The two Weches endemics are annuals that flower in the spring. Baskin and Baskin (1988) noted that light, rather than soil or lack of genetic vari- ability, seems to be the most important factor governing the distribution of annual glade endemics. Another interesting aspect is that narrow endemics such as those of glades seem to produce large seed banks to ensure their continuance (Baskin and Baskin 1978), which also seems to be the case with Lesquerella pallida. During a dry spring on a particular site less than 100 plants will occur as compared to 3000 to 4000 plants during a favorable wet spring. ACKNOWLEDGEMENTS We appreciate the helpful comments and suggestions provided by Jerry M. Baskin. We also thank Melissa Miller for providing us with a Spanish abstract. REFERENCES BASKIN, C. C. and J. M. BASKIN. 1975a. Additions to the herbaceous flora of the Middle Tennessee cedar glades. J. Tennessee Acad. 1975b. The cada glade flora of Bullitc County, Kentucky. Castanea 4:184— 190. BASKIN, J. M. and C. C. BASKIN. 1978. The seed bank in a population of an endemic plant species and its ecological significance. Biol. Conserv. 14:125 — 130 85a. Life cycle ee 2 of annual plant species of cedat glades of southeastern United States. p 371 sae White ed. The population structure of vegetation. Dr. ea Publishers, ce 85b. A floristic study of a st glade in Blue Licks Battlefield State Park, eae Castanea 50:19 — 25 1988. Baemics of rode outcrop plant communities of unglactated eastern United States: an aaa of the roles of the edaphic, genetic and light factors. J. Biogeogr. Fi 829 — 84( BASKIN, J. M., E. QUART aN and C. CAUDLE. 1968. ry check-list of the herbaceous vascular plants of cedar glades. J. Tennessee Acad. COD maa ie U . L. 1950. Deciduous forests of eastern North America. a Bladen Co., Se Pennsylvania IDGES, E. L.a L. ORZELL. 1986. ees pater of the non-endemic flora of Middle ee limestone glades. ASB Bull. 33:155 — 166. CLAUSEN, R.T. 1975. Sedum of North America ce a the Mexican Plateau. Cornell University Pies. Ithaca, New York. CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas ae Foundation, Renner. GOULD, E W. 1975a. Texas plants: A Sie and ecological summary. Texas Agric. Exp. Sa: Publ. MP-585, College Station. 1975b. The grasses of on Texas A&M University Press, College Station. 127 KARTESZ J. T. and R. KARTESZ. 1980. A synonymized checklist of the vascular flora of the United States, Canada and Greenland. The Univ. of North Carolina Press, Chapel Hill. LARKIN, E J. and G. W. BOMAR. 1983. Climatic atlas of Texas. Texas Department of Water Resources, Aus MAHLER, bs ‘ 1987. ie orthia texana (Brassicaceae), a new species from Texas. Sida 9 2 ag MARIETTA, cs cn and E. §. NIXON. 1983. Vegetational analysis of a post oak-black hickory community in eastern Texas. Texas J. Sci. 35:197 — 203. NIXON, E. S. 1986. nies hardwood community structure in East Texas. p 8— 19. In C. A. McMahan and Roy G. Frye, eds. Bottomland Hardwoods in Texas. Wildlife division, Texas ae and Wildlife Department, Austin. K. MARIETTA, R. O. LITT LECH and H. B. WEYLAND. 1980. Woody ene ts American beech (Fagus grandifi ity in eastern Texas. Castanea ee = 180. MATOS, and R. S. HANSEN. 1987. The hee of woody vegeta- tion to a topographic gradient in eastern Texas. Texas J. S 367 J. WARD and . LIPSCOMB. i983. Ree of Lesquerella re (Cruciferae). Sida 10:167 — QUA RMAN, E. 1986. Biota, ee and ecological history of cedar glades. ASB <i ps 124-127. SELLARDS, E. H., W. S. ADKINS and E B. PLUMMER. — The geology of Texas. et I. Stratigraphy. Univ. Texas Bull. No. 3232, Aus SHANNON, C. E. and W. WEAVER. 1949. The mathematic: ‘ don of communication. The a of Illinois Press, Urbana. SOMERS, P, L. R. SMITH, P. B. HAMEL and E. L. BRIDGES. 1986. Preliminary pee i: plant communities and seasonal changes in cedar glades of middle Tennessee. A 33:178 U.S. Bee CnneROn 1987. Endangered and threatened species. Federal Register 52(47):7424 — 7426. WARD, J. R. 1984. Woody vegetation of the dry uplands in East Texas. Master's thesis. Stephen FE Austin State University, Nacogdoches, Texas. BOOK REVIEWS TURNER, C.E., B.S. URBANEK, G.M. WALL, C.W. WALLER. 1988. Cocaine, an Annotated Bibliography. Vols. 1 & 2. University Press of Mississippi, 3825 Ridgewood Road, Jackson, MS 39211. Hardbound $125.00 (set). Vol. 1, 1—798 pp; Vol. 2, 799 — 1364 Pp- “In Volume 1 an introductory section provides a perspective on cocaine and cocaine-related chemistry. The bibliography begins with the pre- 1950 references. The annotated section includes 4,055 annotated citations covering international scientific publications from 1950 through 1986. Entries are arranged alphabetically by author. Unsigned articles are listed under anonymous. Patents, books and book chapters are also listed by author.” “The author index and an extensive, cross-referenced subject index are contained in Volume II. ...The pre-1950 citations are indexed by author only. The 1950 - 1986 references are indexed by author and subject.” MOHLENBROCK, R.H. 1990. The illustrated flora of Illinois flowering plants: Nightshades to Mistletoe. 225 pp, 100 illus. Southern Illinois University Press, P O. Box 3697, Carbondale, IL 62901. ISBN 0-8093-1567-X. Hardbound. This 1s the latest volume in the continuing series on the flora of Illinois. This book treats the following families: Solanaceae, Convolvulaceae, Cuscutaceae, Polemoniaceae, Campanulaceae, Celastraceae, Santalaceae, and the Viscaceae. HUNTER, CARL G. 1984. Wildflowers of Arkansas. 296 pp, 484 color photographs. The Ozark Society Foundation, P O. Box 3503, Little Rock, AR 72203. The purpose of this publication is to include a comprehensive cross sec- tion of the wildflowers from over the entire state. In addition to the descriptions of the colored plates, there are chapters on “History of Botani- cal Investigations,” “Family Descriptions,” and several other chapters covering the state of Arkansas and general botanical information regarding nomenclature and terminology. Sipa 14(1):128. 1990. NOTES A NOMENCLATURAL NOTE ON EUPATORIUM FISTULOSUM (ASTERACEAE)! — Expatorium fistulosum Barratt ranges from “s Me to Iowa, s toc Fla, Ala, Miss, La, and Tex” (Cronquist 1980). Barratt (184 1) separated E. fistulosum from E. purpureum L. He described the former species and few other eupatoriums with whorled leaves in a single folio page pub- lication (fig. 1), titled Eapatoria verticillata. Robinson (1904), probably unaware of Barratt’s Expatorinm fistulosum, described a new Expatorium species from Costa Rica, as E. fistulosum. Robinson's E. fistulosum is a later homonym and illegitimate, which was corrected by B. Robinson (193 1) by renaming his Costa Rican Expatorium sp., as E. angulare B. Robins. King and H. Robinson (1970) treated E. angulare as a synonym of Neomirandea angularis (B. Robins.) King & H. Robins. Barratt intended to distribute his single folio page publication accompa- nied with a set of exsiccatae among his friends and botanists. This informa- tion is found in figure 1. It is evident from ICBN article 31 (Greuter 1988), that the above procedure, practiced by Barratt in this case, would validate a name if done prior to Jan 1953. We believe that Barratt did distribute his single folio page publication to other botanists. Wood (1847) stated that his treatment of Expatorium was adopted from Barratt’s Eupatoria verticillata. Jackson (1881) and B. Robinson (1931) cited Barratt’s 1841 publication. The name E. fistulosum was attributed to Barratt by several authors such as Mackenzie (1920), Weatherby (1921), Wiegand and Weatherby (1937), Deam (1940), Fernald (1950), Gleason and Cronquist (1963), Steyermark (1963), Radford et al. (1968), King and Robinson [1970; as Eupatoriadelphus fistulosus (Barratt) King & H. Robins.}, Correll and Johnston (1970), Strausbaugh and Core (1978), Cronquist (1980), Wunderlin (1982; in Expatoriadelphus), and Kartesz (1990). However, Jones and Fuller (1955) attributed the name E. fistulosum to Barrett ex Wood, and Soil Conservation Service (1982) attributed the name to Barratt ex Hooker (in Expatoriadelphus). We could not locate any Hooker publica- tion of this name. Since Barratt published this name in 1841, Hooker's usage of this name, if any, cannot be prior to 1841 (hence not in his Fl. Bor. Amer., vol. 2. 1840). Weatherby (1921) mentioned that Hooker, for his Fl. Bor. Amer. publication, invited Barratt to contribute the treatment of 'Please send reprint requests to Fryxell. Sipa 14(1):129. 1990. 130 EUPATORIA VERTICILLATA. ae 2°S6 SPECIMENS TO ILLUSTRATE THE NORTH AMERICAN VERTICILLATE SPECIES AND VARIETIES OF THE GENUS EUPATORIUM : With Synonymes and References, By JOSEPH BARRATT, M. D. MIDDLETOWN, CONNECTICUT May, 1841. ey A ra A tk { for distribution among Rot 1 fac. d 1 P 1 UPATORIUM FISTUL = Ep um. Evratoriuw purrurgum Linn. sp. pl. 1173, et Herb. ejuat; | (spe bad, as syn.;) not E. purpureum of ore a acre sp. ‘a 3, a bi tony eal vt et ‘Syn pa Cn .) Pur. gelow, fl. 2d Ed, p. | ei ig e 151, n. ae i = E. macula- Linn. sp. pl. 1174, (in prt) et Fa fide Dr. Gray cE. eum ¥ angustifoli r. & Gray Flor. 2, p. 82. Canad. p. 191. les rubesc (pri pot “Cau en colore suffusi rotundi inanes intus.” Coeaiile (L%.) he specific character given by Willdenow of E purpureum, applies to our E. Gaiulosum, The fal description ssere ta taken f Gone 8p. wl n (Sins E. verticillatum W. ! ae that Will- Semes.bes her can! Tiean aud other desesepancien detected several years a, 0, odie a tos medy wer ants inthe growing state; nor was I able lo settle the synony anaete UIT had made a full collection of specimens. hollow from the cor to ic; hei! Dor 10 feet; smooth bluish purple wud leueai reddish rp ani ntiahes ie neta purple. Tht is in one of the snost mailyi herbaceous plants in N. oe En ale nnd teu a sli tinct from the neat species, n i re instances | have found 2 ova. + joiuta whorls solids ‘The emall artic whieh | anon earaplanila retain the allow stem, and other ¢ leatneteriaties: The Mollovs ofthis |! Ee have been used extensively for blasting rocks atthe Middletown Felds shar “Quarry fistulosum does not pear to pussesa Uae acrid propertics ot F. incl The ori- | gina purpurcum, Lines ix sini (its 4) lo bea diferent plant, pre se i paval ating varictics, but all reins the characteristics of te species, ani iy gradations into other species. The nine here given by we, (since a new name was requited,) | ia sufficiently expressive to remove at anbiguity, The sabia in systematic works on this group of North Aimerican Faypatoriume, ix most extrourdinaty It is hoped that these speci- mens, with ha ies a references, will render the study of thia group easy and perfectly Should our European eorrespondents detect from our specitens | compare with the original Herbaria cited, any etrors, twill give ue plemure tore | ir communications and ¢ ns. ! 1. Evpatorium ristuLosum.* Barratt A small Heaney, 2. epeecanae macuLaTuM Linn. Aman. Ac. 4, p. hee sp. pl. 1174; ie sp. pl. 3 1760; Michaux. Flor. lakhs , p Decand. Prodr. 5, p. 152, n. 69. fect hi im solid striate hispid or pubescent, greenish and purple, with numerous ere and 1 parole lines. The glands on ey stew i wering Linn. 2p. 99; 15; 2,7. There are several varictica in the number of its 2. EvpaToRiumM MACULATU With 4 leaves in a whorl. . Evpatorium macucatum. L. With 3 pee in a whorl. \ 2. EvpaToRiuM MACUL Var. B aciaae Barratt. ht 4-5 fect, stem solid, slender; the leaves arc thin and much longer than the usual form ctr acl Hvis found in the Middletown menor near the ‘banks of the ‘Con- necticut ; ¢ perbaps indebted for many of our Eapatoriums, and other interesting ee this river we ar! c. char Willdenow or Pur == E verticillatum, Willd. sp. pl 3 (et Herb. fol. La) fe Gray Bigel ene 2 2d ed. p. rey Syn. Michx. 2, Stem solid, 6 feet high, glabrous green, sometimes purplish, with a purple ban the joints about 1 inch broad— polaaee in whorla of 3, 4 and 5—emontl ares ee a pabce- cece beneath and along midrib and nerves; length from 9-12 inches, e-ahaped.es: the base, including a ae of 1 or two inches, 4 tnelits wide, wh ade large arte, “The, leaves are thin, oh of greater size than any species pator ge & ine.” Cor tymb lax, flowe y pale purple p aeying | whitish. ie Has. oe The Tincean same be met bets, 2 ne ieee peclea, 02d Maer rear Baie G + bee mixed ap acters of two Ait, 3. EupaTorivum PURPUREUM. Var. 8 album.* alas \ i horl, larg rows in ORY Wood, aad tay ee fa toe ae Stem solid. : 3. EvpaToRiom PURPUREUM. Var. B album® Barratt. With 4 leaves in a whorl—a tall ies upper saa sub-falcate—Suwers dul white, an E. falcatum Mich? Grown in dry woods, Stem 3. EuPaTORIUM PURPUREU Var. 8 albu With 3 lcaves in a whorl; stem ne al a upper leaves gub- » note! sub E. eae ie Ara Elliott Flor, 2, p. 307 = E, Darlington Flor. Castrica 2d 4 lines trifoliatum. 453! Stem solid, variable in size ; leaves with a soft pubescence beneath, deeply serrate, hewers white or sometimes faint purple. Au attention w the number ot ne ves of the Vertiellate Panetoet rather than other characters, has led the older Lotaniste into frequent errovs, and bas duuch perplexity and confusion in systematic works, which requires to be UM TERNIFOLIUM. oo Flor. 2, p, 306! (in part.) 67. pan 5, p: 181, Stem solid, somewhat hispid and glandular; greenish, with purple dota and lines, Height frou 3-3 feet, a rather slender species; the upper and last whorls of leaves smal! and finely serrate. There is a variety ul B lernifolium, with smooth purple stems, which ia sometimes irregularly branched upwards, It has meatly 4 leaves in a whor : ——- —— =r _ ou nedaree —_eorr oe 4. aca TERNIFOLIUM rt, B. vesi eae Height 2-3 fer solid, striate purplish; in m undant—Stada ct est par t of Milton, caine mndeear AnH le arse Papert - purple ee wit the finest pene) The E. bagoraatict | Pr. by p.152, (exe 5 _ y perhaps prove a em seal va Jock pleat? are ® strongly reticu slated, “upper surface presents a vi pearance, i /e able degree, but which ia Eber effaced in the hard a preeee’ aed arcimens, a a large branched an ie ymb, (in “tig bare FIG. 1. Barract’s 1841 publication, citled Expatoria verticillata. Sipa 14(1):130. 1990. ea the genus Sa/ix. Although Barratt declined this offer, he did provide Hooker a synopsis of this genus. Based on Barratt’s synopsis, Hooker described two sections and few taxa in Sa/ix, and attributed the new names to Barratt, but the correct authority for these Sa/ix taxa is Barratt ex Hook (cf. Weatherby; Argus 1986). Probably this situation might have led a few taxonomists to believe that the correct authority for Expatorium fistulosum is Barratt ex Hook. We speculate that taxonomists who attributed the name Eupatorium fistulosum either to Barratt ex Hook. or to Barratt ex Wood, might not have seen Barratt’s single folio page publication or might have considered that publication as ineffective. From Barratt’s single folio page publication, it is evident that he validly described a new species, and from Wood (1847), Jackson (1881), and B. Robinson’s (193 1) references, we believe that Barratt’s name was effectively published. Hence, the correct authority for Eapatorium fistulosum is Barratt. The authors thank Hunt Botanical Institute Library, Barney Lipscomb (Editor, Sida), Hollis G. Bedell (The Botany Libraries, Harvard Univer- sity) for providing relevant literature for this study; and Larry E. Brown (Houston Community College), William R. Anderson (University of Michigan Hebarium) and Martin Cheek (Royal Botanic Gardens, Kew) for helpful suggestions. — Kancheepuram N. Gandhi, Dept. of Biology, Univer- sity of North Carolina, Chapel Hill, NC 27599-3280 and Paul A. Fryxell, U. S. D. A. Research Botanist in colloboration with Texas AGM University, College Station, TX 77843 REFERENCES ARGUS, G. W. 1986. oe — Salix (Salicaceae) in the southeastern United States. Syst. Bot. Monogr. 9:1 — BARRATT, J. 1841. me vertictilata, specimens to illustrate the North American verticillate species and varities of the genus Expatorium: with synonyms and references. | folio ; CORRELL, D. S. and M. C. nea 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renne CRONQUIST, A. 1980. Vascular ae es rhe southeastern United States - Asteraceae. Univ. North Carolina Press, Chapel H DEAM, C. C. 1940. Flora of Indiana. oe of Conservation, Divn. of Forestry, In- dianapolis. cae M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Company, New ork. ree H. A. and A. CRONQUIST. 1963. Manual of vascular Plants of northeastern United States and adjacent Canada. D. Van Nostrand Company Inc., Princeton, N. J. GREUTER, W., Chairman, editorial committee. 1988. International colle of Seance nomenclature, adopted by the Fourteenth International Botanical Congress, Berlin, Germany, Jul-Aug 1987. Reg. Veg. 118:1— 328. Sipa 14(1):131. 1990. [32 JACKSON, B. D. 1881. Guide to the literature of botany. Longmans, Green & Co. London. JONES, G. N. and G. D. FULLER. 1955. Vascular plants of IHinois. The University of Illinois Press, Urbana BAR TESZ, J. 1. 1990: A Page checklist of the vascular flora of the United States, Canada, & Greenland, 2nd ed. SS. KING, R. M. and H. oo io Studies pee (Compositae). XXI. A new genus, Neomirandea. Phytologia 19:305— 31 970. Studies in ee ee XXV. A new genus E i ela eae ple 19:431—4 ON O. ELA. 1971. Plots of tropical Florida. University of Miami ae oe oe MACKENZIE, K. K. ou. Scientific names applicable to our purple-flowered eupato- riums. Rhodora 22:157— 165. DFO E. AHLES, and C. R. BELL. 1968. Manual of the vascular flora of Casslinas. University of North Carolina, Chapel Hill. ROBINSON, B. L. 1904. Diagnoses and synonyms of American aaarerone Proc. Boston Soc. Nat. Hist. ecm — some Mexican and Central :247 — 254. \O Ww . Records ages toa perieral treatment of the Eupatorieae. Contr. Gray Herb. ee v 96327. oe C se aes ION SERVIC E. 1982. National list of scientific plant names, vols. S, Govt. Printing Office, Washington DC. ST EYERMARK J. A. 1963. Flora of Missouri. The ae State University Press, Ames, ST acount: PD. and E. L. CORE. 1978. Flora of West Virginia, 2nd ed. Seneca Books, Inc., Grantsville, WY. WEATHERBY, C., ie a I a 23: 71 WOOD 1847. A - book of botany, ed. 2. Crocker & Brewster Publishers, Boston. es ,K. M. and C. A. WEATHERBY. 1937. The nomenclature of the verticillate eupatoria. Ried 39: ae = 906: WUNDERLIN, R. P. 1982. Guide to the vascular plants of central Florida. University Presses of Florida, Gainesville. Old time Connecticut botanists and their herbaria-Il. CORRECT NAMES FOR THE VARIETIES OF CAREX ALBICANS / C. EMMONSII — Rettig (Sida 13:449—452. 1989) reduced Carex artitecta Mackenzie and C.physorhyncha Steudel to varieties of C. emmonsii Dewey ex Torrey. In doing so, I stated that the oldest specific epithet, C. ee Willd. ex. Sprengel, should not be used under article 69.1 of the N (Greuter et al., eds. International Code of Botanical Nomenclature. rere It has been pointed out, however, that the name may not have been used “widely and persistently” enough to warrant rejection. Regardless, unless a formal proposal to reject the name C. albicans is approved by the Committee for Spermatophyta and the Botanical Congress, the correct names for the three varieties are: Sipa 14(1):132. 1990. 2 CAREX ALBICANS Willd. ex Sprengel var. a/bicans, Syst. veg. 3:818. 1826. Type: CAROLINA (HoLoryee: B, GA {photo!]). Includes C. artitecta Mackenzie (C. emmonsit var. muhblenbergii (A. Gray) J. Rettig). CAREX ALBICANS var. EMMONSII (Dewey ex Torrey) J. Rettig, comb. nov. BasionyM: C. emmonsit Dewey ex Torrey, Ann. Lyceum Nat. Hist. New York -411. 1836. Type: Williamstown, Massachusetts, 1828 (NEoTyPE: NY!). Carex novae-angliae Schwein. var. emmonsii (Dewey ex Torrey) Carey in A. Gray, 1848. Carex varia var. minor at in Hooker, Fl. Bor.-Amer. I1:223, 1839, has neither a description nor a reference to a latent published description and is thus not validly published. The specimen taken as the type for C. emmonsii (Rettig 1989) postdates the name it typifies (C. dala eae by two years an should be called a neotype. CAREX ALBICANS vat. AUSTRALIS (L. Bailey) J. Rettig, comb. nov. BAsIONYM: C. varia vat. australis L. Bailey, Bot. Gaz. (C le)= Lie] 5a 1892. (Lecrorype: Tracy s.n. in 1889, Mississippi, BH!; tsocecrorypr: NY!, US!). Includes C. physorhyncha Steudel (C. emmonsti var. australis (L. Bailey) J. Rettig). For complete synonymy see Rettig (1989). I thank Dr. A. A. Reznicek for pointing out the problem and reviewing this manuscript. —_J. H. Rettig, Department of Biology, Texas A & M University, College Station, TX 77843, A KOELERIA GERARDII (VILL.) SHINNERS (POACEAE) NEW TO LOUISIANA — Koelerta gerardii (Vill.) Shinners was reported to be adventive at a few coastal localities in the United States including Texas by Gould, 1975. Hitchcock, 1951 reported this taxon as Koeleria phleoides (Vill.) Pers. and stated that it was introduced from Europe at Pensacola, Florida., Mobile, Alabama, Cameron County, Texas, Portland, Oregon, and at several points in California. Additionally, he reported it to be culti- vated in nursery plots at Beltsville, Maryland and Tucson, Arizona. A re- cent collection of this taxon from Vernon Parish, Louisiana is apparently the first for the state (Allen, 1980). Koeleria gerardii is usually called annual koeleria and is characterized by its annual habit and pubescent spikelets. The other species of Koeleria (K. pyramidata (Lam.) Beauv.) in Louisiana is perennial and has scabrous spikelets. The collection data are: LOUISIANA. Vernon Parish: disturbed area near Range Control off Texas Ave. on Ft Polk ca 4 mi ENE of Pickering, 12 May 1989, Allen 16433 (LAELSU,NLU). Sipa_ 14(1):133. 1990. 134 — Charles M. Allen and Harland D. Guillory, Division of Sciences, Loutsiana State University at Eunice, Euntce, LA 70535, U.S.A.; Charles H. Stagg and Stephen D. Parris, Environmental Section, Directorate of Engineering and Hous- ing, Fort Polk, LA 71459, U.S.A. REFERENCES ALLEN, C. M. 1980. Grasses of Louisiana. The University of Southwestern Louisiana, Lafayette. 358 pp. GOULD, E W. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station. 653 HITCHCOCK, A. S. 1951. Manual . an grasses of the United States. 2nd ed. revised by . Chase. U.S.D.A. Misc. Publ. RHYNCHOSPORA CAPILLACEA (CYPERACEAE), NEW TO TEXAS — Rhynchospora capillacea Torr., a cespitose perennial; culms deli- cately slender somewhat capillary to 4 dm tall; leaves filiform 0.2—0.4 mm wide, often as long as the inflorescence. Achene 1.7 — 2.6 mm long, long-elliptic, faintly marked horizontally rugose. Perianth bristles usually 6, retrorsely barbed as long as or surpassing the tubercle; tubercle subulate, about as long as the achene body. This species is usually found in calcareous seepage areas, bogs, swamps, shores and ledges ranging from Newfoundland to Saskatchewan, south to New Jersey, Pennsylvania, Virginia, Tennessee, Ohio, Indiana, Illinois, South Dakota and Missouri (Steyermark 1963, Godfrey & Wooten 1979). Waterfall (1966) listed R. capillacea as occurring in Bryan County, Oklahoma. Correll & Correll (1972) also listed R. capillacea as occurring in Oklahoma referencing Waterfall. Neither Correll & Johnston (1970), Stanford (1976), or Johnston (1989) listed R. capillacea as occurring in Texas. A collection of this distinct species, Kerr Co.: 3 Jun 1989, S. & G. Jones 2456 (HPC,SMU,SWT,TAES, TEX), is apparently the first report for Texas. The specimen was collected in an open calcareous hillside seepage at its in- terface with the south fork of the Guadalupe River with a SE aspect. It was locally frequent restricted to or near the seepage area. This site is situated 19 km (11.8 mi) NE on Highway 39 from its junction with Ranch road 187 at its crossing with the south fork of the Guadalupe River. The seepage is at the NE corner of said intersection. Elevation of the collection site ts 575 — 578 m with the geology being Edwards limestone of the Fort Terrett Members (Kft) (Lower Cretaceous). Associated species included Fuzrena Sipa 14(1):134. 1990. 135 simplex, Rhynchospora colorata, Cladium jamaicense, Eleocharis sp., Agrostis semiverticillata, Adiantum capillus-veneris, Justicia americana and Utricularia gibba. This collection site appears to be the southwesternmost United States record. A search in SMU, TAES, and TEX herbaria did not yield any additional specimens. Further investigations in suitable habitats and herbaria research might _ yiel additional Texas distribution records. — Stanley D. Jones, S.M. Tracy Herbarium, Department of Range Sci- ence, Texas AGM University, College Station, TX 77843 and Gretchen D. Jones, Department of Biology, Texas AGM University, College Station, TX 77843, A ACKNOWLEDGEMENTS We thank Andrea McFadden, Executive Director of the Botanical Research Institute of Texas (SMU) and Carole Todzia, Assistant Curator at TEX-LL for checking for specimens. REFERENCES CORRELL, D.S. and H.B. CORRELL. 1972. Aquatic and wetland plants of southwestern United States. Stock #5501-0177. Environmental Protection Agency Research and Monitoring, Washington, D.C. and M.C. eS aneene 1970. Manual of the vascular plants of Texas. Texas Research pare Renne GODEREY, R.K. and J. W. WOOTEN. 1979. Aquatic and wetland plants of southeastern United States. Univ. of Georgia Press, Athens, GA. JOHNSTON, M.C. 1988. The vascular plants f Texas. A list gs dating the manual of the vascular plants of Texas. Published by the author, Austi STANFORD, J.W. 1976. Keys to the vascular plants of cs Tena Edwards Plateau and adjacent areas. Howard Payne University, Brownwood, a ein J.A. 1963. Flora of Missouri. The Iowa ae University Press, Ames, ae tee, U.T. 1966. Keys to the flora of Oklahoma. 6th ed. Oklahoma State Univ. Press, Stillwater, OK. ADDITIONS TO THE FLORA OF TEXAS FROM EL PASO COUNTY. — An intense survey of the flora of the El Paso area of Texas over the past decade has yielded a number of additions to the flora of the state, most of which have now been recorded by Johnston (1988, The vascular plants of Texas, a list, up-dating the manual of the vascular plants of Texas, privately published). The following records have not yet been published for Texas and are worthy of note. Sipa 14(1):135. 1990. 136 Lepipium Latirotium L. (BRASSICACEAE). El Paso Co.: along the Rio Grande at the NM 273 bridge, Worthington 3146, 19 Aug 1978 (UTEP); Country Club Rd. bridge, Worthington 14119, 20 May 1986 (SMU,UCR,UTEP); Borderland Rd. bridge, Worthington 3254, 27 Aug 1978 (UTEP); and at Canutillo, Worthington 3254, 27 Aug 1978 (UTEP). Rollins (1981, J. Arnold Arbor. 62:517—540) notes that the species is now established in widely separated localities in Canada, United States and Mexico. The species 1s also common in sandy areas along the Rio Grande in El Paso County. BACCHARIS SAROTHROIDES Gray (ASTERACEAE). El Paso Co.: lower W slope of the Franklin Mts., along Trans-Mountain Rd. 1.1 mi E of jet. with I-10, 4100 ft elev., 16 Oct 1988, Worthington 17615 (NY, SMU,SRSU,TEX,UTEP); 3 mi E of jct. with I-10, 4800 ft elev., 14 Jan 1978, Worthington s.n. (UTEP); Three Sister Hills (31°52'N- 106°33'30"W), 4100 ft elev., 23 Oct 1988, Worthington 17623 (NY,SMU,UTEP). This species has apparently been recently seeded into the area along roadways where isolated plants are infrequently encounte- red. The species ts also occasionally used as an ornamental in desert land- scaping in El Paso. FILAGO CALIFORNICA Nutt. (ASTERACEAE). El Paso Co.: Franklin Mts., 1.7 mi NW ject. Trans-Mountain Rd. with Gateway South (31°54'50"N-106°27'42"W), 4400 ft elev, 8 Apr 1978, Worthington 4304 (TEX,UTEP); 27 Mar 1983, Worthington 9622 (NY,SMU,UCR,UTEP); 10 Apr 1988, Worthington 13585 (SMU,UTEP); | mi WNW ject. Trans- Mountain Rd. with Gateway South, 4400 ft elev., 16 Mar 1983, Worthin- gton 9463 (UTEP). The species occurs on rocky grante slopes among grasses and small shrubs. Mr. James D. Moorefield kindly brought to my attention an earlier record for “El Paso, Texas” (M.E. Jones 5.n., 22 Apr 1884, NMC,NY, POM). STYLOCLINE MICROPOIDES Gray (ASTERACEAE). El Paso Co.: NW EI Paso 0.3 mi N jct. Belvidere and Westwind (3 1°5 1’ 15”N-106°32'28"W), 4200 ft elev., crest of a low andesite rock hill, 29 Apr 1983, Worthington 10130 (TEX,UTEP). This species has long been suspected to be a part of the Texas flora as the type locality, “hills near Frontera, New Mexico,” might be on the Texas side of the Rio Grande. A smelting operation es- tablished in that area in 1887 has all but eliminated winter annuals from the Cerro de Cristo Rey of New Mexico and Chihuahua and the Campus Andesite hills in El Paso, Texas, making it impossible to duplicate the collection. The discovery of a small population on an isolated outcrop of Sipa 14(1):136. 1990. [af Andesite Rock in NW EI Paso confirms that this species is part of the Texas ora. I wish to express my thanks to James D. Morefield, Guy Nesom and Andrew C. Sanders for helpful comments and determinations. — Richard D. Worthington, Department of Biological Sciences, The University of Texas at El Paso, El Paso, TX 79968, U.S.A.. Sipa 14(1):137. 1990. REVIEWERS FOR VOLUME 13 _ The following individuals have kindly supported Szda through their me and efforts in reviewing manuscripts published in volume 13, 1989 — 1990. Without your interest and support Szd¢ would not be the journal that you all have come to expect. I hope each and every one of you that Sida has come in contact with have enjoyed it over the years. I trust that Sida will continue to improve and with continued support it can remain a top quality journal of systematic botany. Szda’s subscription base continues to expand each year with subscriptions approaching 700 in almost 70 countries. Thanks to all of the authors, reviewers, subscribers, and readers for your continued interest. Allen, Charles M. Haynes, Robert R. Sundberg, Scott Allred, Kelly W. Henrickso Taylor, Elizabeth L. Arp, Gerald K Higgins, lary Thomas, R. Dale Averett, John aa Tucker, Gary Bryson, Charles T. Campbell, Julian Carter, Richard Castaner, David Chamberlain, D.E Chmielewski, J.G. Clewell, Andre E Coile, Nancy C. Comeaux, Barry Ga Helen B. Dent ton, Melinda E Gunn, Charles R. Hall, David Hall, Gustav W. Harriman, Neil A. Hatch, Stephan L. Sipa 14(1):138. 1990. Kimnach, Myron Kron, Kathleen A. Johnson, Dale Johnston, Marshall Lane, Meredith A. Lonard, Bob Morin, Nanc Nauman, Cie E. Ne , Guy ee ae oe H. Powell, A. Michael Pringle, James S. Sie ie K. Reznicek, ee! ree aa Sheviak, anes J. Smith, Edwin Spongberg, ae A. Strother, John Tucker, Gordon C. ‘Turner, Urbatsch, Lowell E. Wells, James R Yatskievych, George ee) BOOK REVIEWS VERGUNOYV, A. P AND V. A. GOROKHOV. 1988. Russian Gardens and Parks. 418 pp. Moscow: Nauka. Text in Russian. The volume begins with an Introduction and is subsequently divided into two sections: 1) Developmental Stages of Garden and Park Art and 2) Creation of Garden and Park Compositions. There are 5 chapters in each of the sections. There are 83 colored photographs and numerous other black and white photographs as well as architectural illustrations. “This book deals with the specialised and little studied art of creating gardens and parks - artistic groups. On the basis of extensive factual material and the results of previous studies, the originality of this art’s historical develop- ment in Russia from the times of Rus, before Czar Peter the Great’s reign, to the turn of the 20th century has been shown for the first time. ...This book is intended for specialists: art experts, architects, artists, dendrolo- gists, park and museum personnel and everybody who is interested in land- scape architecture.” HIGNIGHT, K.W., J.K. WIPFE AND S.L. HATCH. 1988. Grasses (Poaceae) of the Texas Cross Timbers and Prairies. 174 pp. MP-1657. Texas Agricultural Experiment Station, The Texas A&M University System, College Station, TX. This publication contains several descriptive sections prior to the diagrammatic illustrations of the taxa: Introduction, The Grass Plant, Vegetative Parts, Inflorescence Types, and Spikelet Parts. There are 6 Figures of diagrammatic illustrations of the grass plant, inflorescence types, spikelet parts, and spikelets representing major genera and tribes. A checklist of the taxa precedes the Key to the Genera and the Key to Species. The “text” consists of diagrammatic illustrations of the species in alphabet- ical order (pp. 31— 166) followed by a glossary, references, and index. Sipa 14(1):139. 1990. 140 GOODRICH, S. and E. NEESE. 1986. Uinta Basin Flora. 320 pp. + xvii. USDA Forest Service-Intermountain Region, Ogden, Utah. Paperback. The flora contains about 1,600 specific and subspecific taxa of vascular plants. Uinta Basin is bounded on the north by the crest of the Uinta Mountains, on the west by the divide of the Strawberry drainage, and on the south by the breaks of the West and East Tavaputs Plateaus (parts of Colorado and Utah). Contents include preface, introduction, history of collections, the flora, plant communities, acknowledgements, glossary, key to families, descriptive flora, excluded names, addenda, literature cited, and index. MOERMAN, DANIEL E. 1986. Medicinal plants of Native America. Research reports in ethnobotany, contribution 2. Vol. 1, 2. 910 pp. The University of Michigan Museum of Anthropology Technical Reports, Number 19. University Museums building, Ann Arbor, Michigan 48109. A state of the art in al data management. The medicinal uses of plants by 122 tribes from 92 references are summarized in two volumes. The first volume catagorizes the tribal usage by plant taxon, genus and species and concludes with the bibliography. Volume 2 provides informa- tion of the medicinal uses of plants by Native Americans by the basic medi- cal usage, by plant family and by group or tribe. These volumes will certainly be of use to scholars in anthropology, botany, geography, Native American studies, medicine and the allied health delivery sciences. This ts the most important medical reference guide to Native American medical plants ever assembled. John E. Ubelaker. VUILLEUMIER, FRANCOIS and MAXIMINA MONASTERIO (Editors). 1986. High Altitude Tropical Biogeography. Oxford University Press (and American Museum of Natural History), 200 Madison Avenue, New York, NY 10016. Cloth $75.00. 649 pp. This volume is recommended for all those interested in endemic, rare and endangered species, either plant or animal. WFM Stipa 14(1):140. 1990. (Continued from front cover) si aol (Asteraceae: Astereae), a new genus related o Euthamia. Guy L. Nesom, ae Suh, David R. nee and Beryl B. Simpson 101 The herbaceous flora of three Weches formation outcrops in eastern Texas. Robert J]. George and Elray S. Nixon. Liy NOTES. A nomenclatural note on Expatorium fistulosum (Asteraceae). 129—Correct name the varieties of Carex albicans/C. emmonsti. Nees gerard aL ) Shinners (Poaceae) new uisiana 133 Ll. (Cyperaceae), new to Texas. 134—Additions to the flora of Texas feoci El Paso County. 135. Reviewers for volume 13 138. News, announcements, book reviews 78, 86, 100, 128, 139, 140. CONTRIBUTIONS S IDA To BoTANYy VOLUME 14 NUMBER 2 DECEMBER 1990 CONTENTS A new species of Acourtia (Asteraceae-Mutisieae) from southern México. Leticia Cabrera R. 141 Taxonomy of Digitaria section Aequiglumae (Poaceae: Paniceae). Robert D. Webster and Stephan L. Hatch. 145 The Plumbaginaceae in the flora of the southeastern United States. James L. Luteyn. 169 Taxonomic notes on western American Gentianaceae. James S. Pringle. LD A new species of Lzatris (Asteraceae) from the Carolina sandhills. Jon M. Stucky and Milo Pyne. 189 Lectotypification of Lactniarta carinata (Asteraceae). Milo Pyne and Jon M. Stucky. 209 Leaf venation studies in Indian Szda (Malvaceae). A.M. Saibaba and Raja Shanmukha Rao. 215 A new species of [bervillea (Cucurbitaceae) from western México. Rafael Lira 8. and Denis M. Kearns. 223 Gentianella canosoi (Gentianaceae), a new species from Durango, México. Guy L. Nesom and B.L. Turner. Zo} A revision of Paxistima (Celastraceae). Ann M. Navaro and Will H. Blackwell. 231 (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 2, pages 141— 313. Copyright 1990 by Wm. E Mahler A NEW SPECIES OF ACOURTIA (ASTERACEAE-MUTISIEAE) FROM SOUTHERN MEXICO LETICIA CABRERA R. Department of Botany, University ee Texas at Austin Austin, TX 78713 ABSTRACT Acourtia ovatifolia, a new species from Guerrero and Oaxaca, México, 1s described and illustrated. This species is similar in some characters to A. fozantii. RESUMEN Se describe e¢ ilustra una nueva especie, Acourtia ovatifolia, colectada en Guerrero y Oaxaca, México. Esta especie es similar a A. /ozant en algunos caracteres AcOURTIA OVATIFOLIA L. Cabrera, sp. nov. (Fig. 1) Acourtia lozanit (Greenm.) Reveal & King similis indumento glanduloso, foliis ovatis, et phyllariis apicibus adaxialibus reflexis glandulosis sed differt foliorum ad bases rotundatis, et capitulis flosculis paucioribus. Perennial plant up to 1 m tall; stems several from the base, green to dark-purple, striate, densely stipitate-glandular and with scattered crispy- articulated hairs, with a tuft of wooly brown hairs in the leaf axils. Leaves, liptic, 3—8 cm long, — petiolate, with the main blades ovate to ovate-e 1.5 —4 cm wide, progressively decreasing in size towards the infloresc- ence, basally rounded, acute and short-apiculate at the apex, semicoriac- eous, with a dense crispy-articulated indument with some of the hairs with small glandular tips, usually shiny and scabrid on the stipitate- to sessile- glandular upper surface, with both crispy-articulated and_ stipitate- glandular hairs on the veins, prominently veined on both surfaces; the margins entire to denticulate, sometimes slightly sinuate and subrevolute; petioles | — 3.5 mm long, with a dense indument of crispy-articulated and stipitate-glandular hairs. Inflorescence cymose; floral branch nodes with prominent tufts of brownish hairs; heads in clusters of several at the end of the branches or in loosely compound cymes, sessile to shortly pedunculate; peduncles up to 5 mm long, stipitate glandular, with a few leafy, ovate to elliptic and stipitate-glandular scales grading into the phyllaries; involucre cylindric to cylindric-campanulate, 1.5 —2 cm tall; phyllaries in 4—5 series, dorsally stipitate-glandular, with the outer ovate, acute at the Stipa 14(2):141 — 144. 1990. 142 reflexed, leafy-green tip, with the exposed adaxial surface stipitate- glandular, marginally ciliolate and the innermost oblong-lanceolate, gradually apiculate, adaxially glabrous; receptacle scrobiculate, glabrous. Flowers 9 per head; corollas pale pink, bilabiate, 11— 12 mm long, in cluding the 4 —5 mm long outer tridentate lobe, with the inner two ee slightly shorter; anthers 7 — 7.5 mm long with sterile pink tips; style and branches orange, 11 — 11.5 mm long, including the 0.8 —0.9 mm long, papillose, truncate branches. Achenes linear-fusiform, 4—4.5 mm long, stipitate-glandular and hispidulous; pappus 9 — 10 mm long with white bristles in 3 series. ow Type: MEXICO. Guerrero: limestone hill 9 mi by road N of I[guala, in shrubby oak- woods, 1450-1790 m, 7 Feb 1970, W. R. Anderson & C. Anderson 5656 (HOLOTYPE: MICH!). Additional collections ponecne MEXICO. Oaxaca: Road Nacaltepec-Jayacatitlan, 7.8 km SW of Hwy 135, ons slog eyed forest with Ipomoea, Brahea, Lantana hirta, 1600 m, 20 Oct 1989, iia oe 780 (TEX). As a part of a monographic at of Acourtia (in prep.), a study of herbarium specimens revealed this previously undescribed species, collec- ted in 1970 in the state of Guerrero by W. R. Anderson and C. Anderson. Attempts to locate additional material from the same locality were un- successful. Nevertheless, in a trip during October of 1989 to the state of Oaxaca, the same species was found coexisting with another Acomrtia species, the scapiform A. scapiformis (Bacigalupi) B. Turner. Only two individuals of Acourtia ovatifolia were found in the Oaxaca locality, growing in shady and steep places. Both plants had only a few buds, thus the flowering period may start during the winter, as is Common- ly the case for many other species within the genus. The type specimen was collected during the month of February and was in full bloom. Acourtia ovatifolia is similar to A. /ozanii in its phyllaries with reflexed and glandular tips, an unusual character within the genus. Both species have a glandular indumentum and ovate leaves, but in A. ovatifolia the leaves are rounded at the base, and in A. /ozanii the bases are cordate to auriculate. Also, compared to A. ovatifolia, A. lozanii has a campanulate to hemispherical involucre and a greater number of florets per head (25 — 39). The rounded bases of the leaves, as well as the prominent tufts of hairs on the nodes of the floral branches, easily distinguish A. ovatifolia from all other Acourtia. In Acourtia ovatifolia both types of glandular hairs, stipitate and sessile, are of a resinous nature. This characteristic seems to be widespread within the genus. Bacigalupi (1931) recognized 44 species of Acourtia. With the species 143 VX \ X gaps y a5 Poe wth Cok Cane Ys 4 Mey thas Bs yt? , 8 a ie a ine OSS Sy? NAN. IG. 1. Acourtia ovatifolia. a) habit; b) capitulum; c) adaxial view of the involucral brace sat d) detail of the leaf-stem junction 8 the indument. Illustration based on the specimen W. R. Anderson & C. Anderson 5656 (MIC — described here, and others recently or in the process of being described, the number of species within this genus is ca. 60 ACKNOWLEDGEMENTS I thank Guy Nesom for the Latin translation and his critical review of the manuscript. | am grateful to Beryl B. Simpson, Gregg Dieringer and an 144 anonymous reviewer for their constructive comments, and Billie L. Turner for his support. MICH herbarium provided the specimen loan. The illust- ration was drawn by Nancy Webber. The field trip to Oaxaca was suppor- ted, in part, by a B. L. Turner Fellowship granted by the Department of Botany of the University of Texas at Austin. REFERENCES BACIGALUPI, R. 1931. A monograph of the genus Perezia, section Acourtia with a provi- stone s aa to the section Experezia. Contr. Gray Herb. 97:1—81. TURN B. L. 1978. Taxonomic me of the pict species of Acourtia (Asteraceae- ee. Phytologia 38:456 TAXONOMY OF DIGITARIA SECTION AEQUIGLUMAE (POACEAE: PANICEAE) ROBERT D. WEBSTER United States Department of Agriculture Agriculture Research Service, Systematic Botany & Mycology Laboratory, Bldg. 265, BARC-East Beltsville, MD 20705, U.S.A. STEPHAN L. HATCH S. M. Tracy Herbarium Department of Range Science Texas A & M University College Station, TX 77843, U.S.A. ABSTRACT Twelve species are recognized in a taxonomic revision of Digitaria section Aequiglumae. The DELTA computer system was used in the collection, analysis, and presentation of taxonomic data. A comprehensive set of data was gathered from herbarium specimens and analyzed in order to produce a description of the section, key to the species, and comparat- IVE Species ¢ jos escriptions. Given with each species is a complete set of specimen citations and discussion of diagnostic characters and taxonomic relationships. Recognized species in- clude D. aequiglumis, D. connivens, D. costarwensis, D. cayabensts, D. ekmanii, D. ertostachya, D. lanuginosa, D. leucites, D. panciflora, D. sabulicola, D. simpsontt, and D. texand. INTRODUCTION Digitaria Haller consists of approximately 240 species, occurs in temperate and tropical regions of the world, and is commonly recognized as one of the most taxonomically difficult genera of the Paniceae R. Br. This difficulty is a result of the relatively large number of taxa, wide geographical distribution, wide range and complexity of the significant taxonomic characters, and a general lack of knowledge concerning the morphological relationship among the taxa. Digitaria is usually easy to separate from other genera of the Paniceae; however, rarely specimens may be encountered which can be confused with Panicum L. To distinguish these genera one has to ultimately determine whether the margins of the upper lemma are inrolled or flat, and this can sometimes be a difficult or subjective decision. Diagnostic characters of Digitaria include the follow- ing: ligule a membrane or ciliate membrane, primary inflorescence branch with secund spikelets, branches of inflorescence terminating ina spikelet, Sioa 14(2):145 — 167. 1990. 146 spikelets abaxial, and lemma of upper floret with flat, thin-textured margins (Webster 1988, Webster and Valdes 1988). Possible close relatives include Panicum, Anthaenantia P. Beauv. ,and Homolepis Chase; however, detailed quantitative studies of the character distribution for all genera of the Paniceae are required before definitive statements are possible concerning generic relationships. Uncil that time, all statements concern- ing proposed relationships among these genera provide only a limited service to the biological community. Henrard’s (1950) monograph of Digitaria, though 40 years out-of-date, continues to serve as the starting reference for all serious studies in the genus. That study recognized approximately 325 species and grouped these into the following four subgenera; Ev-Digitaria Stapf, Leptoloma Chase, Sefariopsis Stapf, and Solitaria Hack. Approximately 306 of the species were placed in the 32 recognized sections of subgenus Digitaria (= Ex-Digitaria Stapf). The section Aeguiglumae Henrard (described on page 641 of his monograph) was defined primarily on the basis of the second glume being equal in length to the lower lemma and upper floret. Other significant characteristics include paired spikelets and the typical absence of the first glume. Agrasar (1974) and Webster (1983) provide more detailed and comparative descriptive data for this section. Henrard’s monograph recognized the following taxa, all native to the New World, as comprising the Aeguiglumae: D. aequiglumis (Hack. & Arech.) L. Parodi, D. albicoma Swallen, D. campestvis Henrard, D. connivens (Trin.) Henrard, D. cuyabensis (Trin.) L. Parodi, D. distans (Chase) Fern., D. ekmanii Hitche., D. ertostachya Mez, D. laetevirens Mez, D. lanuginosa (Nees) Hen- rard, D. feacites (Trin.) Henrard, D. malacophylla (Hitchc.) Henrard, D. pauciflora Hitche., D. runyonii Hitche., D. sabulicola Henrard, D. Simpsont (Vasey) Fern., D. subcalva Hitchc., and D. texana Hitchc. Since that time, one new species, D. costaricensis Pohl, has been named to this section. The formal taxonomic history of the species in this section began in the early 1800's with the work of Trinius, Nees, and Arechavaleta. These authors named five species and placed them in Panicum or Paspalum. The first North American taxon was recognized by Vasey in 1892, who des- cribed it under Panicum. In the early 1900's Chase named three taxa and placed them in Syatherisma Walter. During this period, Mez named two new species and was the first taxonomist to place a member of the section in Digitaria. This generic concept was followed by Parodi who described one hew species and made two new combinations in the section. Between 1913 and 1934, Hitchcock contributed nine new combinations or new species in the section. The first three of these, those prior to 1926, were described in 147 Syntherisma and the subsequent ones in Digitaria. From 1950 to 1950 Hen- rard provided ten names and formally recognized the section. As with most other sections of Digitaria there has been little significant research to clarify taxonomic relationships among taxa of the Aequiglumae. A cursory analysis of some taxa related to D. fewcites is given in Boonbun- darl (1985), but undoubtedly the most meaningful recent revision ts Agrasar’s (1974) treatment of the Argentine Digitarias. It is not surprising that frequently the best sources of new taxonomic data are regional floristic treatments, however, these have the obvious shortcoming of being limited in aspects of geography and data collected. Floristic treatments of grasses of direct significance to the present study include: Hitchcock's (1927, 1936, & 1951) treatments for Ecuador, Peru, Bolivia, the West Indies, and the United States; Smith, Wasshausen, and Klein’s (1982) treatment for Santa Catarina; Rosengurtt’s (1970) treatment for grasses of Uruguay; and Mc Vaugh’s (1983) treatment for Nueva Galicia. The objective of this research was to clarify the taxonomic relationships among taxa of the Aequiglumae. To accomplish this goal, available herbarium specimens were studied and used to collect a comprehensive set of morphological and geographical data. The specific characters recorded for the recognized taxa are listed on microfiche in Webster et al. (1989). In addition to these characters, an anatomical examination of the leat blade epidermis for all recognized taxa was made. Data for the following characters were recorded: (1) average number of rows of stomates in the intercostal zones, (2) average stomate length, (3) average stomate width, (4) average number of rows of long cells in the intercostal zones, (5) average long cell width, (6) average long cell length, (7) average width of the in- tercostal zones, (8) relative abundance of prickles, and (9) the shape of the apex of the distal cell of the bicellar microhairs. Selection of these characters which we feel to be important was based on previous anatomical studies of species of Digitaria by Webster (1983). Results trom these anat- omical studies on taxa of the Aegwig/wmae indicated that the average values for certain characters could be used to separate some taxa. However, the range of variation within each taxon was great enough to prevent these characters from being useful or reliable for identification, Therefore, the anatomical data is not presented here. An interesting unifying anatomical feature of the members of this section is the presence of a bulbous swelling at the apex of the microhair distal cell. All members possessed this unique character, although it was not well-developed in D. pauciflora. Presence ot this feature in all members of this section and its absence from other species of the genus support a monophyletic interpretation for this section. 148 The morphological and geographical data were stored in DELTA format (Dallwitz 1974 & 1980) and used in the production of a key and descrip- tions. Data for 285 characters (see the microfiche in Webster et al. 1989) were recorded for each of the recognized taxa. INTKEY (the information retrieval program in Dallwitz’s DELTA system) was used to combine the taxonomically significant characters for the section into one description. In the following description of the Aequiglumae, each character state is accom- panied by a number or fraction indicating the distribution of data for the species. For example, ’Plants annual (1) or perennial (11)”, indicates that one of the twelve recognized species is annual and eleven are perennial. The fraction 1/9 indictaes that the character was coded for 9 taxa and 1 possessed the character state. This is followed by a key produced via CONFOR and KEY, which represents an efficient use of characters and placement of taxa for the practical identification of the species. Our con- cepts of the reliability of the characters are incorporated in the key. Descriptions of the recognized species includes a subset of characters of diagnostic value for the section. General methods used with this technol- ogy are adequately described elsewhere, but we consider it important to make the following general statement. Application of DELTA computer technology serves as an efficient means of collecting, analyzing, present- ing, and storing taxonomic data. It also allows for quick experimentation with the production of descriptions, keys, and database interrogation for various taxonomic purposes. However, those parameters that ultimately determine the value or usefulness of taxonomic research (i.e., development of character concepts and taxon concepts and the detailed, precise, and comparative collection of data) are and must be the responsibility of the taxonomist and not the methods applied — whether computerized or not. TAXONOMIC TREATMENT Dicrraria section ArQuiGLuMAE Henrard, Monograph of the genus Digitaria 641. 1950. — Tyrer species: D. aequiglumis (Hackel & Arechav.) L. Parodi. Description: Plants annual (1) or perennial (11). Main axis present or absent (2). Primary branches with appressed secondary branches; not win- ged; with neatly arranged spikelets. Pedicels truncate (8) or cupuliform at the apex (5). Cleistogamous inflorescence present (2) or absent (10). Cleis- togamous spikelets similar to the chasmogamous spikelets (2/2). Spikelets paired; densely (3) or slightly overlapping (11); evenly distributed on the rachis or becoming more concentrated at the rachis apex (1). Firsc glume present (2) or absent (11). Rachilla not pronounced below the second glume. Second glume present; 0.9— 1.0 times spikelet length. Lower 149 floret lacking stamens. Lemma of lower floret with equal internerve spac- ing (7) or with the first internerve space wider than the second (8); the nerves pronounced but not swollen; lacking distinct transverse nerves; glabrous (7) or hairy (9). Lower lemma hairs not forming a distinct hori- zontal line (9/9); overtopping (1/9) or subequal to the upper floret (8/9); smooth and terete (1/9) or flattened and coiled (8/9); without apical mod- ifications (9/9). Palea of lower floret vestigial (3) or absent (10). Upper floret 0.7 — 1.05 times the length of the lower floret. Lemma of upper floret smooth; grey (8), or yellow (7), or purple (3). Remarks: This study recognized twelve taxa in Digitaria section Aequiglumae. All are native to the New World with one species, D. aequiglumis, introduced into the Old World. Significant diagnostic characteristics of this section include the following; most are perennial, the primary branches are not winged, the first glume is typically absent, length of the second glume equals spikelet length, the lower floret lacks a well-developed palea, and when hairs are present on the spikelet parts they are simple, smooth, and unmodified. Additional interesting characters of the section include the presence of cleistogamous inflorescences tn two species and the presence of both cupiliform and truncate pedicel apices. Typically, members of a section within Digitaria will be consistent for the latter character. Recognized taxa: D. aequiglumis (Hackel & Arechav.) L. Parodi, D. con- nivens (Trin.) Henr., D. costaricensis Pohl, D. cvyabensis (Trin.) L. Seas ekmani Hitche., D. ertostachya Mez, D. lanuginosa (Nees) Henr., D. loucites (Trin.) Henr., Dinan pauciflora Hitche., D. sabulicola Henr., D. simpsonit (Vasey) Fern., D. texana Hitche. Key To THE Species Or Diairaria Section AEQUIGULMAE 1(O). Cleistogamous inflorescence ios in the axil of the uppermos leaf 2 Cleistogamous pens BOSCH. tac on ese hee ee 3 2(1). Upper floret 0.7 — times ie length of the lower floret; plants annual; plants ne a MC ects hae neys tee eae ae ee D. aequiglhumis Upper floret 0.88— 1.0 times the length of the lower floret; plants perennial; plants rhizomatous... 2... eee D. lanuginosa 3(1). Spikelets oe Chad. 3 min ONG «22462454546 dbiap eres aaas 4 Spikelets 4. 3-— 3.0) Gite IONE 29 4h c Gb S44eks se eee eae 12 4(3). Spikelets 0. as SMM Wide erads doa ke BG Bede ee eed D. sabuliwcola Spikelets more than 0.5 mm wide .............0.000000005- 5 54). Sorelets 0,9.) 0.99 mm-wide «2.25 io2eess ies Bese esa 6 Spikelets: Sreater than: 0.99 maim wide: vis 2esan ve iene ene Rie 11 6(5). Second aie 7 —nerved; first glume frequently present........ 7 econd glume 5 — nerved; first glume absent ................. 8 7(6). Leat bates filiform, leaf blades about 2 mm wide and purple in color, CUMS WILY eset His eae eeeawac ee ete oa ed D. panciflora 150 Leaf plage linear; leaf blades typically 3—4 mm wide and green in CONG COs NOL Say 22s c1 ppt blesh uae ba oe ee nane se eseary D. simpsonit 8(6). Primary oes branches. 0.2 0.3 tai Wide s.ecdne <3 nyu 9 Primary inflorescence branches greater than 0.3 mm wide ..... 10 8). Leaf ae : labrous; leaf blades with the midrib not obviously dif- fitiated: South Wimerica o2 345 ue gh wer eou preennnecen es D. ertostachya Leaf blades ie leaf blades with the midrib obviously differentiated; Ces nk exe ee eae ere oe Ae oe oe ee ee as . eRmanit 10(8). Leaf blades with the midrib obviously differentiated; spikelets aad densely hairy, with the hairs iii purple early; peduncle usually mere than 10 om. lone: mainly Tease vy 4 aan seen Gwe eee eeG wes D. texana Leaf blades with the midrib not obviously differentiated; spikelets sparsely hairy, the hairs white and occasionally turning purple; ped- uncle usually less than 10 cm long; mainly South America... .... D. cuyabensis 11(5). Lemma of lower floret glabrous; second glume 7-nerved; spikelets ‘abn FI QEIGG <5 ¢ Sede tek ne ae en eo ei ee D. simpsoni Mex shad 4 aratiedoin ay ee sy te aan alee Rae ha ye ad ate epee Sake D. leucites 12(3). Mid- oe leaf blades not reflexed; spikelets hairy; leaves hairy; Meso- GA. Gs had ete ntas fe eer ok Oe eee Te tearing ae eta ae a gs D. costaricens1s Mid-culm leaf blades reflexed; spikelets glabrous; leaves glabrous; SOUCH AMEr CA esis 6 oad pe wd 2 al en nete Saco dadow ne ¥ bcd ude hae D. connivens DIGITaRIA AEQUIGLUMIS (Hack. & Arech.) L. Parodi, Revista Fac. Agron. Veterin., Buenos Aires 4:47. 1922. Syntherisma aequighumis (Hack. & Arech.) Hitche., Contr. U.S. Natl. Herb. 7:211. 1913. Panicum debile Desf. var. aequiglume (Hack. & Arech.) Hack., in Stuckert, Anales Mus. Nac. Hist. Nat. Buenos Aires 11:69. 1904. Panicum aequiglume Hack. & Arech., Gram. Urug. 93. 1894. — Tver: URUGUAY, Montevideo, 3 — 1885, Arechavaleta 220 (HOLOTYPE: W. ioTrypee: US!). Panicum ramosum Arech., Anales Mis, Nac. Montevideo 1:1 11. 1894. Digitaria campes- tris Henr., Blumea 1:97. 1934. — Type: URUGUAY, een (HOLOTYPE: W). Panicum oe Phil., oo. io Chile 93:712. 1896. — Type: CHILE, Curico, (HOLOTYPE: W, # 4 Digitaria chillanensis Phil. ex Henr. Monogr. Digitaria 29. 1950. — Type: CHILE, HOLOTYPE: ) oe laetevirens Mez, Bot. Jahrb. Syst. 56:8. 1921. Digstaria aequiglumis var. laete- urens (Mez) Henr., Monogr. Digitaria 370. 1950. — Tyre: Evidently destroyed. Description: Plants annual; scoloniferous; lacking rhizomes (rarely with poorly developed compacted rhizomes). Nodes glabrous (rarely sparsely pilose). Auricles 1-2 mm long. Sheaths glabrous or hairy. Ligule 1.2 —2.2 mm 14 long. Leaf blades flexuous; spreading; mostly 2— 12 cm long; 2—6 mm wide; usually glabrous on the lower surface; glabrous or ISI hairy on the upper surface; with the midrib not obviously differentiated. Main axis 5 — 30 mm long; with quaquaversal primary branches. Primary branches appressed to to spreading from the main axis; whorled only at the lower nodes; (2 — )3 — 6(-8) on the main axis; 0.3 — 0.5 mm wide. Pedicels 2—3.5 mm long. Cleistogamous inflorescence present. Spikelets 26 — 42 on a typical primary branch; lanceolate; (3—)3.2—4.2 mm _ long; 0.6—0.8 mm wide. First glume absent. Second glume | times spikelet length; 5 or 7-nerved; glabrous or hairy; acuminate. Lemma of lower floret 7-nerved; acuminate; glabrous or hairy. Lower lemma hairs shorter than the upper floret; white. Upper floret 0.7 — 0.87 times the length of the lower floret. Lemma of upper floret grey (yellow when immature); acum1- nate. Distribution: Brazil, Paraguay, Uruguay, Chile, and Argentina. Remarks: Digitaria aequiglumis is the only representative of this section known to occur in the Old World, in that it has been introduced 1n Europe and naturalized in a few localities in southern Australia. This species, as treated here, is relatively easily to recognize; however, previous authors have placed some specimens under D. /anuginosa and D, cuyabensis. Usually it can be recognized at a glance by the presence ofa relatively high degree of branching, dark-colored nodes, and yellow-green leaves. Taxonomically significant sptkelet characters include the relatively long acuminate second glume and lower lemma which overtop the upper floret by 0.5 — 1.0 mm. Presence of the cleistogamous inflorescence in the upper leaf sheath was positively correlated with the other primary characters used to define this species. As far as we are aware, this is the only species of this genus where this feature serves as a significant character for differentiating a species. This species lacks rhizomes and well-developed stolons and was therefore described as an annual, but it is believed that plants survive for more than one year. Henrard (1950) differentiated D. campestris from D. aequiglumis based primarily on variation in pubescence of the nodes and leaves. Our studies of specimens in this section have shown that such variation cannot be correla- ted with important diagnostic characters and since there were no other dif- ferentiating characters we are treating D. campestris asa synonym. Digitaria laetevirens Mez appears to be identical to D. aequiglumis except for the complete absence of hairs on the second glume and lower lemma. This pattern of pubescence, where there exists essentially identical glabrous and hairy forms, is not uncommon in Digitaria. Henrard frequently gave formal taxonomic rank, usually varietal, to this variation, however, recent revisions by Veldkamp (1973), Webster (1983), and Webster (1987) ten- ded to minimize the taxonomic importance of this character variation. [52 Representative specimens: ARGENTINA. (locality uncertain) T. Stackert 13862 (US #557989). Buenos Aires: Dock Sur, 8 Apr 1909, S.I. No. 12524 (US). Cordoba: Herbarium Hackel (US #297953); Bezirk Rio Primero, Villamonte, Jan 1904, Stuckert 545 (MO, US); Corrientes: Santa Fé, Villa Ocampo, 20 Jan 1895, C. Quarfn 1895 (US). Entre Rios: Parana: Chana Mini, L. Parodi 4926 (US); Punilla: Orilla NE del Lago San Roque, 29 Feb 1976, Hunztker 22925 (NY). Tucuman: Capital, Rio Sali, 16 Dec 1923, 8. Venturi 2167 (US); Cerro de Campo, 15 Mar 1930, 8S. Venturi 10212 (US). BRAZIL. Brasilia: RGS, Cristal, Porto eke 31 Mar 1949, B. Rambo J. 40741 (US). Curitiba, Parque Rio Iguacu, 27 Dec 1979, R. Kummrow 1297 (MO, NY). Rio Grande do Sul: Dom Pedrito, 15 Apr 1946, ane 9102 a ); Santa Victoria do Palmar, 27 Apr 1946, Swallen 9207 (US). ei carina: ie ec 1972, R. M. — 10.458 (US). CHILE. Palguin, Nov 1928, . Joseph 4840 (US); Santos de Chillan, (US #1126084). URUGUAY. Campos del sae i ity unknown) J. Aha sn. (US #927949, #927950, #927951, #927952). Hackel 30 (US #927958). rro Largo: Dec 1935, B. Rosengurtt 1049 (US). Durazno: coe ia Las Palmas, 1926, coe 18743 (US); San Gregorio, La Paloma, Osten 19536 (US). Canelones: Arroyo Sarandi sobre el rio de la Plata, Costa Azul, 26 Feb 1956 B. Rosengurtt B-6524% (BF). Flores: rio fi y Arroyo Marindro, 10 Apr 1937, B. Rosengurtt B-1508b (US). Florida: Campo experimenta de Pastos, Estancia Rincon de Santa Elena, 23 Feb 1948, B. Rosengurtt 5960 (E US); Mausavillagra, 31 Dec 1936, B. Rosengurtt B- 890 (US). Montevideo: | Apr 1888, Arechavaleta (US #927960). Rocha: Laguna Negra, 20 Mar 1938, B. Rosengurtt B-2615'% (US). Soriano: Monzén-Heber, Juan Jackson, | May 1940, B. Rosengurtt PE-4385 (E US, NY) DiGitARIA CONNIVENS (Trin.) Henr., Meded. nea Heths-O iG, oo connivens Trin., Mém. hon Imp. Sci. St. Petersbourg 6(3): 206. 34. — Tyee: BRAZIL (HoLotype: LE; isorypr: PW). eee Plants perennial; stoloniferous; lacking rhizomes. Nodes : labrous. Sheath auricles 0.5 — 1.5 mm long. Sheaths glabrous. Ligule 1.5 —3 mm long. Leaf blades straight to flexuous; reflexed; 2 —5 cm long; 1-6 mm wide; glabrous on the lower surface; glabrous on the upper surface; with the midrib not obviously differentiated. Main axis 3— 15 mm _ long; with quaquaversal primary branches. Primary branches appressed to the main axis; not whorled; 2—4 on the main axis; 0.4 — 0.5 mm wide. Pedicels 2—6 mm long. Cleistogamous inflorescence absent. Spikelets 16 — 34 ona typical primary branch; lanceolate eee el- liptic); 3.3 —5 mm long; 0.9— 1.2 mm wide. First glume mostly absent (occasionally present as a vestigial scale ca. 0.2 mm long). Second glume 1 times spikelet length; 7(-9)-nerved; glabrous; acuminate. Lemma of lower floret aa acuminate; glabrous. Upper floret 0.94— 1 times the length of the lower floret. Lemma of upper floret grey or yellow; acumt- nate. Distribution: Southeast Brazil. Remarks: Digitaria connivens occurs in southern coastal regions of Brazil, where it is commonly associated with coastal sand dunes. Important diagnostic characteristics possessed by this easily recognizable species in- 153 clude the relatively short, reflexed, and obviously distichous leaf blades; an inflorescence consisting of only a few primary branches appressed to the central axis; relatively large and completely glabrous spikelets; and a caryopsis which completely fills the interior of the upper floret. Representative specimens: BRAZIL. (location and collection date uncertain), Schott 1961, 4844 (US). Parana: Caioba, Praia do Mendanha, June-1 — R. Braga 1627 (US). Rio de Janeiro: Rio de Janeiro, ey dare unknown, ae s.n. (US). Rio Grande do Sul: Sacco du Mangeira, 1902, G. O. Mea/me 1561 (US); Brasilia, Torres, 11 Feb 1954, B. Rambo 8. J. 54777 (US); Rio en 3 May 1946, J. R. Swallen 9241 (US); Camaqua, Fazenda Aguada, Lagoa dos Patos, 12 May 1946, J. R. Swallen 9287 (US). Santa Catarina: Jureré, - aa 1966 Klein, Souza S. & Bresolin 6.057 (US), Voyage A. St. Hilaire 1711, 1816 — , (US). Sao Paulo: Mun. Cananeia, IIha Comprida, 16 Feb 1965, W D. Clayton a Biren 4688 (US); Mun. Cananeia, IIha Comprida, 16 Feb 1965, W. D. Clayton & G. Eiten 4710 (US); Mun. [guape, East point of Iguape Island, 19 Feb 1965, W. D. Clayton & G. Eten 4761 (US); Santos, Guaruja, 23 Feb 1938, F C. Hoehne 39261 (US). DIGITARIA COSTARICENSIS Pohl, Fieldtana, Bot. 38:5. 1976. — Tver: COSTA RICA, Prov. Cartago, 10—-8— 1986, Pohl & Davidse 11215 (HOLOTYPE: ISC!) Description: Plants perennial; stoloniferous; with poorly developed rhizomes. Nodes hairy. Sheath auricles about 1.5 mm long. Sheaths hairy. Ligule 1-2 mm long. Leaf blades flexuous; spreading; 4— 15 cm long; 3—7 mm wide; hairy on the lower surface; hairy on the upper surface; with the midrib not obviously differentiated. Main axis 10 — 20 mm long; with quaquaversal primary branches. Primary branches appressed to the main axis; not whorled; 3 —6 on the main axis; 0.3 —0.4 mm wide. Pedicels 2—4 mm long. Spikelets 18 — 50 ona typical primary branch; lanceolate; 3.3 —4.1 mm long; 0.85 — | mm wide. First glume absent (or present as a hyaline scale to 0.3 mm long). Second glume | times spikelet length; 7- nerved; hairy; acuminate to acute. Lemma of lower floret 7-nerved; acumi- nate to acute; hairy. Lower lemma hairs shorter than the upper floret; white. Upper floret 0.95 — | times the length of the lower floret. Lemma of upper floret yellow; acuminate to acute. Distribution. Costa Rica. Remarks: Pohl (1980) correctly placed this species in the Aeqguiglumae on the basis of the relative sizes of the spikelet parts. In addition, he indicated that it seemed cto be most similar to D. aequiglumae but differed on the leaf pubescence, leaf width, presence of a first glume, and the number of nerves on the second glume and lower lemma. Even though there are only three known collections, it is obvious that they represent a distinct taxon. The pattern and type of leaf pubescence is similar to the hairy form of D. cuyabensis. The spikelets are relatively large with pronounced nerves and scattered fine hairs on the second glume and lower lemma. 154 Representative specimens: COSTA RICA. Providence Cartago: Rio Macho Reservoir, $ of Orost, 3 Oct 1968, Pohl & Davidse 11190 (SC); 25 km SW of Tejar along the Carretera Interamericana, 8 Oct 1968, Pohl & Davidse 11215 USC); 2 km W of Paraiso, 21 Apr 1969, Pohl & Davidse 17789 USC). DiGiraRia CUYABENSIS (Trin.) L. Parodi, Physis 8:378. 1926. — ree fanuginosa (Nees) Henr. var. cvyabensis (Trin. ) Henr. Fes oe 164— 165. 19 Contr. U.S. Natl. Herb. 22: L922, Poe ea Trin., Mém. Acad. a Ga ot eet. 3:206. 1834. — Type: Trin nen (HOLOTYPE: LE) Syntherisma cuyabensis (Yrin.) Hitche. ,) Syntherisma malacophylla Hitche., Contr. = S. Natl. Herb. 22:466. 1922. Digitaria oe (Hitchce.) Henr., Meded. ks-Herb. 61:4. 1930. — Tyre: BRIT- NA, 31 Dec 1919, seas oe (HOLOTYPE: US!). Description: Plants perennial; stoloniferous (frequently not pronounced); rhizomatous. Nodes glabrous or hairy (the upper nodes usually glabrous). Sheath auricles 0.8 — 1.5 mm long. Sheaths glabrous or hairy. Ligule | — 2 mm long. Leaf blades straight; spreading; 2.5 — 20 cm long; 3 — 6(-8) mm wide; glabrous or hairy on the lower surface; glabrous or hairy on the upper surface; with the midrib not obviously differentiated. Main axis 5 — 40 mm long; with quaquaversal primary branches. Primary branches appressed to the main axis to spreading; usually whorled at the lowermost node; 3 — 8 on the main axis; 0.31 — 0.4 mm wide. Pedicels 1.5 — 2.5 mm long. Spikelets 30 — 60 on a typical primary branch; lanceolate to elliptic; 2.4—3 mm long (-3.5); 0.6-—0.75 mm wide. First glume absent (or present as a minute hyaline scale 0.1 —0.2 mm long). Second glume 1 times spikelet length; 3 to 5-nerved; hairy (rarely glabrous); acuminate to acute. Lemma of lower floret 7-nerved; acuminate to acute; glabrous or hairy. Lower lemma hairs shorter than the upper floret; white or purple. Upper floret 0.95 — 1.05 times the length of the lower floret. Lemma of upper floret grey; acuminate to acute. Distribution: South America (Guiana, Surinam or French Guiana, Brazil, Paraguay, Uruguay, and Argentina) and Central America. Remarks: Digitaria cuyabensts occurs in Argentina, Paraguay, Uruguay, Brazil, and extends up the east coast of South America to Central America. It has been frequently confused with D. aequiglumis; however, these species, as defined here, are clearly distinct. Digitaria cuyabensis is an obvious perennial, spikelets are usually less than 3.1 mm long and less than 0.8 mm wide, and the leaf blades and spikelet bracts are usually hairy. The complete absence of a cleistogamous inflorescence and relative length of the florets are the most important diagnostic characters for distinguish- ing between these species. Hitchcock (1922) originally described D. malacophylla in the genus Syn- 155 therisma and differentiated it from D. cvyabensis on the basis of leaf pubesc- ence and the spreading nature of the primary inflorescence branches. His concept of the species was based only on the type specimen. Henrard (1950) transferred it to Digitaria and separated it from D. lanuginosa, which included the variety cvyabensis, based on a shorter spikelet length and its proposed annual nature. Our studies indicate that typical specimens of D. cwyabensis vary in the amount of leaf hairs from glabrous to densely hairy. Generally, specimens from northern part of South America, specifically from the Brazilian state of Pernambuco, are hairy and those from southern parts tend to be glabrous; however, there are numerous obvious exceptions. The primary branches of the inflorescence are spread- ing at maturity in both forms. In addition, other specimens referable to D. malacophylla show its perennial nature and spikelet lengths intergrade setween the two forms. Therefore, it was concluded that D. malacophylla is best treated as a synonym of D. cuayabensis. Most of the specimens in the D. aequig/umis complex fit well within the classification system proposed in this paper; however, a few specimens Steinbach 0877 (US), Parodi 8323 (US), and Parodi 9259 (US) from Argen- tina; Gunes s.n. from Venezuela; Hitchcock 8245 from Panama} were morphologically intermediate. These specimens are perennials and lack a cleiscogamous inflorescence; the spikelets are about 2.8 mm long, acumi- nate, with an overtopping second glume and lower lemma. Therefore, they do not fit well into the concepts of D. aequiglumis, D. lanuginosa, D. cuyabensis. It was concluded that these specimens do not represent a new taxon but are the products of hybridization and introgression within this complex. They were annotated as intermediates. a specimens: ARGENTINA. Buenos Aires: Puerto Nuevo, 15 Apr 1928, rodi 8524. Corrientes: Parada Pucheta, Ruta Nac. No. 127, 17 Feb 1979, O. es 2551 (MO). Formosa: (location not given), Jan 1918, Jorgersen 2434 (US); (loca- tion not given), Jan 1928, L. R. Parodi 8338 (US); (location not given), 23 Jan 1928, L. R. Parodi 8326 (US). BRAZIL. Ceara: Campo Grande, 12 May 1934, J R. Swallen 4533 (US). Mato Grosso: Mun. de Caceres, Faz. Descalvados, 4 Nov 1978, A. Allem et al 2386 (MO): between Campo Grande and Dourados, 14 Feb 1930, Chase 10923 (US); between Campo Grande and Dourados, 14 Feb 1930, Chase 10923+ (US). Minas Geraes: Serra de San Antonio, Diamantina, 27 Dec 1929, Chase 10328 (US); Serra de San Antonio, Diamantina, 27 Dec 1929, Chase 10417 (US). Pernambuco: Recife, 12 Nov 1924, Chase 7670 (US): eee 20 Nov 1924, Chase 7763 (US); Tapéra, Feb 1929, B. Pickel 1968 (US); Tapéra, Jan 1930, B. Pickel 2241 (US); Tapéra, 4 ao er B. Pickel 3171 (US); Tapéra, 9 Jan 1935, B. ae 3769 (US), Tapéra, 30 May 1935, B. Pickel 3794 (US). Rio Grande do Norte: Estremoz to Natal, 1 Jun 1934, J. R. ae 4788 (US). GUYANA. North Guiana, Rockstone, 13 Jul 1921, H. A. Gleason 636 (US); Rockstone, 31 Dec oo. Hitchcock 17284 (US); (location unknown), 1838, Leprienr s.n. (US). PARAGUAY. Central: Asuncion, Banco San Miguel, Rio Paraguay, Jan 1949, Rosengurtt 5442 (US); rd 156 sé aga du Rio Paraguay, Feb 1877, B. Balansa 147 (US); Loma Para, (Chaco), 15 May , LT. Rojas 2068 (US); Pilcomayo River, May 1906, T: Rojas 92 (US); Pilcomayo River, ae — 1890, T. Morong 962 (US); Puerto Casado, Jan 1917, To Rojas 2723 Nes Puerto Colon, May 1943, 7) Rojas 10525 (US); R ie lacus Ypacaray, Jan 191: ead 12464 (US), reece (collection date unknown) E. Hassler 13018 (US). ae Chirinqui: between El Hato and Cerro Punta, Ps ee 960, E. Ebinger 765 (US); vicinity a El Boquete, 7 ae L911, Hitchcock 8262 (US). DictrariIA EKMANU Hitche., U.S.D.A. Misc. Publ. 243:176. 1936. — Tyee: CUBA, Pinar del Rio, Herradura, 26 Jun 1922, Ekman (HOLOTYPE: US! isoryprE: MO!) Digitaria ekmani Hitche. var. curtisst? Henr., Monogr. Digitaria 213. 1950. — Tyee: [UB Isla de Pinos, | Jun 1904, Cn 521 (HOLOTYPE: HAC; tsorypes: F!, Y!, US) Description: Plants perennial, lacking stolons; rhizomatous. Nodes glabrous or hairy. Sheath auricles 0.5 — 2.5 mm long. Sheaths glabrous or hairy. Ligule 1.5 — 2.5 mm long. Leaf blades flexuous; spreading; 5 — 22 cm long; 3-6 mm wide; hairy on the lower surface; hairy on the upper surface; with the midrib obviously differentiated (on the lower surface). Main axis mostly 15 —40(-60) mm long; with quaquaversal primary branches. Primary branches appressed or spreading from the main axis; not whorled; 4—9 on the main axis; 0.2 — 0.3 mm wide. Pedicels about 2.2 mm long (with relatively long narrow lateral pedicels). Spikelets 40 — 120 on a typical primary branch; oblong or elliptic; 2.2—2.5 mm _ long, 0.6—0.7 mm wide. First glume always completely absent. Second glume | times spikelet length; 3 to S-nerved; glabrous or hairy; acute. Lemma of lower floret 7-nerved; acute; glabrous or hairy. Lower lemma hairs shorter than the upper floret; silvery. Upper floret 1 times the length of the lower floret. Lemma of upper floret yellow (soon becoming purple); acuminate to acute. Distribution: Cuba. Remarks: Digitaria ekmanii is currently known only from Cuba, however, we feel that future collections will likely show that it occurs elsewhere in the Caribbean. Spikelets of this species, specifically on the lower lemma and second glume, may be either glabrous or hairy. As with other species of Digitaria especially in this section, little or no taxonomic significance can be applied to this variation since it does not correlate with other characteristics. Specimens with glabrous spikelets (e.g. E. L. Ekman 1052 & 14086) are otherwise essentially identical to those with a fine line of white villous hairs in the internerve spaces (e.g. E. L. Ekman 10938 & 11310) varies relative to the density of the hairs. Additional important characteristics of this species include the presence of a pronounced raised 157 midnerve on the lower surface of the leaf blades, relatively long and filiform pedicels, and the complete absence of the lower glume. Finally, the spikelets are narrowly ovate to elliptic or nearly oblong. Representative specimens: CUBA. Isla de Pinos, Apr 1904, A. H. Curtiss 5.n. (NY); Isla de Pinos, Nueva ce Pee! 1904, A. H. Curtis 521 (E MO, NY, US); Isla de Pinos, Santa i 2 Nov 1920, E. L. =n 12021 (E NY). Pinar del Rio: Herradura, 26 Jun 1922, E. L. Ekman 1052 Loe NY, US); Bea 21 Jun 1922, E. L. Ekman 14086 (NY), i rroyo Nantua, Damuje, 27 May 1920, E. . Ekman 10938 (E NY); between Remates and La Fé, 17 Jun 1920, E. L. Ekman 113 oe NY, ee Santa Clara: Sabanas des Manaca, near Asiento Viejo, 25 Jul 1920, Bro. Leon 9294 (NY DicitrariA ERIOSTACHYA Mez, Bot. Jahrb. Syst. 56, Beibl. 125(4):80. 1921. — Typr: PARAGUAY, Balansa 146 (HOLOTYPE: L). Digitaria fallens L. Parodi, Revista. Soc. Arg. Ciencias Naturales 8:375. 1926. — Typr: ARGENTINA, Parods 7130 (HOLOTYPE: BAA; tsoryPe: US!) Description: Plants perennial; stoloniferous; rhizomatous or lacking rhizomes. Nodes glabrous. Sheath auricles 1-2 mm long. Sheaths glabrous. Ligule 1-3 mm long. Leaf blades flexuous; spreading; 3 — 20 cm long; 3—8 mm wide; glabrous on the lower surface; glabrous on the upper surface; with the midrib not obviously differentiated. Main axis 20 — 40 mm long; with quaquaversal primary branches. Primary branches appressed to the main axis to spreading; not whorled; 4—7 on the main axis; 0.2 — 0.3 mm wide. Pedicels 2— 3 mm long. Cleistdaamons inflores- cence absent. Spikelets 36 — 60 on a typical primary branch; lanceolate or elliptic; (2.2 —)2.4—2.9 mm long; 0.6—0.8 mm wide. First glume absent (occ. present as a minute scale ca. 0.1 mm long). Second glume | times spikelet length; 3 to 5-nerved; hairy; acuminate to acute. Lemma of lower floret 7-nerved,; acuminate to acute; hairy. Lower lemma _ hairs ee the upper floret (by 0.2—0.5 mm); white. Upper floret ().92 — 1 times the length of the lower floret. Lemma of upper floret grey or yellow; acuminate. Distribution: Paraguay and Argentina. Remarks. Presence of long silver hairs, which turn purple at maturity, makes D. eriostachya a distinctive easily recognizable species within this section. This hair type, the absence of hairs between the mid-nerve and first lateral nerve, spikelet shape, relatively long slender pedicels, and the presence of secondary branching are features of D. eriostachya that indicate a relationship with section Trichachne; however, other characteristics of D. eriostachya support its retention in the Aeqaig/umae. Additional important diagnostic features of this species include the pronounced long stolons, glabrous leaves, and spikelet length and shape. 158 Re eee specimens: ARGENTINA. Corrientes: ee -Grande, 4 F ee 126, Parodt 7130 (US ); Mburucuya, "Santa Maria”, | Nov 1950, T. M. Petersen 835 (US); San Miguel, Loreto, 12 Mar 1972, C. Quarin 567 (US); Em ee ee Gonz ez y Ruta 12, 2 b 1974, C. Quarin et al, 2229 (US); Saladas, Estancia Bovril, ¢ 1949, G. Schwarz 9143 a. Encre Rios: Federacion Santa Ana, 7 Feb 1963, A. nee 22416 (US). PARAGUAY. Caazapa: Barrento, 14 Mar 1950, E. Anderson 1138 (US): Pindafoy, Yutt, Jan 1949, Rosengurtt 546] (US). Central: In regione lacus Ypacaray, 1913, E. Hassler 11565 (US); Prope ees Sears date unknown), E. Hassler 13018 (US); Trinidad, Asuncion, Mar 1942, T: Royos (US). Misiones: Santiago, 19 Jan 1952, Rosengurtt B-6117 (US). Seas Yacare, Pilar, 22 Jan 1949, Rosengurtt B-5525 (US). Paraguart: Kn 198+ ruca Asuncion - era 16 Jan 1949, Rosengurtt B-5412 (US). Dictraria LANUGINOSA (Nees) Henr., Meded. Rijks-Herb. 61:5: 1930. Paspalum lanuginosum Nees, Agrost. Bras. 63. 1829. — Type: ORIGINALLY AT ERLIN NOW EVIDENTLY DESTROYED. Description: Plants perennial; stoloniferous or lacking stolons; rhizoma- tous. Nodes hairy (usually pilose). Sheath auricles about 1.5 mm long. Sheaths hairy. Ligule 1.5 — 2.2 mm long. Leaf blades flexuous; spreading; 3— 12cm long; 3 — 6 mm wide; usually hairy on the lower surface; usually hairy on the upper surface; with the midrib not obviously differentiated. Main axis 10 — 20 mm long; with quaquaversal primary branches. Primary branches appressed or spreading from the main axis; whorled at the lower nodes or not whorled; 4 — 8 on the main axis; 0.4 — 0.5 mm wide. Pedicels 2—4 mm long. Cleistogamous inflorescence present. Spikelets 20 — 40 on a typical primary branch; lanceolate to ovate; 2.6 —3.1(-3.5) mm long; 7—0.8 mm wide. First glume absent. Second glume | times spikelet length; 5 to 7-nerved; hairy; acuminate. Lemma of lower floret 7-nerved; acuminate or acute; usually hairy. Lower lemma hairs shorter than the upper floret; white. Upper floret 0.88 — | times the length of the lower floret. Lemma of upper floret grey; acuminate to acute. Distribution: Southern Brazil, Uruguay, and northeastern Argentina. Remarks: Digitaria lanuginosa is closely allied to D. aeguiglumis and occa- sionally it may be difficult to distinguish between these taxa. Both have cleistcogamous inflorescences at the upper leaf nodes. In addition, these taxa possess similar spikelet shapes and the same relative length of the upper floret and lower lemma. The most significant difference between the taxa 1s that D. /anuginosa has short compacted rhizomes and is considered as a perennial, whereas D. aequiglumis is morphologically annual. Correlated with this difference is a smaller spikelet size and the presence of more hairs on the leaf blades and spikelet parts in D. danuginosa. When the basal parts of the plant are missing from a specimen it can be difficult to distinguish between these taxa. 159 Representative specimens: ARGENTINA. Corrientes: General Paz Pueblo Cercanias, 16 Oct 1945, T. 8. Tharrola 3544 (US); Mburucuya, "Santa Maria”, 28 Noiv 1965, R. git? ig (NY). Entre Rios: Isla del ie (freute a Rosario), 15 Dec 1937, A. Burkart 8860 (F). BRAZIL. Rio Grande do Sul: (precise location unknown), 1902, Ma/me 1570 SN 11 Apr 1946, J. R. ae 9045 ¢ (US); Fazenda pxpetimmental de Criacaé Bagé, 2 Dec 1945, J. . Swatlen 7579 (US); Itaqui, ek que vai de Itaqui para Alegrete, 25 Nov 1980, R. M. Klein G U. Pastore 11 —971 (US); Peloras, 4 May 1946, J. R. Swallen 9267 (US); ae eae do Palmar, 27 Apr i JR. Swallen 9232 (US); Uruaguatana, 9 Dec 1945, J. R. Swallen 7646 (US). URUGUAY. Montevideo: Cauasco, 27 Feb 1937, Rosengurtt B-1183 (US). Rocha: Laguna Negra, 20 Mar 1938, pee B-2615 (US). Salto: rio Uruguay y San Antonio Grande, 27 Feb 1937, Rosengurtt B- 943; Tacuarembo, Picada del Cuello, 12 Mar 1945, Rosengurtt B-4755 (US). Dicrraria Leucrres (Trin.) Henr., Meded. Rijks-Herb. 61:6. 1930. Panicum lencites Tein., Gram. Pan. 85. 1826. — Tyre: Type specimen not located. Milium velutinum DC., Cat. Plant. Horci Bot. Monsp. 126. 1813. ee Lag., sen. & Sp. Nov. 2. 1816, non ace firm (L.) Koeler, 1 Syntherisma aaa ae Proc. Biol. . Wash -191. 1906. Disttaria velutina (DC.) ch Proc. Biol. Soc. ca. “40:84. on non Digitaria velutina (Forsk.) . 1812. — Tver: from cultivated material derived from Mexican seeds (HOLOTYPE: fragment US!). Syntherisma velutina glabella Chase, Contr. U.S. Natl. Herb. 17:220. 1913. a leucites. (Yrin.) Henr. var. glebella (Chase) Henr., Monogr. Digitaria 395. 1950. Type: MEXICO, Michoacan, 16 Sep 1910, Hatchcock 6989 (HOLOTYPE: US! isorype: LL!, NY!). Digitaria distans (Chase) Fern., Rhodora 22:103. 1920. Syntherisma distans Chase, Contr. U.S. Natl. Herb. 17:220. 1913. — Tver: MEXICO, Jalisco, vicinity of Orozco, 29 Sep 1910, Hitchcock 7376 (HOLOTYPE: US!). Description: Plants perennial; stoloniferous; rhizomatous. Nodes glabrous or hairy. Sheath auricles 0.7 — 1.5 mm long. Sheaths glabrous or hairy (the lower sheaths usually hairy). Ligule 2.3— 3.5 mm long. Leaf blades flexuous; spreading; 6— 20 cm long; 2-5 mm wide; glabrous or hairy on the lower surface; glabrous or hairy on the upper surface, with the midrib not obviously differentiated. Main axis 15 —45 mm long; with quaquaversal primary branches. Primary branches s preading; whorled at the lower nodes or not whorled; 4—9 on the main axis; 0.3—0.4 mm wide. Pedicels 2—4 mm long. Spikelets 20-60 on a typical primary branch; elliptic; 2.4— 3.2 mm long; 0.9— 1.1 mm wide. First glume present or absent (mostly present as a hyaline truncate scale ca. 0.3. mm long). Second glume | times spikelet length; 5-nerved; hairy, acute. Lemma of lower floret (5 —) 7-nerved; acute; hairy. Lower lemma hairs subequal to the upper floret; usually purple. Upper floret 0.92 —~ | times the length of the lower floret. Lemma of upper floret grey or yellow; acute. Distribution: Mexico. 160 Remarks: Digitaria leucites is a distinctive perennial species occurring 1n south-central mountainous regions of Mexico. The most characteristic fea- ture of this species is the relatively plump spikelets in which the second glume and lower lemma do not tightly enclose the upper floret at maturity. The second glume and lower lemma are hairy with purple villose hairs, but frequently the internerve space between the midnerve and first lateral nerve is glabrous. This pattern of pubescence is common in other sections of this genus. Digitaria distans is known from two collections (Hitchcock 7376 & 7372), both collected on September 29, 1910 at Orozco, Jalisco, Mexico. Chase (1913) recognized these as a new species and used the distant and glabrous spikelets as key characters. McVaugh (1983) differentiated D. distans from D., feucites based on the absence of spikelet hairs in D. distans. Our study of all the available specimens of these taxa resulted in the follow- ing observations. The second glume and lower lemma of D. distans is glabrous whereas these structures in D. /eucites possess a line of mostly purple hairs between the lateral nerves. However, within specimens normally accepted as D. /eucites there exists a wide range of variation in these characters. For example, Lyonnet 1879 shows clearly distant nearly glabrous spikelets. It was concluded that D. distans is best treated as a syn- onym. Representative specimens: MEXICO. Chiapas: Mun. de Zinacantan, 5 Oct 1966, R. M. Laughlin 2325 (ENCB, TAES). Distrito Federal: Contreras, Primer Dinamo, 14 Jan 1969, I. Garcia S. 128 (ENCB); Pedregal de Tlalpan, 1932, E. Lyonnet 975 (MEXU); San Angrés, D: E; Aue 1930.2, Lyonnet 975 (MEXU), Carretera Cuernavaca, 23 Oct 1937, E. Lyonnet 1879 (ENCB, CHAPA, MEXU); base of Sierra de Ajusco, 29 Oct 1896, C. G. Pringle 6623 (ENCB, MEXU); Pedregal de San Angel, cerca de Eslava, 19 Oct 1952, J. Rzedowski 2008 (ENCB, MEXU). Hidalgo: 10 km al Este de Metepec, 7 Aug 1980, R. Hernandez M. & R. Hernandez V. 4716 (MEXU). Jalisco: Los Guaybos, (collection date not given), A. A. Beetle G Guzman M. 5440 (CHAPA); Sierra de Tigre, 3 mi S of Mazamitla, 18 Sep 1952, R. McVaugh 13029 (MEXU). Mexico: Mun. de Villa Allende, San Cayetano, Oct 1963, J. M. A lcocer s.n. C(ENCB); Chapingo, Terrenos de la E.N.A., Lomas de San Juan, 1 Oct 1965, R. Bonilla B. s.n (CHAPA),; Terrenos de la E.N.A., Xaltepa, 29 Sep 1966, R. Bonilla B. 5.n. -HAPA); Chapingo, Mun. de Tezcoco, Molino de las Flores, 19 Oct 1976, José Cantu s.n. (CHAPA); Chapingo, Mun. de Texcoco, 22 Aug 1968, ]. Flores Crespo s. n. (ENCB); 2 km E of Temamatla, 22 Aug 1972, J. Elias 203 (ENCB); roadside from San Juan del Rio to Mexico City, 6 Nov 1962, EF W. Gould 103 16 (ENCB, TAES); Chapingo, Mun. de Tezcoco, 2.5 km al E de Tezcoco, 14 Oct Oct 1976, E. Garcia M. 5.n. (CHAPA, TAES, US); Toluca, 13 Sep 1910, Hitchcock 1560 (LL, NY, TAES); Villa de Allende, 5 Oct 1952, E. Matuda 20429 (MEXU),; Valle de Bravo, 21 Nov 1952, BE. Matuda 27791 (MEXU); Mun. de Ixtapaluca, Cerro del Pino, 30 Oct 1976, S. Morelos O. 44 (ENCB); Mun. de Ixtapaluca, Ladera Sureste del Cerro del Pino, 3 Oct 1976, S. Morelos O. 116 CENCB); Mun. de Huehu- x 96 ENCB, TAES); Mun. de Ixtapaluca, Cerro del Pino, 3 Oct 1976, L. Rico R. 51 (ENCB): , Mun. de Ixtapaluca, laderas inferiores SE del Cerro del Pino, 3 Oct 1976, Rzedowski 34423 161 (CHAPA, pa Chapingo, Edo. Mexico, (collection date not given), J. Tovar G E. Garcia s.n. (CHAPA), alrededores de San Pedro Nexapa, 2 Nov 1963, Marina Villegas D. 276 (ENCB). Tata Mun. Villa Escalante, 24 Oct 1981, ]. Garcia P1555 (CHAPA, ENCB); 20 km S Zamora, 28 Sep 1946, E. Hernandez ae (CHAPA), Rae caee . Sep ae Hitchcock 1561 (NY, | fe an, Hitchcock 6989 (US); Patzcuaro, 19 Oct 8 . D. Holway 3212 (US); Mun. Tangancicuaro, Las Canas, 19 Nov 1971, Eee y eae 612 (ENCB); NE side. ef the Voledn de pea : Oct 1953, E. R. Sohns 809 (TAES); NE side of the Volcan de Paricutin, 4 Oct 1953 R. Sohns 822 (TAES). Morelos: =: mi N of Toll gate, Cuernavaca, 10 Nov 1962, F aie 10388 (TAES, US); 60 km x.-Cuernavaca, “Campo Turista,” 7 Sep 1952, FE Gallegos Harkings 499 (MEXU),; Trés Marias (Camino de Cuernavaca), Jul 1927, E. Lyonnet 58 (MEXU)); Valle del Tepeite, 17 Sep 1938, E. Lyonnet 2442 (MEXU). Oaxaca: 42 km de Putla rumbo a Tlaxiaco, 23 Jun 1980, A. A. Beetle M-4721 (CHAPA), Campamento Rio de Molino, 4 km al SW de San Miguel Suchistepec, 21 Sep 1965, J. Rzedowski 21025 (ENCB); District of Ixtlan, a Cumbre del Cuartel, 2 Nov 1944, J. V. Santos 3619 (CHAPA, NY). Tlaxcala: Mun. de Salvador, Tzompantepec, 6 Sep 1982, H. Vibrans 1187 (ENCB). DictrarRia PAUCIFLORA Hitchc., Proc. Biol. Soc. Wash. 41:162. 1928. — Type: U.S.A., Florida, Jenkins to Everglade, 10 Nov 1903, Eaton 207 (HOLOTYPE: US!) Description: Plants perennial; lacking stolons; rhizomatous. Nodes most- ly glabrous. Sheath auricles about 1.5 mm long. Sheaths hairy (becoming glabrous with age). Ligule 1.5 — 2.0 mm long. Leaf blades flexuous or twisted; spreading; 7— 18 cm long; 1.0—2.2 mm wide; hairy on the lower and upper surface (becoming glabrous with age); the midrib not obviously differentiated. Main axis 10-80 mm long; with quaquaversal primary branches. Primary branches appressed or spreading from the main axis; not whorled; 2 — 8 on the main axis; 0.43 mm wide. Pedicels 2— 3 mm long. Spikelets 30 — 60 on a typical primary branch; lanceolate; 2.7 — 3.0 mm long; 0.7 —0.9 mm wide. First glume commonly present. Second glume | times spikelet length; mostly 7-nerved; glabrous; acuminate to acute. Lemma of lower floret 7-nerved; acuminate to acute; glabrous. Upper floret 1 times the length of the lower floret. Lemma of upper floret becoming purple; acuminate to acute. Distribution: Southern Florida. Remarks: Digitaria pauciflora is known from the Everglades region of southern Florida. Inflorescence and spikelet characteristics are similar to those of D. simpsoniz. However, they differ significantly on vegetative characters. Representative specimens: U.S.A. Florida: Dade Co.: Everglades 7 ational Park, 6th glade, 19 Jun 1978, G. N. Avery 1928 (F); Everglades National Park, Long Pine Key, W edge of 3rd glade, Block H, 20 Jun 1978, G. N. Avery 1929 (F);, E dea National Park, Long Pine Key, Block D, 16 Jun 1978, G. N. Avery 1932 (F); Everglades National Park, S$ of Long Pine Key Road in 6th glade, 20 Oct 1978, G. N. Avery 1979 (F); Jenkin’s Home- 162 stead, 14 — 20 mi S of Cutler, (collection date unknown), A. A. Eaton s.n. (E US); Evergla- des National Park, Long Pine Key, Glade #6, 28 Apr 1986, A. Herndon 1519 (F); In pinelands, South Miami, 2 Oct 1939, W. A. Silveus 5285 (TA between Cutler and Longview Camp, 9 Nov 1903, /. K. ai & J. J. Carter 916 (NY DictraRiA SABULICOLA Henr., Blumea 1:108: 1934. — Tyrer: BRAZIL, Provincia de Espirito Santo, 1816— 1821, (HOLOTYPE: P; IsoTyPE US)). Description: Plants perennial; lacking stolons; rhizomatous. Nodes glabrous. Sheath auricles about 0.6 mm long. Sheaths glabrous. Ligule 1 —2 mm long. Leaf blades flexuous; spreading; 2—5 cm long; 1.5 — 2.5 mm wide; glabrous or hairy on the lower surface; hairy on the upper surface; with the midrib obviously differentiated. Main axis 10-25 m long; with quaquaversal primary branches. Primary branches appressed to the main axis; whorled at the lower nodes or not whorled; 3 — 9 on the main axis; 0.2 mm wide. Pedicels 1.5 — 2.5 mm long. Spikelets 32—70 ona typical primary branch; lanceolate; 2.1—2.3 mm long; 0.4—0.5 mm wide. First glume absent. Second glume | times spikelet length; 5-nerved; hairy; acuminate. Lemma of lower floret 7-nerved; acuminate; hairy. Lower lemma hairs shorter than the upper floret; white. Upper floret 0.9 times the length of the lower floret. Lemma of upper floret yellow; acumi- nate. Distribution: Brazil. Remarks: The present concept of D. sabulicola is based on two collections from Brazil. Further collections are needed to better understand the full range of morphological variation for this species and its affinities within the section. The principle diagnostic character is spikelet length which is shorter than that found in the apparently closely related taxa, which in- clude D. aequiglumis, D. cuyabensis, and D. lanuginosa. Representative specimens: Brazil. Bahia: Joazeiro, near Rio Sao Franciso, 13 Dec 1924, Chase 7910 (US). Espirito Santo: Santo, Voyage d’Auguste de Saint Hilare, from 1816 — 1821 (precise location and colleciton date not given) (US). DiGtraRIa SIMPSONII (Vasey) Fern., Rhodora 22:103. 1920. Panicum hs inale vat. simpsonti Vasey, Contr. U.S. Natl. Herb. 3:25. 1892. Panicum simpsonii ae y) Beal, Grasses of N.A. 109. 1896. Pre He (Vasey) Nash, bal Torrey Bot. Club 25:297. 1898. — Type: U.S.A., Florida, Manatee, 1890, oe (HOLOTYPE: NY!; tsorype: US!). Description: Plants perennial; lacking stolons; rhizomatous. Nodes most- ly glabrous. Sheath auricles 0.7 — 1.5 mm long. Sheaths hairy (becoming glabrous with age). Ligule 1.5—2.5 mm long. Leaf blades flexuous; spreading; 6 — 20 cm long; 3—55 mm wide; hairy on the lower surface; hairy on the upper surface (becoming glabrous with age); with the midrib 163 not obviously differentiated. Main axis 40—70 mm long; with qua- quaversal primary branches. Primary branches spreading from the main axis; not whorled; 6 —9 on the main axis; 0.3 mm wide. Pedicels 1.5 — 2 mm long. Spikelets 40 — 50 on a typical primary branch; ovate to lanceo- late; 2.9—3.1 mm long; 0.7—0.9 mm wide. First glume typically absent. Second glume | times spikelet length; mostly 7-nerved; glabrous; acuminate to acute. Lemma of lower floret 7-nerved; acuminate to acute; glabrous. Upper floret 1 times the length of the lower floret. Lemma of upper floret becoming purple; acuminate to acute. Distribution: Florida. Remarks: According to Nash (1898) the type material for D. s¢mpsonit was taken from cultivated material originally collected from Long Key southwest of Sarasota Bay, Florida. The length of time that the plants were in cultivation before the type material was collected is unknown. The name has been applied to two collections from Florida and material from Cuba. The Cuban collections are D. ekmanii. The two collections from Florida are the holotype from Manatee, Florida and Curtiss 6422 from St. Augustine, Florida. Spikelet and vegetative characters differ between these collections and it is obvious that they belong to different species. Curtiss 0422 is a perennial with a decumbent base, the leaves are hairy with pilose or setac- eous indumentum, and the spikelets are about 2.5 mm long and glabrous or with a few short purple hairs between the lateral nerves. All characteris- tics found in Cyrtiss 6422 are also found in D. texana. Specific examples of D. texana that possess characteristics of Curtiss 0422 include Swallen 1533, H.R. Reed s.n., and Swallen 10574. It was concluded that Curtiss 0422 ts a disjunct collection of D. texana. Therefore, this species is only known from the type material and its present status under natural conditions ts un- known. Representative specimens: U.S.A. Florida: Manatee, garden of J. H. Simpson, 1890, Simpson s.n. (NY, US) DictrariA TEXANA. Hitchc., Proc. Biol. Soc. Wash. 41:162 1928. — Tyee: U.S.A., Texas, 6 — 27 — 1910, Hitchcock 5479 (HOLOTYPE: US!). Digitaria runyontt oe , J. Wash. Acad. Sci. 23:455. 1933. — Type: U.S.A., Texas, 21 Apr 1929, ae 188 (HoLorype: US!). Digitaria subcalva Hitchc., Amer. J. Bot. 21:138. 1934, syn. nov. — Tyee: U.S.A., Florida, Plant City, 26 Oct 1932, C. p eer 1956 (HOLOTYPE: US!; IsoTYPE: a Di Digitaria albicoma Swallen, J. W ad. Sci. 30:214. 1940, syn. nov. TYPE: U.S.A., Florida, 18 Nov oe ee 5644 (HOLOTYPE: US!). 164 Description: Plants perennial; stoloniferous; rhizomatous. Nodes glabrous. Sheath auricles 0.5 — 1.5 mm long. Sheaths glabrous or hairy. Ligule |— 2 mm long. Leaf blades flexuous; spreading; 2.5 — 20 cm long; 2—6mm wide; glabrous or hairy on the lower surface; glabrous or hairy on the upper surface; with the midrib obviously differentiated. Main axis 10 — 70 mm long; with quaquaversal primary branches. Primary branches spreading; whorled at the lower nodes or not whorled; 5 — 12 on the main axis; 0.31 —0.4 mm wide. Pedicels |— 2.5 mm long. Spikelets 18 — 65 on a typical primary branch; lanceolate to ovate; 2.3 — 3(-3.2) mm long; 0.51—0.7 mm wide. First glume absent (occ. present as a small hyaline scale). Second glume (0.9—)1 times spikelet length; (3 —)5-nerved; glabrous or hairy; acuminate to acute. Lemma of lower floret 7-nerved; acuminate to acute; glabrous or hairy. Lower lemma hairs shorter than the upper floret; white. Upper floret 0.95 — | times the length of the lower floret. Lemma of upper floret yellow or purple; acuminate to acute. Distribution: Texas and occ. introduced in Florida and Mesoamerica. Remarks: The Digitaria texana - runyonit complex occurs on sandy coastal areas of southern Texas from Calhoun to Cameron County. One collection reported from Brazos Co., Texas {Reeves 1040 (TAES)} probably represents an incorrectly labeled specimen. There is one collection from Veracruz, Mexico [Hitchcock 6554 (LL, NY, US)]}, and future collections may show that this complex commonly occurs on sandy coastal areas in Tamaulipas. Hitchcock (1950) and Gould (1975) ¢ distinguished D. texana from D. runyont on spikelet length and vestiture of second glume and lower lemma; however, Correll and Johnston (1970) united these species. Our study of this complex indicated an interesting correlation between the morphological forms and habitat. These taxa are commonly associated with coastal sand dunes but extend inland for about 75 miles. The inland form has spikelets usually 2.3 — 2.6 mm long with the second glume and lower lemma glabrous to sparsely hairy. Examples of this form include H. R. Reed s.n. (US), Swallen 1533 (US), Cory 28346 (TAES), Runyon 2753 (NY), and W. A. Sz/veus 7310 (TAES, US). This small-spikelet form occurs on the coast but differs in that the second glume and lower lemma is dis- tinctly hairy with villous hairs between the lateral nerves. A number of specimens, including the holotype of D. fexana, are intermediate between these forms. The other morphological form identified in this complex occurs in coastal sandy areas. Vegetative characteristics overlap with the inland form; however, the spikelets are about 2.8 — 3.2 mm long and the outer bracts distinctly hairy. The holotype of D. rvmyonii falls into this group. Intermediates [Lundell 15029 (NY), Swallen 10563 (US), and Suwallen [0611 (US)} are common between the small and large spikelet 165 forms. Based on these observations it was concluded that D. runyonii is best placed in synonymy under D. texana. Digitaria albicoma is known from two collections, the holotype (collected in 1938) and a second incomplete specimen collected five years later at the same locality (Chinsegut Hill Sanctuary, Brooksville, Florida). These specimens possess the following significant characteristics: perennial with densely villose leaf sheaths; leaf blades long, narrow, with a pronounced mid-nerve; primary inflorescence branches lacking spikelets at the base; spikelets ca. 2.6 mm long, nearly glabrous but with a few purple hairs in the internerve spaces. These characters can be found in the range of vari- ation accepted in D. texana. Some specific examples of D. texana exhibiting these characters are Swallen 1856, 1533, & 1408 —A, all of which occur on sandy coastal areas. It was concluded that D. albicoma is best treated as a synonym of D. texana. A similar situation exists for D. subcalva, which is also treated as a synonym. The presence of these specimens in Florida in- dicates that D. texana is occasionally introduced but fails to persist. It ts interesting to note that these specimens were collected close to the 28th latitude, which is the same latitude where D. texana is native and con- centrated in the Texas coastal Bend region. Representative specimens: MEXICO. Veracruz: Veracruz, 31 oe ee A.S. aie 6554 (LL, NY, US). U.S.A. TEXAS: Aransas Co.: Aransas near Bay, 24 Nov 1932, W. Silvews 847 (US); Port Aransas Pass, 24 Nov 1940, W. A. Sr/vens 67 70) ea id port, 15 Oct 1941, W. A. Silvens 7320 (TAES); Port Aransas, 10 Nov 1941, W. A. Sifvens 7450 (TAES); Copano Bay, E side, 25 Nov 1931, B. C. Tharp 7908 (US, NY). Brazos Co.(?): Fall 1940, R. G. Reeves 1040 (TAES). Brooks Co.: 4 mi SE of Encino Division, King Ranch, 18 Nov 1954, FE W. Gould & J. Morrow 6728 (TAES), Falfurrias, 26 Jun 1936, H.R. Reed s.n. (US); Santa Fe Division, King Ranch, 3 Nov 1949, J. R. Swallen 10597 (US). panes Co.: sand below Seadrift, | Dec 1928, Tharp 5073 (US). Cameron a 0.: mouth o inde, 21 Apr 1929, R. Runyon 188 (US), Brazos Santiago Island, 1938, RK. ae 1878 (US); Brazos Santiago Island, 7 Oct 1938, R. Ranyon 2010 (F . Kenedy Co.: Sarita, 27 Jun 1910, A. S. Hitchcock 5479 (US); Rags Ranch, 8 mi S of Sarita, 15 1946, Lundell & Lidell 14701 (US); Kenedy Ranch, N of Mifflin, 3 Nov 1949, C. i oe 15029 (NY); between the South border and Los Norias, 11 May 1941, R. Runyon 2783 (NY); near Encino, 30 Apr 1932, W. A. Silveas 575 (US); Sarita, 14 Nov 1941, W. A. Silvens 7311 (TAES, US); Sarita, 6 Apr 1931, J. R. ru 1408 — A (US); Sarica, 17 Apr 1931, J. R Scaler a 3 (US); Sarita, 17 Apr 1931, J. R. Swallen 1533 (US); Riviera co Riviera each 8 Jun 1931, 7. R. Salle 1856 (US); King eee Norias ma , San Jose pasture, 2 Nov 1949, J. : Swillen 10574 (US); King Ranch, Norias Div., San Jose | rasture, 2 1949, J. OR. Swatlen 10579 (US); between Mifflin and pee 2 Nov 1949, J. R. Swallen {0581 (TAES); 1% m1 S$ of Mifflin, 2 Nov 1949, J. R. Swallon ae (US); N of cae 3 Nov 1949, J. R. Swallen 10610 ee of ee 3 Nov 1949, J. R. Swallen 10611 (US). Kleberg Co.: Padre Island, 25 N 1932, W. alate 848 (NY, US); ee Ranch, —_~ oo Div., | Nov 1949, J. R. Staller 10563 Hee I} mi Sof Riviera, 8 Oct . Parks & V. L. Cory 16989 (TAES). Nueces Co.: ae a oe Nov 194( ie 20936 (TAES, US); 15 mi S of Corpus Christi, | Oct 1931, W. A. Silvers 3 pe 166 Mustang Island, 18 Oct 1975, S. R. Hill 3843 (TAES); 10 mi S of raed ee oS 1931, J. R. Swallen 1829 % (US). Refugio Co.: Copano Bay, 26 F G, rp 43189 (NY). Willacy Co.: Raymondville, 4 Apr 1938, V. L. Cory sie eee . mi N of Raymondville, 14 Nov 1941, W. A. Silvews 7305 (TAES); between Laguna Madre and Raymondville, 14 Nov 1941, W. A. Svlvews 7310 (TAES, US) ACKNOWLEDGMENTS Appreciation is extended to John Wiersema and Jay Wipff for making suggestions to improve the wording of this manuscript. Vittoria Hess provided assistance in all aspects of data collection, storage, analysis, and presentation. REFERENCES AGRASAR, Z. 1974. The species of the genus Digitaria (Gramineae) in Argentina. Darwi- niana 19(1).65 — 166. BOONBUNDARL, S. 1985. A biosystematic study of the Digitaria leucites complex in North America. oo Department of Range Science, Texas A & M University, College Station BURKART, A. ae a Ilustrada De Entre Rios (Parte II, mune Distributed by Librart S.R.L., Corrientes 127, Buenos Aires, Argentin CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual vee vascular plants of Texas. Texas Research eae Renner, esas. DALLWITZ, M. J. 1974. A flexible computer system for generating identification keys. Syst. Zool. eee: 1980. A general system for coding taxonomic descriptions. Taxon 29:41 — 46. GOULD, F W. and T. W. BOX. 1965. Grasses of the Texas Coastal Bend. Texas A & M University Press, College Station, Texas. GOULD, E W. 1975. The grasses of Texas. Texas A & M University Press, College Station, Texas. ar J. E 1950. Monograph of the genus Digitaria. Universitare Pers, Lieden, = 992. se. A. S. 1922. Grasses of British Guiana. Contr. U.S. Natl. Herb. 22:466. 1922 1927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24(8):291—556. 1936. Manual of the grasses of the West Indies. U. S. Dept. of Agric. Misc. Publ. No. 243, Washington D. C. 1951. Manual of che grasses of the United States, 2nd ed. Revised by A. Chase. U.S. Dept. Agric., Misc. Publ. No. 200, Washington, D.C. McVAUGH, R. 1983. Flora Novo-Galiciana: A descriptive account of the vascular plants of Western Mexico (William R. Anderson, ed.), Gramineae (Volume 14). The Univer- sity of ae Press, Ann Arbor, Michigan. NASH, — Genus Syntherisma in North America. Bull. Torrey Bot. Club 2a POHL, R. on a aad in William Burger, ed., Flora Costaricensis. Fieldiana Bot., New Ser., No. 167 ROSENGURTT, B., B. ARRILLAGA M., P IZAGUIRRE A. 1970. Gramineas Urugu- ayas. Universidad de La Republica, Departmento de Publicaciones, Montevideo. SMITH, , D. C. WASSHAUSEN, and R. M. KLEIN. 1982. Gramineae in Raulino Reitz, i Hes Hustrada Catarinense, National Museum of Natural History, Smithso- nian thas. ares eC, VELDKAMP, J. FE 1973. A re of Digitaria Haller in Malesia. Blumea 21:1 — 80. WEBSTER, R. 1983. A revision of the genus Digitaria (Poaceae: Paniceae) in Australia. Brunonia 6(2):131— WEBSTER, R. 1987. Taxonomy of Digitaria section Digitaria in North America (Poaceae: Pp ee hae 12¢1):209 — 222 WEBSTE 1988. Genera of he North American Paniceae (Poaceae: Panicoideae). Syst. 13(4):576— fee WEBSTER, R. and J. VALDES R. 1988. Genera of Mesoamerican Paniceae (Poaceae: Panicoideae). Sida 13(2):187 — 221 WEBSTER, R., J. H. KIRKBRIDE, and JESUS VALDES R. 1989. New World genera of the Paniceae (Poaceae: Panicoideae). Sida 13(4):393 —417 Book REVIEWS Copy, WitiiaM J. 1988. Plants of Riding Mountain National Park, Manitoba. 320 pp. 323 illus. of selected species. Price unknown. Agriculture Canada Publication 1818E, Canadian Government Pub- lishing Centre, Supply and Services Canada, Ottawa, Canada K1A OS9. With keys to the species and a checklist of the species in addi- tion to the Index (species epithets and common names). FRANKTON, CLARENCE AND GERALD A. MULLIGAN. Weeds of Canada. 217 pp. 101 plates include complete or partial drawings of 154 species. There are 230 species described in the text. $16.95 paper. Agriculture Canada Publication 948, NC Press Limited, Box 4010, Station A, Toronto, Ontario MSH 1H8. Keys absent, identification by excellent tllustrations. Erskine, D. S. 1985. The plants of Prince Edward Island (Reprinted rom Agriculture Canada 1088, 1960), with new records, nomenclatural changes, and corrections and deletions. Catling, P M., D. S. Erskine, and R. B. MacLaren. 272 pp. plus map. $16.50 in Canada; $19.80 for other Countries. Agriculture Canada Publication 1798, Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Canada K1A 089. This popular flora 1s reproduced with the updated version given in the initial pages iv — Xxil. Aiken, 8S. G. AND S. J. DarBysuire. 1983. Grass genera of Western Canadian cattle rangelands. 173 pp. 46 figures of drawings with one photograph. $9.00 in Canada; $10.80 for other Countries. Agri- culture Canada Monograph 29, Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Canada K1A 0S9. Keys and descriptions accompany the diagrammatic illustrations. Sipa 14(2):168. 1990. THE PLUMBAGINACEAE IN THE FLORA OF THE SOUTHEASTERN UNITED STATES JAMES L. LUTEYN The New York Botanical Garden Bronx, NY 10458-5126, U.S.A. ABSTRACT The Plumbaginaceae is represented in the native southeastern flora by Limontum carolint- co. anun, Limon limbatum, and Plumbago scandens. Plumbago auriculata is widely cultivatec and has become naturalized. This paper describes, illustrates, and maps these species in the southeastern flora. Las Plumbaginaceas estan representadas en la flora nativa del sureste por Limoninm caroli- manum, Limontum linbatum, y Plumbago scandens. Plumbago auriculata es cultivado ampli- amente y se ha naturalizado. En este papel se describen, iluscran, y presentan mapas de cn eee para estas Cuatro especies para la flora del sureste The Plumbaginaceae contains 12 genera and about 400 species distrib- uted throughout the world. It is best developed in the Mediterranean and the Middle East, mostly in xerophytic situations, on saline and calcareous soils (Luteyn 1990). In the United States, three genera, Armeria, Limonium, and Plumbago, occur natural y. Limonium is either a salt marsh plant with one species found along the entire Atlantic and Caribbean sea- board {L carolinianum (Walter) Britton} and another along the Pacific coast from southern California to northern Oregon {L. californicum (Boiss.) Hel- er], or an inland species in sale flat areas in Texas, Oklahoma, New Mexico, Arizona, and northern Mexico (L. /imbatum Small) (Luteyn 1976). Several species are cultivated and used in dried floral arrangements; two of these have escaped and become naturalized in southern California [viz., L, perezit Hubb. and L. sinvatum (L.) Miller}. Plumbago scandens L. is native to southern Florida, Texas, and Arizona, and ranges south through Central and South America. Plambago auriculata Lam. (= P. capensis Thunb.), a nat- ive of South Africa, is widely cultivated and has become naturalized locally in Florida. In the United States, Armeria maritima Willd. is native on bluffs and sandy places along the Pacific coast as far south as San Luis Obispo County, California. Several other species of Armeria are cultivated, mostly as rock garden plants. This treatment was originally prepared in 1976 (Luteyn, in press) for the “Vascular Flora of the Southeastern United States” (Massey et al., Editorial Board). It follows the basic format for that flora as outlined in Radford et — Sipa 14(2):169— 178. 1990. 170 al. (1967), although herein | have included illustrations and distributional maps. For geographical completeness, | have also included the entire state of Texas within the range of coverage fully realizing that some parts of the state (specifically that west of the 100th meridian) may not be phytoge- ographically “southeastern.” TAXONOMY PLUMBAGINACEAE. THE LEADWORT FAMILY Perennial herbs, subshrubs, or climbing plants. Leaves simple, alter- nate, basal or cauline, entire; petioled with bases persistent and often sheathing the stem; exstipulate. Inflorescence of terminal or axillary racemes or panicles, often spike-like; floral nodes bracteate. Flowers perfect, actinomorphic, 5-merous, hypogynous, bracteate, homostylous or heterostylous; calyx synsepalous, plicate, 5-ribbed, sepals scarious; corolla sympetalous or of nearly distinct petals, marcescent, exserted beyond the calyx, lobes convolute-imbricate; stamens 5, distinct, sometimes epipeta- lous, then borne on the corolla tube opposite the lobes, anthers introrse; stigmas 5, linear, styles | or 5, ovary S-carpellate, |-loculed, usually 5- ribbed, ovule solitary, anatropous to circinotropous. Fruit a capsule or utr- icle, partly or totally enclosed by the persistent calyx. Seed with straight embryo, endosperm white, mealy. Channell, R.B. & C.E. Wood, Jr. 1959. The genera of Plumbaginaceae of the southeastern United States. J. Arnold Arbor. 40:391— 397. — 1. Leaves basal; corolla of nearly distinct petals, less chan | cm long; styles 5, PSC tO base iy soe ede tates sere eee tee eee ees 4 1. Limonium Leaves cauline; corolla salverform, more than | cm long; style 1, with 5 SH OMaS sic.cx pipe ecan ee een hee sane eee See bisa ei ees Fe 1. LIMONIUM Miller Sea-LaveNbEeR Scapose, acaulescent, vegetatively glabrous herbs, with short to el- ongate, thick taproots. Leaves basal, equitant, coriaceous, long attenuate. Inflorescence of terminal panicles or corymbs, with the ultimate branch tips bearing secund, solitary or usually few-flowered spikelets. Flowers homostylous, hypogenous, sessile or nearly so, subtended by 3 (or more) sheathing bracts; calyx tubular-funnelform, 5-ribbed, limb plicate, lobes oblong to triangular, usually with smaller intervening lobes or dentate- erose sinuses; corolla with petals nearly distinct, long-clawed; stamens adnate to the base of the corolla, included; styles 5, distinct to the base, with 5 linear-clavate, papillate stigmas. Fruit a brownish-green utricle, usually exserted from the persistent calyx, usually capped by the marces- cent corolla and style bases. 171 Luteyn, J. L. 1976. Revision of Limoninm (Plumbaginaceae) in eastern North America. Bricttonia 28: 303 — 317 Calyx limb erect at maturity; spikelets loosely to moderately or even Ll. L. carolintanum ea aggregated; plants of coastal sale marshes or flats. ...... Calyx limb wide-spreading or flaring at maturity; spikelets always densely 2. L. limbatum aggregated; plants of inland alkaline areas Ll. L. CAROLINIANUM (Walter) Britcon. — Stems 1—9.5 dm tall. Leaves elliptic, spatulate, obovate oblanceolate, rarely linear or semior- bicular, 5 — 15 (30) cm long, 0.8 — 4 (7.5) cm wide, apex rounded or acute to retuse, deciduously cuspidate. Inflorescence with spikelets loosely to moderately densely aggregated, the floral internodes 0.5 — 10 mm long. Flowers perfect, rarely male-sterile, solitary or 2—3 (5) clustered, bracts 3—6 mm long; calyx obconic to funnelform, (3.8) 4.5 —6.5 (7.5) mm long, tube glabrous to densely pilose along |1—5 ribs, limb erect at aturity, lobes oblong to narrowly triangular, 0.4—1.5 (2) mm long petals lavender, rarely white. Fruit 3 — 5.5 mm long. (n = unreported) Spr- Salt marshes and salt-flats along Atlantic and Gulf _ Incl. Limonium angustatum (Gray) Small; L. obtust- . var. trichogonum Blake; L. n. var. t. £. Teg: ing, summer, early fall. seacoasts. Figs. | and 2 lobum Blake; L. nashit Small. L. n albiflorum (Rat.) House; L. carolinianum var. angustatum (Gray) Blake; var. obtustlobum (Blake) Ahles; L. x. var. angustatum (Gray) Ahles. My studies (Luteyn 1972, 1976) including field observations from much of its extensive geographical range (Labrador and Bermuda to Tamaulipas, Mexico), indicate that numerous local populations have resulted in a polymorphic species. Morphological variation is almost continuous throughout the range, and therefore, the recognition of several taxa is un- justified. Elaboration of the different morphologies is unnecessary; how- ever, a word about the reproductive biology is in order and may shed light on community structure. Limonium carolinianum reproduces sexually and is self-compatible (Baker 1953). Its seeds are dispersed by birds and ocean currents. However, seedlings are very rare, and the spread of populations is vegetative from the horizontal rhizomes and short laterals of the branched woody stock (pers. observ.). With time, extensive clones (each possibly with distinctive morphologies) may spread and coalesce within the community. 2. L. uimparum Small. — Stems to 6 (10) dm tall. Leaves spatulate, oblong-spatulate, obovate to elliptic, 4— 16 cm long, 1.5 — 6.5 cm wide apex rounded or retuse, shortly mucronate. Inflorescence with spikelets distichously and densely aggregated, the floral internodes 0.5 — 3 mm long; calyx funnelform to salverform, 3.3 —5 mm long, tube glabrous to 3 172 L montium carolumanum Li Wanveaavl AAA Lis ate : — Limontum carolinianum and Limonium limbatum. A — FE, Limonium carolinianum. A. Habit Banioa . inflorescence. C. Calyx, gla ete D. Calyx, pubescent, and floral brace. E. Petal with adnate stamen. FE Fruit (right) and seed (left). G—L, Limonium limbatum. G. Portion of inflorescence. Close-up - flowering spikelets and floral ae [. Floral bracts. J. Flower. K. Longitudinal section ar flower with piscil removed. L. Pistil. (A A drawn from Luteyn 2703 and 28 20, both NY; B from Lateyn er i D and E from Luteyn 2777, —I from Ferris G Duncan 3446, NY; and J—L from Paerens 17938, NY.) S fom = a o) 3 =) K = = z S = No ™N ww wn Bee FIG. 2. States. 173 limbatum Le ° oO 6 L. carolinianum w uo 58 28 = Oo " an 7 % 9 “ ee. » 7 Yo ao @\ aN 2 Y : ~. eS ~ e eae ° ray oO Distribution of Limoninm limbatum and Limoninm carolinianum in the southeastern United 174 densely pubescent along 2 — 3 ribs, the other 2 ribs moderately pubescent, pubescent only at base or glabrous, limb wide-spreading or flaring at maturity, lobes broadly triangular-ovate, 0.5 — 1.2 mm long; petals blue to nearly white. Fruit 2.5 — 3 mm long. (n= unreported) Summer, early fall. Wet meadows, gypsum soils, salt-flats and alkaline depressions in the interior, 1400 — 5800 ft elevation. Figs. | and 2. Incl. Limonium limbatum var. glabrescens Correll. Although L. /imbatum, from the Trans-Pecos and Panhandle regions of Texas, 1s very distinct from L. carolinianum in its inland, higher elevation habitat, morphologically they are quite similar. The type and degree of variation within L. /imbatum is exactly similar to that in L. carolinianum, therefore, no infraspecific taxa are recognized. 2. PLUMBAGO L. Leapwort Perennial shrubs or suffrutescent herbs; stem ribbed, often elongate and climbing. Leaves cauline, alternate, membranaceous. Inflorescence of terminal or axillary, spike-like racemes or panicles. Flowers sometimes het- crostylous, hypogenous, short pedicellate, pedicels bibracteolate; calyx tubular, capitate-glandular along the 5 ribs, lobes triangular, |— 2 mm long; corolla salverform, lobes obovate, rounded or truncate, mucronate; stamens free from the corolla, included or exserted; style 1, included or exserted, with 5 linear stigmas. Fruit a capsule, included, long-beaked, the valves coherent at apex Corolla white, tube mostly less than twice the length of - calyx; calyx with glandular hairs only; inflorescence elongate, 3— 11 (30) cm long; ELE VEL ESEE G1: 10 bcp ae ane Vena a a aR so re a 1. P. scandens Corolla pale blue, tube twice or more than twice the length of the calyx; calyx with glandular and eglandular hairs; inflorescence compact, 2.5 — 3 (5) cm long; plants cultivated and naturalized locally in Florida ....2. P auriculata 1. B scanbens Linnaeus. — Erect, prostrate or climbing suffrutescent herbs; stems glabrous. Leaves ovate, lance-elliptic, spatulate to oblanceo- late, (3) 5 —9 (13) cm long, (1) 2.5 — 3.5 (5) cm wide, apex acute, acumi- nate or obtuse, base attenuate. Inflorescence elongate, leafy, 3-11 (30) cm long, rachis glabrous but glandular and viscid, floral bracts lanceolate, 3 —6.5 mm long. Flowers heterostylous; calyx 7 — 11 (13) mm long, tube glabrous but with glandular hairs along length of ribs; corolla white, a 33. mm long, tube 12.5—28 mm long, lobes 5— 12 mm long, 3 — 3.5 mm wide; stamens included. Capsule 7.5 — 8 mm long. Seed redd- ish- or dark brown, 5—6 mm long. (n=unreported) All year. Palm groves, thickets, shady hammocks, shell mounds, and rocky places in open > areas. Figs. 3 and IG. 3. Plambago scandens. A. Habit. B. Close-up of leaves and leaf bases. C. Portion of inflorescence showing glandular rachis, floral bracts, and flower. D. Distal portion of stamen. (Drawn from Hudson 745, M O.) Plumbago scandens is a widespread, tropical American species which reaches its northern distribution in Florida, southern Texas, and Arizona. It is morphologically quite stable throughout its geographical range. 176 e A MILES fe) | P. scandens P. auriculata FIG. 4. Distribution of Planbavo scandens and Plambago auriculata. = | FIG. 5. mnibrer) wulat _B. Close-up of leat bases. C. Portion spay Sag ha showing eco en floral bracts, ee a in bud. D. Longitudinal section of corolla. E. Distal portion of stamen,. E Distal portion of style. New York Boranical Garden. ) (Original illustration drawn from living materi ul cultivated at the 178 2. P. auricutata Lam. — Perennial shrub, erect, trailing or climbing; stems glabrous below becoming pubescent above. Leaves elliptic, oblanceolate, obovate to spatulate, 2.5-—-9 cm long, 1—2.5 cm wide, apex acute or obtuse, mucronate, base long attenuate or sometimes auricu- late. Inflorescence compact, 2.5 — 3 (5) cm long, rachis short pilose, eglan- dular, floral bracts lanceolate, 4—9 mm long. Flowers tristylous; calyx 10— 13 mm long, tube usually short pilose and also with glandular hairs along upper 1/2 — 3/4 length of ribs; corolla pale blue, 37 —53 mm long, tube 28 — 40 mm long, lobes 10 — 16 mm long, 6— 15 mm wide; stamens included or exserted. Capsule 8 mm long. Seed brown, 7 mm long. (2n= 14, 16, 28) All year. Escaped from cultivation and naturalized in Florida in hammocks, thickets, and disturbed sites in dry soil. Figs. 4 and 3. P. capensis Thunb. ACKNOWLEDGMENTS [am grateful to Bobbi Angell for the beautiful illustrations; to Maria L. Lebron for help with the maps; to Barney Lipscomb and two anonymous reviewers for helpful comments on the manuscript; and to the curators of the following herbaria for loans of their material: AAU, AN : DUKE, E FLAS, FSU, FTG, G, GA, GB, GH, K, LAE MEXU, MO, NCS, NCU, NO, NY, OKLA, PH, RUT, S, ‘SMU, TEX, UC, US, USCH, USE REFERENCES BAKER, H. G. 1953. Dimorphism and monomorphism in the ea Seca IT. Pollen stigmata in the genus Limontum. Ann. Bot. (London) | : 433 —445, LUTEYN, J. L. 1972. A taxonomic study of the genus ae (Plumbaginaceae) in eastern oe America Master's thesis, Duke University, Durham, North Carolina ——____.. 1976. Revision of Limoninm (Plumbaginaceae) in eastern North America. Brittonia 28: ai S17. 1990. Plumbaginaceae. Pp. 37 — 47. In: Harling, G. and L. Andersson (eds.), Flora of Ecuador 39. n press. Plumbaginaceae. In: Massey, J. et al. (eds.), Vascular flora of the southeastern United States. The University of North Carolina Press, Chapel Hill. RADFORD, A. E., C. R. BELL, J. W. HARDIN, and R. L. WILBUR 1967. Con- tributors’ guide for the "Vascular flora of the southeastern United States.” Department of Botany, University of North Carolina, Chapel Hill jes anc TAXONOMIC NOTES ON WESTERN AMERICAN GENTIANACEAE! JAMES S. PRINGLE Royal Botanical Gardens, Box 399 Hamilton, Ontario CANADA LSN 3H8 In the manuscript on the Gentianaceae recently prepared for The Jepson Manual of the flora of California, departures from previous treatments of certain taxa require further explanation than would be feasible within the manual itself. The appropriate discussions, along with a nomenclatural combination that must be validated for use in the manual, are presented HE CIRCUMSCRIPTION OF SWERTIA Wood & Weaver (1982) have called the circumscription of Swertia L. “perhaps the most controversial in the Gentianaceae.” Over the years about a dozen genera have been segregated from Swertia sen. lat. Recent authors have generally treated the Eurasian and African representatives of this complex as one genus, but, with regard to the North American species, the status of Frasera Walt. remains unresolved. Until recently the segregation of Frasera was rejected in most standard floras, following Die natiirlichen Pflanzenfamilien (Gilg 1895) and St. John’s (1941) monograph of Swertia s. lat. in North America. During the past thirty years, however, generic status for Frasera has gradually been gaining acceptance. This has been based in large. part on unpublished studies by _M. Post ca. 1948— 1957, summarized by Hitchcock (1959) and Threadgill & Baskin (1978). Frasera has also been accepted in Wood & Weaver's (1982) recent discussion of generic delimitation in the Genti- anaceae of the southeastern United States. My preliminary discussion of this question in 1979, in contrast, advocated the inclusion of Frasera in Swertia. Basic chromosome numbers have been emphasized in support of generic status for Frasera, although a suite of morphological characters has been presented as being correlated with chromosome number. Wood & Weaver (1982) described Frasera as having x = 13, whereas “the perennial species of Swertia s. str. have numbers mostly based on 14.” Post (paraphrased by ‘Contribution No. 66 from the Royal Botanical Gardens, Hamilton, Ontario, Canada Sipa_ 14(2):179 — 187. 1990. 180 Hitchcock 1959) went so far as to suggest “that Frasera, with a basic chromosome number of 13, is perhaps more closely related to Gentiana (which has a similar number) than to Swertia.” Toyokuni (1965) expanded Frasera to include those Japanese species then known to have x = 10, es- sentially basing this treatment on chromosome numbers alone, but this circumscription has not been adopted by subsequent authors. Wood & Weaver (1982) commented that “this alignment is contradicted by other characters.” Among the Japanese species, for example, S. psendochinensis Hara, with » = 10, 1s especially similar in morphology to S. perennis L., the type species of the generic name Swertia, with n = 14, Unfortunately, chromosome numbers were known for only a few species in this complex until recently. Since the papers cited above were written, chromosome counts have been published for many more species of Swertia s. lat. (compiled 1973 — 1988 in Regnum Veg. 90, 91, 96, Monogr. Syst. Bot. Missouri Bot. Gard. 5, 8, 13, 23, and Taxon 35, 36; see also Table 1). Also, as is often the case, a few early counts have not been substantiated by further investigations of the same species and must be considered unreli- able. With chromosome numbers now known for over 40 species in this complex from all parts of its range, a much clearer picture