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 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 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 of the distribu- tion of chromosome numbers in Swertia s. lat. is now available. This pic- ture 1s considerably at variance with some extrapolations made from early counts. It is now evident that the majority of species in Swertia s. lat. have x = 13 (usually » = 13, occasionally » = 26 or 39). A smaller number of Asiatic species have x = 10 (usually » = 10; » = 30 in S. tashiroi (Maxim.) Makino). Associated with each of these groups are a few aneuploid species having one more or one fewer (in S. paniculata Wallich two fewer, 7 = 8) chromosomes per genome than the prevailing number. Of these, the amphi-Pacific S. perennis L. (including S. cuspidata (Maxim.) Kitigawa, S. obtusa Ledeb., and S$. stenopetala (Regel & Tiling) Pissjauk., segregates seldom accorded specific rank) is the only species known to have v = 14 or any multiple thereof. The generic name Frasera Walt. is typified by S$. caroliniensis (Walt.) Kuntze (as F caroliniensis Walt.), a species of eastern North America with » = 39. Wood & Weaver (1982) followed all of the earlier interpretations of Frasera except Toyokuni’s in restricting this genus to North American species, implicitly assigning to it all species of Swertia s. lat. native to this continent except S. perennis. They characterized Frasera morphologically by: cauline leaf bases weakly connate, not decurrent (free, long-decurrent in Swertu); flowers always tetramerous with one nectary pit per corolla lobe (Swertia usually with pentamerous flowers and two nectary pits per lobe, n 181 some species either with tetramerous flowers or with solitary nectaries); and a distinct filiform style (stigma sessile or nearly so in Swertia). Threadgill & Baskin (1978), following Post, listed axis type, general anatomy, phyllotaxis, nodal anatomy, number of flower parts, and pres- ence of a style as morphological features by which Frasera and Swertia s. str. were differentiated. Swertia bimaculata (Sieb. & Zucc.) C.B. Clarke and S§. swertiopsis Makino, both of Japan, correspond ideally to Wood & Weaver's concept of Swertia s. str. in morphology, being perennial and having striate internodes, pentamerous corollas with paired nectaries, and, in S. bimaculata, a sessile stigma, yet both have x = 13. Swertia bimaculata has the same number, 7 = 13, as the taxon called Frasera pahutensis Reveal (probably best included in S. albicanlis (Griseb.) Kuntze), an ideal “Frasera” with tetramerous corollas and single nectaries. Pentamerous corollas with paired nectaries also occur in species with x = 10, ¢.g. S. japonica (Schult.) Makino and S. pseudochinens1s Hara, but so does the combination of tetramery and single glands, as in S. tashiror. The significance of the number of nectaries per corolla lobe can clearly be discounted. In western North America, S. radiata (Kellogg) Kuntze (FE speciosa Douglas ex Griseb.) most closely resembles S$. caroliniensis in general aspect, life history, stem structure, phyllotaxy, foliar and floral morphology, and chromosome number (both » = 39). It has consistently been placed in Frasera by all who have accepted the genus, but it has paired nectaries. In Asia both single and paired nectaries can be found associated with both tetramerous and pentamerous flowers. Paired nectaries — i.e., the actual groups of nectariferous cells at the base of a pit — may open into a single pit, as in the Asiatic species S. alata (D. Don) C.B. Clarke, S, angustifolia Buch.-Ham. ex D. Don, S. nervosa (G. Don) C.B. Clarke, and S. ciliata (D. Don ex G. Don) B.L. Burtt: or the nectariferous zone may be w-shaped, as in the American species §. neglecta (H.M. Hall) Jeps. Con- versely, many species have solitary nectaries opening into pits with u- shaped or notched openings. Even within a single species or on an in- dividual plant, as in the Asiatic S. atroviolacea H. Sm., there may be various degrees of fusion of the nectary pits. Pentamery is not constant in S. perennis; individual specimens have been found to be variable in this study, with tecramerous flowers being quite common. Occasional tetramerous flowers were also noted on specimens of S. bimaculata and S$. japonica, and have been reported in other species that normally have pentamerous flowers. Stylar differences between S. perennis and “Frasera” are a matter of relative length rather than being qualitative. In this study, distinct albeit slender styles were observed in S. perennts, 182 Taste |. Partial list of Swertia species for which chromosome sources), with floral characters and phyllotaxy numbers have been publi shed (see text for SPECIES RANGE N COROLLA pirs/ PHYLLOTAXY LOBES LOBE Ay ne (D. Don) Asia 13 4 l opposite Clarke Ss, liaise (Griseb. N.Am. 13 4 l opposite intze (S ie as J. a 5S: be Buch. Asia 13 4 | opposite Ham. ex D. Don S. beddomei C.B. Clarke Asia 13 4 l opposite S: peer oe & Asia 13 5) 2 opposite . Clark S: rein teak ex Asia 13 4 2 Opposite Fleming) Karsten S. cordata (G. Don) Asia 13 5 l opposite C.B. Clarke S. corymbosa (Griseb.) Asia 13 4 | opposite C.B. Clark S. densifolta (Griseb.) Asia 13 4 l opposite Kashyapa Ay ee — Don ex Asia 13 4 2 Opposite n) C.B. Clarke S. nervosa (G. Don) Asia 13 4 l opposite C.B. Clarke S. petiwolata Royle ex Asia 13 5 2 opposite D. Don S. speciosa (D. Don) Asia 13 5 2 opposite C.B. Clarke S. thomsonn C.B. Asia 13 5 2 opposite Clarke S. trichotoma (Wight) Asia 13 4 l opposite C.B. Clarke S. swertiopsis Makino Asia 26 5 2 opposite S. caroliniensts N.Am. 39 4 l whorled ale.) Kuntze S. radiata (Kellogg) N.Am. 39 4 2 whorled Kuntze S. perennis L. ; ms 14 > 2 opposite Eurasia or 4 or alt Ss: Hae aes Dor Asia 10 5 l opposite ) BE: oan (. bnpnaan a Dor Clar S. te (Turcz.) Asia 10 5 2 opposite Benth. & Hook. fil. (S. tosaensis Makino) 183 S. japonica (Schult. ) Asia 10 5 2 opposite Makino S. minor (Griseb.) Asia 10 4 2 opposite Knobl. S. psendochinensis Hara Asia 10) 5 2 opposite S. tashiroi (Maxim.) Asia 30 4 l Opposite Makino S. tetragona (Edgew.) Asia 9 5 2 opposite C.B. Clarke S. paniculata Wallich Asia 8 5 l opposite or 4 scarcely exceeded by those of some “Frasera” species. Swertia swertiopsis, which as noted above is otherwise an ideal “Swertza” in morphology, has a relatively long, slender style. Sessile stigmas do occur in Swertia s. lat., but are variously combined with pentamery and tetramery, single and paired nectaries, and x = 10 and 13. Both whorled and opposite leaves occur within North American “Frasera” and within several other species-groups. Connate leaf bases, although prominent in S. carofiniensis and S. radiata, are not present in some opposite-leaved North American species, at least at the upper nodes. Post's (1958) published study of nodal anatomy, specifically the number of gaps in the stele associated with leaf traces at successive nodes, divided the North American species into five groups. The difference in nodal anatomy between his group V, which comprised S. perennis alone, and group IV of “Frasera,” does not appear significantly greater than the difference between groups I and IV of “Frasera.” Its citation by later authors as supporting a generic division was based on a supposed correlation with basic chromo- some numbers and morphological features. No Eurasian or African species were included in Post's study. The striking difference in aspect between the familiar S. carolzmiensis and S. radiata, which have tall, stout, hollow stems, and S. perennis, which is of lower stature with slender, solid stems, cannot be used to support the seg- regation of Frasera from Swertia (unless Frasera were to be restricted to the two stout-stemmed species). Most North American species have slender, solid stems and and narrower leaves, and are of much lower stature than the two hollow-stemmed hexaploids. The basic inflorescence type throughout Swertia s. lat. is a thyrse or verticillaster, comprising a determinate (usually elongate) axis bearing opposite or whorled branches that terminate in dichasial cymules. Within North American “Frasera” there 1s considerable diversity in total inflorescence size, height at which the lowest infloresc- ence branches are produced, and length of internodes in the infloresc- ence — as much within “Frasera” as between “Frasera” and residual Swertia 184 species. Conversely, within residual Swertia there is extreme diversity in general aspect and in inflorescence branching. Many Asiatic species are similar in these respects to some of the slender-stemmed “Frasera” species of North America, including some species with narrow, crowded in- florescences and others with diffuse inflorescences (see Pringle 1979 for examples). The Asiatic and Indonesian taxa also include, however, such highly dissimilar species as §. dichotoma L., with small flowers and slender, decumbent stems giving the plants a chickweed-like aspect (or, as the old name Anagallidium Griseb. for a generic segregate suggests, an Anagall1s- like aspect); S. zeylanica (Griseb.) C.B. Clarke, with a flat-topped inflores- cence; S. acaulis H. Sm., witha greatly reduced inflorescence axis, the long pedicels appearing to arise from the caudex; S. pubescens Franch., with each flower subtended by a pair of large, ovate bracts; and other species respec- tively resembling Gentianella and Deianira species in general aspect. These diverse habits show no correlation with the floral characters discussed above. Nilsson (1967) found that all species of Swertia s. lat. that he studied had basically similar pollen, the grains being separate, prolate, and tricolpo- rate, with the exine striato-reticulate. He did find that all North American species of Swertia s. lat. except S. perennis shared an exine structure unusual for the genus, with finer striations and more closely spaced baculae than those of most other species. Such pollen, however, was also found in one Japanese and two Himalayan species. Of these, S. japonica has n = 10, pentamerous corollas, and paired nectaries; and S. handeliana H. Sm. (chromosome number unknown) combines the “Frasera” characters of tetramerous flowers and single nectaries with the subsessile style attributed to Swertia s. str. (The third species has not been described, at least not under the unpublished name by which it was known to Nilsson.) Among the Asiatic species there was considerable diversity 1n exine structure. Nils- son retained the broad concept of Swertia. Differences in xanthone chemistry have been alleged to support the seg- regation of Frasera, but the early studies (see Threadgill & Baskin 1978 for citations) dealt with too few species to reveal patterns within Swertza s. lat. Later investigations of the oxygen-substitution positions of xanthones in the Gentianaceae by Jossang et al. (1973) included nine species of Swertia s. lat., the North American species being represented by S. a/bicaulis and S. caroliniensis. Species represented by more than one specimen were generally variable in xanthone chemistry. Substitution at position 2 was restricted to S. albicanlis and S. caroliniensis among the species studied, but was not found in all specimens of either of these species. At least one sample of S. caroliniensis was identical in xanthone chemistry to samples of S. perennzs as 185 well as S. chirayita (Roxb. ex Fleming) Karsten (v7 = 10, lobes 4, nectary 1), S. diluta (Turcz.) Benth. & Hook. fil. var. tosaensis (Makino) Hara (S. tosaensis Makino; » = 10, lobes 5, nectaries 2), and S. swertiopsis. Their study, therefore, does not provide support for generic status for Frasera. In summary, data now available show that the morphological characters associated with “Frasera” are restricted neither to North American species nor to species with x = 13, nor are they consistently correlated with one another in their occurrence. Conversely, rather than differing in basic chromosome number, most species of Swertia s. str. and “Frasera” have the same basic number. Although, as indicated particularly by Nilsson’s studies, the North American species of Swertia exclusive of S. perennis may have a monophyletic origin, this group does not appear to have differentia- ted from its Asiatic relatives sufficiently to justify its recognition as a genus or even to permit the characterization of such a genus. No suite of correla- ted characters nor, as far as evidence is available, even one character reliably separates “Frasera” from Swertia. The same criteria by which Wood & Weaver rejected the segregation of Japanese taxa as Frasera species and restored them to Swertia (above) also preclude the segregation of the North American species. Therefore Swertia will be retained in the broad sense in The Jepson Manual. THE STATUS OF SWERTIA UMPQUAENSIS According to Peck & Applegate (1941), their Frasera umpquaensis was much like F fastigiata (Pursh) Heller (Swertia fastigiata Pursh) in general aspect and had “quite similar foveae” (nectary pits), “differing in the setae and in the characteristics of the calyx.” The former reference was evidently to “setae” (trichomes in the present paper) on the corolla in the “area below the [nectary} pit” similar to those surrounding the opening of the pit. Such trichomes would be designated corona trichomes in the terminology now prevalent for floral descriptions in this genus. Peck & Applegate may have assumed that their presence constituted a distinction because Card’s (193 1) “Revision of the genus Frasera” lacked any mention of such trichomes in his description of F. fastigiata or any representation of them in his illustration of corolla details (although the corona trichomes of other species were shown). Peck & Applegate described the calyx lobes of E wmpquaensis as being “linear cto lance-linear, somewhat unequal, 9—12 mm _ long,” whereas Card described those of F fastigiata as “somewhat subulate, 2 cm long.” Actually, S. fastigiata does have a corona of trichomes near the base of the corolla, as is clearly shown in Abrams’ (1951) and Hitchcock’s (1959) illustrations of this species. Hitchcock described the calyx lobes of F fastz- 186 giata as lanceolate, 5— 13 mm long, although as illustrated they could be termed linear. Supposed differences in calyx-lobe shape appear merely to reflect different authors’ interpretations of descriptive terms. The only distinction between S. fastigiata and S. wmpquaensis cited by Abrams was in the apex of the corolla lobes, described in S. wmpquaensis as being narrowed “rather abruptly at the apex to a slender apiculation with l-few minute teeth.” Such characters tend to be variable within species of Swertia, and their aspect may be affected by maturity and by preparation of specimens. An “apiculation” appears in Hitchcock’s illustration of the corolla of F. fastigiata. In the present study, comparison of specimens from California identified as S. or EF umpquaensis with specimens from the Blue Mountains of Oregon and from Idaho identified as S. or F. fastigiata disclosed no differences by which two taxa could be distinguished. A NEW COMBINATION IN GENTIANA Extreme forms of Gentiana newberry: A. Gray s. lat., treated as G. newberryi s. str. and G. tiogana Heller by Mason (1960), are connected by too many intermediates to permit their interpretation as two ordinarily well-differentiated species that occasionally hybridize where their ranges overlap. Intermediate forms outnumber specimens of G. newberryi s. str. and prevail to the virtual exclusion of either extreme in some regions (note distribution of “hybrids” as mapped by Mason 1960). Nevertheless, the relatively tall plants with blue corollas that occur in the Klamath Ranges of California and in Oregon (G. newberry: sensu Mason) do appear to merit some taxonomic distinction from the more widespread “tiogana” plants with white or faintly blue-tinged corollas in the Sierra Nevada of California and adjacent Nevada. The following combination is therefore made: — GENTIANA NEWBERRY! A. Gray var. TIOGANA (Heller) J. Pringle, comb. nov. — BastonyM: Gentiana tiogana Heller, Leafl. W. Bor. 2:221. 1940. Gentiana newberryi var. newberry: in this concept corresponds to G. newberryi sensu Mason (1960). Some plants in the Cascade Ranges and the northernmost Sierra Nevada of California appear to be genuine in- termediates. Most of the “hybrids” and “intermediates” of earlier identi- fications, however, should probably be included in var. tiogana as low- altitude forms. REFERENCES ABRAMS, L. 1951. Illustrated flora of the Pacific States. Vol. II]. Geraniaceae to Scrophu- lariaceae: Geraniums to Figworts. Stanford: Stanford University Press. viii + 866 pp. Mi » H.H. 1931. A revision of the genus Frasera. Ann. Missouri Bot. Gard. 18:245 — 280, pl. 14 187 GILG, E. 1895. Gentianaceae. In: Engler, [H.G.JA., & K.{A.E.]} Prantl. Die oo Pflanzenfamilien... Leipzig: Verlag von Wilhelm Engelmann. 4(2):50— 1 HITCHCOCK, C.L. 1959. genie Gentian family. In: cae CL. A. oe M. Ownbey, & J.W. Thompson. Vascular plants of the Pacific Northwest. Part 4: eas through Campanulaceae. Univ. Wash. Publ. Biol. 17(4):57 — 76, 80. JOSSANG, P.,, J. CARBONNIER, & D. MOLHO. 1973 [“1972”}. Les xanthones de Gentianacées. ies - taxinomie [sic] numerique au niveau moleculaire. Trav. Lab. La Jaysinia 4:143 MASON, C.T., Te: 960 iene in the perennial gentians: G. newherry: and G. tiogana. Madrono 15:233 — NILSSON, S. 1967. He morphological studies in ia Gentianaceae-Gentianinae. Grana Palynol. 7:46— 145. (Incl. appendix by H. Sm PECK, M.E., & E.]. APPLEGATE. 1941. A new Frasera i Oregon. Madrofo 6:12. POST, D.M. 1958. Studies in the Gentianaceae. I. Nodal anatomy of Frasera and Swertia perennis. Bot. Gaz. (Crawfordsville) 120:1— 14. PRINGLE, J.S. 1979. New combinations in Swertia (Gentianaceae). Phytologia Frasera. Amet. Midl. Nat. 26:1— 29. THREADGILL, PE, & J.M. BASKIN. 1978. Swertia caroliniensis or Frasera caroliniensis? Castanea 43:20 — 22. TOYOKUNI, H. 1965. Systema Gentianinarum novissimum. Symbolae Asahikawensis lel47 = 138. WOOD, C.E., Jr, & R.E. WEAVER, Jr. 1982. The genera of Gentianaceae in the sbucherstenn United States. J. Arnold Arbor. 63:441— 487. Book REVIEWS Looman, J. anp K. E Best. 1987. Budd’s flora of the Canadian Prairie Provinces. (Revised). Title changed from “Wild Plants of the Canadian Prairies,” published in 1957 and revised in 1964 and 1969 with input from Budd’s records. 863 pp. 230 figures. Cloth, $38.50 in Canada; $46.20 for other Countries. Agriculture Canada Publication 1662, Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Canada K1A 0S9. A popular manual with keys and diagrammatic illustrations. KANNowskI, Paut B. 1989. Wildflowers of North Dakota. 126 pp. with last 6 blank for Notes. Colored photographs of 161 species. $12.95 paper. University of North Dakota Press, Order from: Wildflowers, Department of Biology, University of North Dakota, Grand Forks, ND 58202-8238. Every state should have wildflower books with colored plates of representatives of the flora of the region and this one ts an excellent, yet inexpensive book. MatHew, Brian. 1989. The genus Lewisia. 171 (151 numbered) pp. with 28 color photographs, 12 botanical watercolors, and numerous maps and drawings. $29.95 plus $3.00 shipping & handling, Hardbound. Order from: Timber Press, Inc., 9999 SW Waulshire, Portland, OR 97225. Tele: 800/327-5680; Fax: 503/292-6607. This is a Kew Magazine Monograph Series published by The Royal Botanic Gardens, Kew in association with Christopher Helm and Timber Press. From the flyer accompanying the book: “This latest in 1 the celebrated series of Kew Monographs treats one of the genera most esteemed by alpine enthusiasts everywhere, even though the ten on only 19 species and its range is confined to western North America. Within the species there 1s much natural variation, and ee and hybridization has ae mine he scope of plants available.” “The book presents a comprehensive taxonomic account of the genus, with much inform- ation on the history, morphology, and relationships of Lewisias. Since it is written for horticulturists as well as boranists, details of cultivation are provided, < well as a useful list of currently available cultivars. C hristabel King’s lovely watercolors a complement Brian Mathew’s authoritative text. Sipa 14(2):188. 1990. A NEW SPECIES OF LIATRIS (ASTERACEAE) FROM THE CAROLINA SANDHILLS' JON M. STUCKY AND MILO PYNE Department of Botany, Box 7612 North Carolina State University, Raleigh, NC 27095-7612, U.S.A. ABSTRACT Liatris regimontis (Small) Schumann has been regarded as a species widely distributed in the western Piedmont and adjacent provinces of ae North Carolina, South Carolina, and Georgia. Principal | t lyses (PCA) show that the holotype and an isotype of L. regimontis from the western Piedmont of NC are close eee approximates of the holotype of L. eraminifolia var. smallit (Britton) Fern. & Griscom from the mountains 0 and that the two types of the former species fall well within the range of morphological variability described by a sample of the latter variety. It is concluded that L. graminifolia var. smallii and the types of L. reg/montis represent the same taxon. PCA and cluster analyses show that specimens from the Fall-line sandhills of North and South Carolina previously determined by other investigators as L. regLINONELS are eaecee: discontinuous with material from other portions of the species range, including the holotype and isotype, and with collections of L. graminifolia var. smallit. This aon sandhills plant is here described as a new species, Liatris coker Pyne & Stucky. In a geographical zone in the Coastal Plain of the Carolinas, where the ranges of L. cokers and L. graminifolia become contiguous, specimens that are morphologically intermediate between the two species have been collec- ted. Liatris regimontis (Small) Schumann ha sido considerada una pe ampliamente dis- tribuida en la = occidental del Piedmont y en provincias proximas de los estados de Virginia, Carolina del Norte, Carolina del Sur, y Georgia. Los aie de componentes principales (PCA) muestran que el holotipo y un isotipo de L. regimontis de la parte occiden- tal del Piedmont en Carolina del Norte son proximas en morfologia al es de L. graminifolia var. smallii (Britt.) Fern. & Griscom de las montanas de Virginia, y que los dos tipos de la primera especie se hallan dentre del rango de variabilidad morfologica circu scrita por un muestréo de la seguna variedad. Se concluye que L. graminifolia var. ne y los tipos de L. regzmontis Bepeesentan el mismo taxon. PCA y analisis de grupos (“cluster analysis”) muestran que especimencs del area Fall-line Gadhills en Carolina del Norte y del Sur, anteriormente identificados como L. ree/montis por otros investigatores, son morfoldgi- camente discontinuos con colectas de otras areas de la distribucion de la especie, incluyendo Y el holotipo ¢€ isotipo, y con colectas de L. halien var. smallii. Esta planta distinta de las colinas arenosas se describe aqui como una espeet , Liatris cokeri Pyne & Stucky. En una zona geografica donde la distribucion de L. ee y hae L. graminifolia son contiguas, especimenes que son morfol6gicamente intermedios entre las dos especies han sido colec- tados 'Paper No. 12: oe of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC 27696-76 Stipa 14(2):189 — 208. 1990. 190 INTRODUCTION In preparation for a taxonomic study of Liatris Series Graminifoliae, herbarium material (GH, NCSC, NCU, NY, USCH) of the five species in the series was examined. A problem with the circumscription of L. regrmontis (Small) Schumann was revealed. Most treatments (Godfrey 1948; Fernald 1950; Cronquist 1952, 1963, 1980) follow Gaiser’s (1946) by recognizing L. regzmontis as a widely distributed species (Atlantic Coastal Plain to western Piedmont, Va to Ga) which inhabits a variety of substrates. In contrast Ahles (in Radford et al. 1968) applies this name to plants only of the Carolina Fall-line sandhills as defined by Duke (1961). The plants from outside the sandhills region included in L. regimontis by the previous workers are included in L. graminifolia by Ahles. Small (1898) based the original concept of Lacinaria regimontis { = Liatris regimontis (Small) Schumann} on his collections from King’s Mountain, Cleveland County, NC. Alexander (in Small 1933) defines the range of this species as “outliers of the Blue Ridge in the Piedmont, also in adjacent provinces.” It 1s, therefore, surprising that the majority of collections to which the name Lvatris regimontis has been applied are plants from the Fall- line sandhills. If the species concept of Gaiser and subsequent workers is followed, then L. regémontis appears to include two distinct morphological types; one which occurs throughout the Fall-line sandhills and another which occurs in the western Piedmont of Virginia and North Carolina and the Piedmont and Coastal Plain of South Carolina and Georgia. If the species concept of Ahles is followed, then a morphologically more homogeneous species results, but this concept excludes the Cleveland County, NC type location from the species range. If, in fact, L. regimontis sensu lato Comprises two morphological variants deserving of recognition, then the one from the sandhills must bear a name other than L. regimontis. Most collections compatible with the type material of Liatris regimontis have previously been determined as L. graminifolia Willd. var. smallii (Britton) Fern. & Griscom. Cronquist (1980) lists this variety in synonymy under L. regimontis. Numerous collections from the Fall-line sandhills have been determined by other investigators as L. regimontis despite their morphological incompatibility with the type material. Since the present study of Liatris regimontis is based largely on specimens determined as L. graminifolia var. smallii, it is necessary to demonstrate that the types of the two taxa are compatible and that these two names apply to the same plant. Accordingly, the purposes of this study are to (1) demonstrate that the names Liatris regimontis and L. graminifolia var. smallii apply to the same plant, (2) determine if material from the Fall-line sandhills is morphologically distinct from material of the Piedmont and 191 Coastal Plain portion of the range of L. regzmontis, and (3) present the most appropriate taxonomic treatment. (Henceforth in this paper, unless other- wise indicated, the element of the Fall-line sandhills will be called the “sandhills plant;” the more widely distributed element of a western Piedmont and Coastal Plain distribution will be called Liatris graminifolia var. smallit). MATERIALS AND METHODS Collections of Liatris regimontis and L. graminifolia var. smallit obtained from G, NCSC, NCU, NY, and USCH were examined. LIATRIS REGIMONTIS - L. GRAMINIFOLIA VAR. SMALLIT COMPARI- son — Data for principal components analysis (PCA) was obtained from the holotype and an isotype specimen of L. regzmontis [NORTH CAROLINA. Cleveland Co.: King’s Mt., 27-30 Aug 1894, J. K. Small s.n. (HoLoTYPE: NY!; tsorype: NY!)}, the holotype of L. graminifolia var. smallii {VirGINIA. Smyth Co.: along Dickey Creek on Iron Mtn., 2900,’ 8 Aug 1892, J. K. Small s.n. (Hovoryee:NY!)}, 22 specimens of Lratris graminifolia var. smallii from western Piedmont sites, 16 specimens of L. graminifolia var. smalli from Coastal Plain sites, and five specimens of L. graminifolia var. graminifolia (Table 1). The last taxon was included to provide outgroup comparison. tates of seventeen characters (Table 2) determined for each specimen constituted data set A. The OTU (specimen) X character matrix was standardized by characters and a character correlation matrix was derived from the standardized matrix. PCA was performed on this correlation ma- trix. LIATRIS REGIMONTIS TYPE SPECIMENS - SANDHILLS PLANT COMPARI- SON — Univariate comparison of the holotype and an isotype of Liatrzs regimontis (see above) and 65 specimens of the sandhills plant was performed. Characters utilized were among those mentioned in Small’s type description of Lacinaria regimontis (1898). LIATRIS GRAMINIFOLIA VAR. SMALLIL - SANDHILLS PLANT COMPARI- sON — A data set was compiled for 25 sandhills plant specimens, 22 L. graminifolia var. smallii specimens from western Piedmont sites, 16 L. graminifolia var. smallii specimens from Coastal Plain sites, the holotype of L. graminifolia var. smallii, the holotype and isotype of L. regzmontis, and 5 specimens of L. aan var. graminifolia included as outgroup representatives (Table 1). The full data set comprising 17 characters, data set B, and a subset of data comprising six characters (Table 2,), data set C, were each subjected to PCA which was performed as described above. Data set C comprised quantitative expressions of those characters included in the 12 univariate comparison. Cluster analyses were performed on data sets B and C. Taxonomic resemblance between OTU’s was measured using the chord distance equation (Pielou 1984) and Gower’s coefficient of similarity (Gower 1971). The resulting distance matrices were subjected to UPGMA clustering (Sneath and Sokal 1973). Results for those analyses utilizing Gower'’s coefficient will be presented as phenograms. Taste 1. Group designation, geographic origin, collection number, and OTU number for specimens included in this study Group Geographic origin Designation* (Co./State Collection No. OTU L. graminifolia var. smallit, WP? Avery/NC Ables & Duke 49602 38 Cleveland/N( Small s 59 Cleveland/NC! Small s.n 18 Gaston/NC Fox 5426 39 Iredell/NC Veerhoff s.n 44 Lincoln/NC Bell 19346 42 McDowell/NC Beaman 04 47 McDowell/ Beaman 210 48 McDowell/NC Began eee 34 McDowell/NC 32 Mecklenberg/NC Ables : ‘Duke 50000 45 Rutherford/NC Fox 5273 37 tokes/N¢ Godfrey & Fox 48575 51 Stokes/N¢ Radford 41403 3] Surry/NC Godfrey & Fox SOI81 35 Transylvania/NC Bannister & Anderson 702 2 Transylvania/NC Cooper 2373 33 Transylvania/ NC Godfrey & Fox 49919 4] Transylvania/ NC Hardin 2222 50 Oconee/SC Powell & Patton s.n. 18 Oconee/SC Radford 1776 49 Pickens/SC Radford 16457 36 Union/S¢ ff 1 46 ork/SC Ables 34488 43 Smythe/VA‘ Small s.n. 60 L. graminifolta var. smatlit, CP! Elbert/GA Cotle 1384 75 art/GA McCarthy s.n 67 Allendale/S¢ etl 522 68 Bamberg/S Ahles 37615 74 Bamberg/SC Ables 37634 72 keley Ables 35525 79 Calhoun/S¢ Ables 353 70 Charleston/S¢ Abhles & Haesloop 38132 80 Colleton/S¢ Rayner 1840 78 Florence/SC Bartlett 2856 69 Hampton/SC Ahles & Bell 18274 73 Tanie 1 (continued) Jasper/SC Bell 5117 Lexington/SC Hutto 199 Orangeburg/SC Abhles 34949 Richland/SC Godfrey 50747 W illiamsburg/SC Radford 3115 Sandhills Cumberland/NC a es G& Leisner 33 f Harnett/NC Fox & Whitford 183 Harnett/NC Rock O61 Hoke/NC Ables 36348 Hoke/NC Ables 30491 Hoke/NC uke R-3289 Hoke/NC rey G Fox 50551 Montgomery/NC Radford 19636 Moore/N Godfrey SOO98 Moore/NC Duke O-3 Moore/NC Wicken s.n. Richmond/NC Preeman 56708 Richmond/NC Radford 19324 Robeson/NC Fox 5508 Scotland/NC Duke 2507 Scotland/NC duke 3240 Wayne/NC Bruton 400 Chesterfield/SC radley & Sears 3505 Chesterfield/SC Duke & Ahles 2200 Darlington/S¢ Coker 5.0 Darlington/S¢ Smut ¢ Dillon/SC Ahles 37096 etna Duke 2313 rshaw/SC Duke Q-2936 ere Duke Q-3110 Intermediate Bladen/NC Ables 37366 laden/NC Crutchfield 5591 Columbus/NC Bell 1583 Columbus/NC Bell 159 Cumberland/NC Ahles 36528 Johnston/NC eee & 2c 48703 Robeson/NC sritr 2983 Wayne/NC ee 28830 Horry/SC Duke 0199 L. graminifolia var. graminifolia See Massey & Massey 2979 Pender/N Ables 36171 cee Ables 340 Warren/NC Bozeman & n Rad 11549 Washington/NC Radford 42 “Group a came at initiation of study. *Western Piedmont. ‘Holotype of L. regemontis (Small) Schumann. eoky ype of L. regzmontis (Small) Schum Holotype of L. graminifolia var. small (Britton) Fern. & Grisc. ‘Coastal Plat 194 Nine herbarium specimens appeared morphologically intermediate (Table 1) and could not be designated with confidence as either Liatris graminifolia var. smallii or the sandhills,plant. Data from these specimens ed to data sets B and C yielded data sets D and E, respectively. PCA was performed on both D and E RESULTS LIATRIS REGIMONTIS - L. GRAMINIFOLIA VAR. SMALLIL COMPARI- SON — The first axis of the PCA explained 24.1% of the data variation. The characters loading heavily on this axis pertained to head and flower size and head density along the inflorescence axis (Table 3). The second axis explained 16.0% of the data variation and was interpreted primarily as a phyllary shape axis (Table 3). Although somewhat distinguished by the second PCA axis, the PCA scores of the type specimens (OTU’s 18, 59, 60) were relatively compatible (Fig. 1). In relation to the total array of PCA scores, the scores for the types were not centrally located; however, they were clearly not disparate. Although there was not a discernable discon- tinuity between PCA scores for western Piedmont and Coastal Plain Taste 2. Characters and character states used in the multivariate study. Pedicel length (mm) Number/3 cm inflorescence axis Orientation: 1, strongly divergent; 2, weakly divergent 3, strongly & PEDICEL: HEADS: NOR Wo INVOLUCRE 4. Height (mm 5 PHYLLARIES: 6. Outer phyllary planation: |, flat; 2, cupped; 3, keeled* Te Inner phyllary length (mm 8 Inner phyllary width (mm) oe Inner phyllary shape index: [length (mm) - distance from apex to point of greatest width]/ length (mm)* 10. Inner a apex shape: 1, truncate; 2, obtuse; 3, acute; 4, acuminate 1. Inner p ae apex reflexion: : none; ee weak; 5 strong 12. Inner ey ae apex planation: 1, flat; 2, involut 1 Extent of scarious margin on inner ee , a 2/3; 2, basal 2/3 an not around apex; 3, complet FLOWERS: 14, Number/head? 15. Corolla tube length (mm) 16. Pappus length (mm) PUBESCENCE: 17. ensity on petioles, inflorescence bracts, and phyllaries: (Density was assessed on each part and the three assessments summed. ): Character states for individual parts were 0, glabrous; 1, sparse; 2, moderate; 3, dense “Included in data sets C and E. Taste 3. Character loadings with absolute values greater than 0.5 for the first principal componen t axes. Data PCA Set Axis Character Loading A I no. heads/3 cm 0.670 involucre height -0.736 involucre width -0.738 phylary length -0.820 corolla length -0.805 pappus length -0.811 I inner phyllary apex shape 0.754 extent scarious margin on phyllary -0.604 B I no. heads/3 cm -0.761 involucre width 0.690 inner phyllary apex shape -0.691 inner phyllary reflexion -0.519 no flowers/head 0.788 outer phyllary planation -0.722 inner phyllary planation -0.749 I Involucre height 0.683 hyllary length 0.868 corolla length 0.767 pappus length 0.797 C I no. heads/3 cm -0.782 involucre width 0.742 inner phyllary apex shape -0.626 no. flowers/head 0.843 outer phyllary planation -0.745 inner phyllary planation -0.779 II involucre width 0.506 inner phyllary apex shape 0.662 specimens, material of these geographical ranges constituted two phases of the distribution of OTU’s in two-dimensional space. PCA scores for the five outgroup OTU’s were discontinuous with the body of scores for the 41 other OTU’s. LIATRIS REGIMONTIS TYPE SPECIMENS - SANDHILLS PLANT COMPARI- sON — The univariate comparison of the type specimens of Liatris regimontis with specimens of the sandhills plant suggested a morphological distinction between the two (Table 4). LIATRIS GRAMINIFOLIA VAR. SMALLI - SANDHILLS PLANT COMPARI- SON — The first axis of the PCA performed on data set B explained 26.0% of the data variation. Characters loading heavily on this axis pertained to head size and density in the inflorescence and phyllary shape (Table 3). The 196 0.675 A 0.4 1 +3 a 4 J a 5 - 0.24 7 “a4 ; o ® ; 60 | 4a a 4 . © 0.07 ‘. aa as 4 4 SMALLII WP Se] q a i a a 4% = aa 4 SMALLII CP —-0.? 4 = ] : x GRAMINIFOLIA -0.44 x = x -0.6 x x -0.8 = 0.0 1.0 1 24.1% FIG. CA of ee set A showing PCA scores of OTU’s on axes [and I]. OTU 18 = isotype of Liatris 9=holotype of L. i sunilae baa = aaa of L. graminifolia var. smallit. regimontss a, K. ) Fer es ae Western Piedmont; CP = Coastal Plain 0.6 x a A aa tak “We = SANDHILLS 7 77 a 4 SMALLII WP ze fs a ® om 8 Ba aah Oy 4 SMALLII CP A480 a a woe aX x GRAMINIFOLIA 4 A a A 4 —— ae 0.0 0.2 0.4 0.6 I 26.0% FIG. 2. PCA of data set B showing PCA scores of OTU’s on axes I and II. Numbered OTU’s are interpr fered as see between the sandhills plant and Liatris cen var. smallt1. OTU's are identified in Table 1. WP= Western Piedmont; CP = Coastal Pla a second axis explained 15.6% of the data variation and was interpreted primarily as a head and flower length axis (Table 3). A discontinuity between the PCA scores for the sandhills plant and those for Liatris gram- inifolta vat. smallii and the types of L. regimontis was evident along the first axis (Fig. 2). This discontinuity was greater than that between L. &ram- intfolia var. smallit and the outgroup OTU’s. This discontinuity was broached somewhat by two disparate specimens of the sandhills plant from 197 Taste 4. Comparison of the holotype and isotype of Liatris regimontis with the sandhills plant. tatrts Sandhills Character regimontis plant Inflorescence heads frequently widely heads closely spaced spaced along inflorescence along inflorescence axis; axis; not secund frequently secund Involucre shape obconic narrowly obconic Inner phyllary apex acute, not involute acute to acuminate, involute Outer phyllaries cupped strongly cupped to keeled Flowers/head 9—12 4—9 (10) Robeson (OTU 10) and Hoke (6) counties, NC, and three specimens of L. graminifolia var. smallit from Charleston (80), Florence (69), and William- sburg (77) counties, SC. The first axis of the PCA performed on data set C explained 57.1% of the data variation. Characters loading heavily on this axis pertained to head size and density and phyllary shape (Table 3). The second axis explained 15.6% of the variation and was interpreted as a phyllary shape and head size axis (Table 3). The discontinuity between the sandhills plant and Liatris graminifolia var. smalli plus the types of L. regimontis along axis one was approximately equal to that between the latter taxon and the outgroup OTU’s (Fig. 3). The sandhills specimen from Robeson (OTU 10) County, NC, and the specimens of L. graminifolia var. smallii from Charleston (80), Florence (69), and Williamsburg (77) counties, SC, were, again, in- termediate. The cluster analysis performed on data set B indicated two major clusters; one composed of 24 sandhills plant OTU’s and the other composed of 39 Liatris graminifolia var. smallii OYU’s from both Piedmont and Coas- tal Plain sites including the holotype, the two type specimens of L. regimontis, the five outgroup OTU'’s, and one sandhills plant OTU (Fig. 4). The cluster analysis on data set C also indicated two major clusters; one comprised entirely of L. graminifolia var. smalli and outgroup OTU’s and the other comprised of 25 sandhill plant OTU’s plus three OTU’s of L. graminifolia var. smallii from Coastal Plains sites (Fig. 5). Cluster analyses that utilized chord distances agreed closely with those presented here; the primary differences being the distances at which OTU’s clustered with each ef. Of the nine specimens that initially appeared morphologically in- termediate, specimens from Bladen (OTU’s 23, 25) and Columbus (22) 198 0.3 10 . ao re 0.2 & rY a A 2 4 ry 0.14 sas : is = SANDHILLS a be a, 4 eo at Ae on 4 SMALLIL WP . A («0.0 " :* S aa Pe 4 SMALLII CP — A aa "48 : x GRAMINIFOLIA -—- -01- "os a . a A . Ne a 4 -0.27 4 x x x -0.3 T r r T T T T y — T r -0.6 -0.4 -0.2 0.0 0.2 0.4 l 57.1% FIG. 3. PCA of data set C showing PCA scores of OTU's on axes I and I]. Numbered OTU'’s are interpreted as intermediate between the sandhills plane and Liatris eee var. smallit. OTU’s are identified in Table 1. WP = Western Piedmont; CP = Coastal P counties, NC, and Horry (24) County, SC, were shown to be intermediate by PCA (Figs. 6 and 7). The specimens from Robeson (26) and Cumber- land (20) County, NC, could, possibly, also be interpreted as intermediate. PCA indicated that the specimens from Johnston (11) and Wayne (19) counties, NC, were compatible with L. graminifolia var. smallit. The spect- men from Williamsburg (77) County, SC, not initially felt to be in- termediate and initially annotated as L. graminifolia var. smallii, was also intermediate according to PCA. Additional initially annotated specimens that could, possibly, be interpreted as intermediate include those from Florence (69) and Charleston (80) counties, SC. DISCUSSION LIATRIS REGIMONTIS - L. GRAMINIFOLIA VAR. SMALLII COMPARISON — The PCA showed that the type specimens of Liatris regimontis were reason- ably congruent with the holotype of L. graminifolia var. smallii and that all three types were included within the range of variability collectively ex- hibited by the 38 other specimens of L. graminifolia var. smallit. These results suggested that L. graminifolia var. smalli and L. regimontis refer to the same plants. The use of specimens determined as L. graminifolia vat. smallii in this study of the circumscription of L. regimontis was justified. Although the Coastal Plain collections of Liatris graminifolia var. smalli appeared to be somewhat differentiated from the western Piedmont collec- tions, these two aspects formed one continuum of variation. We recom- mend that these two regional elements not be taxonomically distinguished 1? SCP 79 $1 34 SCP 69 35 SCP ?4 SCP 78 G 61 G 64 G 65 G 63 6 62 | | | FIG. 4. Phenogram of cluster analysis of data sec B. SWP=var. sma/lii of Western sage SCP = var. smallii . Coastal Plain; S=sandhills plant; G=var. graminifolia (outgroup). OTU’s identified in Table SCP FIG. SCP = var. smallir of saa Plain; S=sandhills plant; G=var. graminifolia (outgroup). =" ec C1 A Oo Ww eSeunan pe a os] r=) — =a ds 7 On C) ad CAN NN AA wWwAAOMMAMWH OHM MH OMMOMOMYM MM SaBDAmoanmrnvenanawWhv hUONN Oo =—_ OT OF SCP 69 Phenogram of cluster analysis of data set C. SWP= identified in Table > | var. smallit of Western ss OTU’'s a 0.6 J 25 0.44 oe 4 id a a gg . we (02 se 2 ws = SANDHILLS r 23 m ae : 4 8 4g = : 2 INTERMEDIATE 1 77. 7 | Ld nl 0.0- x By 22° So 4 SMALLII WP AY as = Bs va 20 . x a ar Ves 80-g8- =o 4 SMALLII CP -0.24 xA 4% 4 a. 0 a. x GRAMINIFOLIA —-0.4 a -0. T T de i -06 -0.4 -0.2 0.0 0.2 0.4 0.6 I 23.8% 6. PCA Ofdareet showing PCA scores of OTU’s on axes I and II. OTU's are identified in Table OTU'’s initially determined as inte a ite between Liairis graminifolta var. smallit = Coastal Platr irermediate ee a sandhills tne WP = Wes tern Piedmont; C ae : , a 0.24 4 =| aa 014+ * a“ ‘: 23 Ba?* om "t Be ; : a a " = SANDHILLS a” 4 4 77 «a wT o 4 a 22 Fr ir ~ (0-0 : ‘. 4 80 a ® INTERMEDIATE a 4 me A 4 -o.14 * a4 et ee 4 SMALLIT WP a A A = | a a Bf 4 SMALLII CP 190 -0.2 ae e x GRAMINIFOLIA x -0.3 8 -0.4 ¥ T i T -0.4 -0.2 0.0 0.2 0.4 | 52.9% FIG. 7. PCA of data set E showing PCA scores of OTU's on axes Land I. OTU’s are identified in Table L. Intermediate = and the sandhills plant; WP = Western Piedmont, CP = Co ITU's initially determined as intermediate between Liatris graminifolta var. smallii vastal Plain. at this time and that future study of the relationships between the two is needed. Additionally, study of the distinction between L. graminifolta var. smalli and L. graminifolia var. graminifolia is warranted to determine if the former would most appropriately be recognized as a variety or as a species, L. regimontis (Small) Schumann. Liatris regimontis TYPE SPECIMENS - SANDHILLS PLANT COMPARI- SON — Each character suggested a morphological discontinuity between 202 the types of Liatris regimontis and the sandhills plant. Distinguishing the different involucre shapes and the cupped vs. keeled nature of the outer phyllaries exhibited by the two groups was strongly subjective. The characters that most objectively distinguished the two groups were the spacing of heads along the inflorescence axis, involute vs. non-involute na- ture of inner phyllary apices (Figs. 8 and 9) and number of flowers/head. LIATRIS GRAMINIFOLIA VAR. SMALLIL - SANDHILLS PLANT COMPARI- SON — The distinction between Liatris graminifolia vat. smallii and the sandhills plant was equal to or greater than that between L. graminifolia var. smallii and the outgroup OTU’s representing L. graminifolia var. gram- inifolia, according to the two PCA’. This distinction was also indicated by 04 Oran Rh 35 6 Sa oeeenpe cB 08ar%o ? o° Ocean 9 x 000 B2s0 RATT ees E5888 Op seo (J eo8 B e 2 FIG. 8. An outer (A) and an inner (B) phyllary of the sandhills plant. Bar = 1mm. 203 the two cluster analyses. According to PCA, specimens from a continuous north-south geographical zone from Cumberland County, NC, to Willi- amsburg County, SC, were morphologically intermediate between L. graminifolia vat. “mall and the sandhills plant (Fig. 10). All specimens which initially appeared intermediate prior to the analyses were include in the final PCA while only a sampling of those specimens that appeared typical for the sandhills plant and L. graminifolia var. smallii were included. In view of this “heavy sampling” of potential intermediates, it is our opinion that the relatively few OTU’s that were shown by the numerical analyses to be truly intermediate do not obviate the overall discontinuity between the sandhills plant and L. graminifolia var. smallii. Both PCA and cluster analyses suggested that the affinity of the sandhills plant is stronger with the Coastal Plain aspect of Liatris gram- inifolia var. smallii than with the western Piedmont aspect. If Gaiser (1946) was correct in suggesting that the widely distributed, morphologically variable L. graminifolia is the evolutionary ancestor to the other geographi- cally more restricted, less variable taxa in series Graminifoliae, the results of the current study suggested that the sandhills taxon evolved from ances- ANNOTATED BY R. K. GODFREY ; a fortx (Welt: ) While, pan ih cd PP 7 ee 9 (9Ee. FIG.9. Series of phyllaries, outer (shortest) to inner (longest), of Liatris regimontis. This is the drawing that is on the holotype of L. regimontis (Small) Schumann 204 Gaieeeye eee & ae Q y Breer ee aS Ge SS 4 Liatris graminifolia var. smallii e Liatris cokeri aa Olntermediate 10 FIG. 10. Distributions of Ligtris cokert, L. gramintfolia var. smalliz, and intermediates between the two taxa. tral L. graminifolia populations of the Coastal Plain. The current study does not provide an adequate basis for determining if the more likely ancestor is L. graminifolia var. smallit or L. graminifolia vat. graminifolia, the latter variety common in the Coastal Plain of North Carolina, because so few specimens of the latter variety were included in the analyses. Investigations of the relationships of the sandhills plant with the two varieties are warran- ted. The intermediate specimens could indicate a zone of primary in- tergradation between the sandhills plant and its ancestral Coastal Plain populations (Fig. 10) or they could indicate hybridization between dif- ferentiated populations. Our results show that the sandhills plant should be recognized as a 205 species since it is morphologically distinct from plants representative of a species concept in which it has previously been included. We are not the first to recommend its taxonomic recognition. The label of an R. K. God- frey collection of the sandhills plant (Godfrey 50098, Moore Co.: NC, 15 Sep 1949) states, “This is considered by Gaiser to be L. regimontis (Small) Schumann, but is considered by the collector to be an entirely different taxon.” Godfrey annotated this and other specimens (NCSC) as Liatris carinata (Small) Coker. Laciniaria carinata Small (1903), the basionym of Liatris carinata (Small) Coker, has been lectotypified to a specimen of Liatris secunda Ell. (1822); therefore, Liatris carinata (Small) Coker 1s a syn- onym for L. secunda Ell. and is not available for the sandhills taxon (Pyne and Stucky 1990). A name for the sandhills species must be published NEW SPECIES OF LIATRIS 1. Liarris coKeri Pyne & Stucky, sp. nov. pecies nova similiter L. reesmontis (Small) Schumann optimo distinguitur a capitulis eau apices phyllaris intimis s involutis, floribus Beanee per capitulo. Species nova similiter L. secanda Ell. © optimo distinguitur a caule glabrate, phyllariis carinatis, cae ad reflexis, involucro at brevior et corolla et pappo multo brevior. Perennial herb; rootstock corm-like, globose, 0.8 — 3.0 cm wide. Stems 1—5 per corm, usually unbranched, frequently drooping or upright, glabrous, usually sparsely minutely glandular, 25—85 cm tall. Leaves linear, densely punctate on both surfaces, occasionally sparsely hirsute along midvesn on either or both surfaces, margins irregularly ciliate near base or occasionally glabrous, (1.8) 2.0 — 4.8 (5.0) mm wide X 0.5 — (2.0) dm long, length gradually reduced upwards. Inflorescence a spike or compact raceme; heads imbricate along rachis, frequently secund, sessile or on bracteate peduncles to 6.0 mm long, closely ascending or diverging particularly when heads secund. Involucres narrowly obconic, 4.8 — 10.5 mm long X 4.0—7.8 (8.0) mm wide at tips of phyllaries during anthesis; phyllaries imbricate in several series, punctate, scarious-margined, usually minutely ciliate or occasionally glabrous; inner phyllaries strongly acute to acuminate, apically involute and spreading to reflexed, 5.0 —8.8 (9.0) mm long; outer ahaa acute, strongly cupped to keeled. Flowers 4—9 (10) per head, corolla tube pink, glandular autsiGe, pilose basally inside, 4.2—7.0(7.5) mm long. Pappus barbellate, 4.0 — 7.0 mm long. Mature achenes obconic, 2.7 — 3.8 mm long, 0.8 — 1.2 mm wide at apex, angular in cross section, longitudinally ribbed, densely hirsute with ascending trichomes, gray to black. Type: NORTH CAROLINA. Harnett Co.: 0.2 mi E jet. NC rt. 27 and co. rt. 1242 along NC 27 on S side road; sandy roadside and margin of longleaf pine/turkey oak/ _ 206 wiregrass vegetation; 23 Sep 1989, J.M. Stucky 511 (HOLOTYPE: NCU; isorypes; GH NCSC, NCU, NY, US, USCH). This species is named in recognition of Dr. W. C. Coker who contributed significantly to the botany of the Carolinas and who included this species, calling it Liatris carinata (Small) Coker, in The Plant Life of Hartsville, §.C. (1912). Although the resolution of a lectotypification problem makes his combination incorrect for this species (Pyne and Stucky 1990), Dr. Coker should be recognized. As far as can be determined, the only vascular plant presently bearing the epithet cokers is Lycopus coker? Ahles. As stems of Liatris cokeri grow longer and as heads mature and become heavier, the degree of drooping of the stems usually increases. On these drooping stems, the heads respond phototropically, causing the secund na- ture of the inflorescence. Due to its phenological basis, the secund inflores- cence becomes more prevalent as the growing season progresses. The non- secund nature of an inflorescence should carry little diagnostic signific- ance, particularly for specimens collected early in the growing season. Liatris cokert and L. secunda Ell. frequently form mixed populations in the Fall-line sandhills of the Carolinas and thus the species have often been confused. The basis of this confusion undoubtedly is their shared habitat and the secund inflorescence. Several characters do, however, distinguish L. cokeri from L. secunda in this area (Table 5). KEY TO SPECIES OF LIATRIS OF THE CAROLINA PALL-LINE SANDHILLS AND ADJACENT OUTER CoasTaL PLAIN _~ aie ee seg errata see oe sig rey ayy ee ee oer ean L. squarrosa ea barbe 2 _ fiddle ie outer oie squarrose; heads tending to be turned away from the axis, not sec Tasie 3. Distinctions between Liatris cokert and L. secunda. Character Liatris cokeri Liatrits secunda Stem pubescence Lacking Usually densely, minutely, I & ¥ y NA | hirsute basally Involucre length Phyllary Sek and reflexior Corolla tube lengel 7 Inner corolla cube pubescenc Corolla lobe length 4.8—10.5 mm Outer frequently distinct . Heads pedicellate; basal leaves 3: Smit Wide length 15. Calyx lobe width Taste 2. Cumulative variance accounted for by the first eight principal components. Principal component 1 Principal component 8 ).371551 0. 86 2849 Taste 3. The first three principal components (eigenvectors) and the amount of variance in each character. Eigenvectors 2 3 Characters 1 0.198363 0.503841 0.048926 2 0.347826 0.301759 0. 148375 a) 0.384297 0.025179 0.014257 4 0.336087 0.212749 0.134065 5 0.360205 0.220944 0.079408 6 0.216563 0.001895 0.188342 7 0.261845 ).074394 0.0437 14 0.227593 0.170500 0.178999 9 0.317034 ).118163 0.140525 10 0. 188620 0.249411 0.038666 11 0.196223 0.044555 0.176109 12 0.110797 0.173056 0.621414 13 0.005623 0.426972 0.506604 14 0.215398 . 284063 0.348118 15 0.220742 -393276 0.255808 236 width, length of blade co its widest point, length of blade toothed and the number of leaf-pairs per cm per branch (see Table 1 and Table 3). In the second principal component, the presence/absence of adventitious roots and the length of the central inflorescence axis are most important. Three more or less distinct groups can be recognized in the ordination produced by principal components analysis (Fig 1): one corresponds to Paxistima canbyi, the taxon endemic to areas of the central Appalachian Mountains and its foothills; another corresponds to P. myrsinites, a species widespread throughout the Rocky Mountains; a third is circumscribed by specimens, not previously studied together, collected in mountainous areas of northeastern Mexico. The range of these Mexican specimens 1s not con- tiguous with the range of the Rocky Mountain taxon. Although geographically disjunct, there is, however, some intergradation in morphology, and consequently overlap in the ordination, between specimens of P. myrsinites collected in the United States and the Mexican specimens. Therefore, we are designating the Mexican populations as a subspecies of P. myrsinites (following the concept of Du Rietz 1930), rather than recognizing them as distinct species. The Mexican populations con- stitute a significant geographic facies of P. myrsinites, and consequently subspecies rather than varietal rank seems appropriate (see Du Rietz). It 1s interesting that a (lesser) tendency toward intergradation also occurs between the Mexican populations and P. canby/; possible interpretations of this observation will be discussed under Distribution and Geofloristic His- tory. The FASTCLUS program of SAS provided further insight into group structure within the genus Paxistima. FASTCLUS ts a disjoint clustering (but non-tree producing) procedure which employs nearest centroid sort- ing, i.e. cluster seeding, techniques (Anderberg 1973); preassignment of number of groups is requisite to the procedure. We ran this procedure for one, two, three and six groups respectively — constituting all putative divisions previously recognized within Paxistima. The — principal components analysis demonstrated that no more than two taxa, 1.e. F canbyi and P. myrsinites, are clearly distinct at the species level, although three groupings may be discerned from the analysis. When the principal components were entered into FASTCLUS, the most favorable clustering score (cubic clustering criterion value), indicative of the optimum number of clusters, suggested the existence of three groups as well. Hence, results of the FASTCLUS procedure support the recognition of two subspecies (myrsinites and mexicana) within PB. myrsinites, as well as the existence of P. canbyi. Our delimitation of three taxa of Paxistima — P. canby1, P. myrsinites subspecies myrsinites and P. myrsinites subspecies mexicana — is thus = = : = = = = = = = a x = = = = = x = x = = = x = = = = = a2 = 2 x 2 = x 3+ x . 2.5 x = = x = = > as v 2 3 eS : = z = ~ ‘. = x = == =z x = z 7 = = = U »< = = U == U VU = x Uu =< U Pad = U U ~< »< ~< U ~< OU U “< a4 U »< Uu »«< U u U ood «KU Uu ~< Ww ~< U o) U c PNaN geo velo FIG. |. Bivariate plot of first rpholovi P. Myrsimites subsp. myrsznites, X = Pp rapes A MEXICANA, io = P canbyi PRINCIPAL COMPONENT 1 ilanalysis of Paxiitima. M = ss 238 substantiated by the numerical phenetic analyses performed, i.e., when the results of both PRINCOMP and FASTCLUS are considered in consort. If one examines the numerical data, the numerical analysis, the keys to taxa, and the descriptions, it will be apparent that all three taxa of Paxis- tima differ only by a number of seemingly minor characters, with overlap- ping character states. Although obviously debatable, if taken collectively, and considered in context of the disjunct nature of major super-groups of populations of Paxzstima, we believe that the data (as analysed by compu- ter) support the recognition (or continued recognition) of three taxa, as opposed to the submergence of all taxa into a single, fragmented, polymor- phic species. Although the taxa of Paxistima are what we would term “statistical taxa,” not distinguished by any one or a few infallible, totally clear-cut characters, the taxa are nonetheless rather readily recognized by their overall patterns when viewed on herbarium sheets, or in the field as we have seen them. As alluded to in the concluding section on Distribution and Geofloristic History, the taxa of Paxistima may well represent the now disjunct and somewhat divergent descendants of a single, wide-ranging, polymorphic ancestral species of the North American Arcto-Tertiary flora. Should all taxa survive, we would predict only a greater divergence of taxa through time, given their present geographic isolation and scant oppor- tunity for gene exchange. SYSTEMATIC TREATMENT THe Genus PANISTIMA PAXISTIMA Raf., Sylva Telluriana 42. 1838. (spelled Pachistima by Rafin- esque, 1818, 1819a and b, a nomen nudum; Pachystima by Endlicher, 1841; and Pachystigma by Meisner, 1843). — Type: Paxistima myrsinites (Pursh) Rafinesque. Oreophila Nutt. ex Torrey & Gray, A Flora of North America 1:258. 1838 (Celastraceae); non Oreophila D. Don, Trans. Linn. soc. of London 16:178. 1833 (Compositae). Oreophila T.& G. is thus a later homonym. Low, evergreen, glabrous, much branched shrub or subshrub with subterranean rhizomes; adventitious roots often present on lower portions of stems; branches terete, with rough bark. Leaves simple, smooth, serru- late to crenulate (rarely subentire), coriaceous, opposite (decussate), short- petioled, with small caducous stipules. Flowers small, perfect, axillary, solitary or in simple dichasia (rarely fascicled or in compound dichasia); calyx lobes 4, imbricate, green, widely ovate, small; petals 4, maroon (occasionally green), trullate, longer than calyx lobes; stamens 4, inserted in the edge of a broad nectar disc, the anthers introrse, the filaments short, awl-shaped (occasionally longer and thread-like); ovary 2-loculed, superior 252. but sunken in the disc; style short to obsolete; stigma capitate to linear- clavate (rarely obscurely 2-lobed). Fruit an oblong, 2-loculed capsule. Seeds 1 or 2, oblong, erect, enclosed in a membranaceous, white, cleft aril, endosperm fleshy. Flowers and fruits developing from early spring to early summer; flower buds formed the preceding summer, although some un- dergo anthesis prematurely (later in the season in which they are formed). ORTHOGRAPHY AND NOMENCLATURE: As noted by Wheeler (1943) and Uttal (1986), the spelling of the genus name should be Paxestzma. Rafinesque provided no description in his early publications (1818; 1819a,b) when he spelled the name “Pachistima.” His references variously to Pursh’s and Nuttall’s descriptions in these publications might appear to achieve validation by direct reference, but do not because neither Pursh nor Nuttall were attempting to describe new genera or sections of genera in this particular case (cf. Article 41.2, International Code of Botanical Nomen- clature, Greutcer et al. 1988). The first generic description or diagnosis legally attachable to the genus occurred in 1838 (7n situ in Silva Telluriana) when Rafinesque employed the spelling Paxistima. It is plausible that Rafinesque (1838) may have written the Greek “chi” or “x” for the “ch” in Pachistima leading to an accidental change to the “x” (Paxistima) spelling; but this is only speculation and not justification for a change back to the “ch” spelling, although Merrill (1949) indicated “Pachistima” to be “universally accepted.” Regarding meaning and gender, Paxistima may be a corruption of pachys (thick) and stzgma (Genaust 1976). Since stigma is neuter, Paxistima could as well be interpreted as neuter. However, this again is difficult to prove, and consequently we are following Wheeler's (1943) apparent recognition of Paxistima as feminine. Concerning the name of the original species, Pursh’s (18 14) inclusion of a question mark in I/ex(?) myrsinites does not invalidate the publication of the epithet myrsinites. Although Wheeler's (1943) interpretation of J. myrsinites aS a provisional name may have been reasonable at the time, according to the present edition of the code the use of a question mark does not obviate publication when the author (Pursh) accepted the species, but merely expressed taxonomic doubt as to which genus it belonged (cf. Art- icle 34.2, International Code). The valid combination Paxistima myrsinttes was made by Rafinesque in 1838. The correct name and citation of the original species is thus Paxzstima myrsinites (Pursh) Rafinesque (1838), not Paxistima myrtifolia (Nutt.) Wheeler (1943). SPECIES AND SUBSPECIES OF PANISTIMA A. Shrub or subshrub 20 to 100 cm high (typically not prostrate); leaves usual- ly 1 —2 pairs per cm of branch length; inflorescences averaging 6 — 10 per branch; western U.S., southwestern Canada, northeastern Mexico... 1) P. myrsinites 240 B. Shrub or subshrub 30 — 100 cm high; leaves | —2 pairs per cm of branch length; blades lanceolate to obovate or oblanceolate, typical- ly 11—27 mm long; inflorescences averaging 10 per branch; western United States, southwestern Canada ................ sity GR PE ees Diatare ou a utee Dene ane ees Ee ees lA. P We subsp. myrsinites — ee. . Shrub or subshrub 20 — 45 cm high; leaves 2(3 — 4) pairs per cm o branch length; blades lanceolate, typically 8— 12 mm ee in- florescences averaging 6 per branch; northeastern Mexico ...... eae ere eo a eats ols Sees ane fea ae 4 eae 1B. P. myrsinites subsp. mexicana A. peuer a (tending to be prostrate) 10-40 cm eh. leaves 2 — 4 pairs per 1 of branch length; inflorescences averaging 4 per branch; eastern United ee Seer GOH a AGH ho Gg grie-a eet tom ese Reeds Basen Sones Ve ae tants 2) P. canbyi — . PAXISTIMA MYRSINITES (Pursh) Raf., Sylva Telluriana 42. 1838. Shrub or subshrub, usually densely branched, 20 to 100 cm high; the lower portion of the stems sometimes prostrate; adventitious roots may be present. Leaves approximate, | — 2 (occasionally 3 — 4) pairs per cm; bla- des ovate (elliptic) to lanceolate (oblanceolate), (6-)8 — 27(-40) mm long, (3-)4 — LO(- 15) mm wide; blade margins serrulate to crenulate (occasional- ly entire), revolute to subrevolute or not revolute (sometimes thickened when not revolute); teeth pointed or rounded, extending from apex to 1/3 to 4/5 of blade length; blade secondary veins indistinct below (occasionally evident); blade apex obtuse, apical angle 90° — 165°; petioles (0.8-)1 — 2(- 2.5) mm long. Inflorescence axillary or terminal, averaging 9.3(3 — 21) per branch, generally composed of | — 2(-3) flowers each; length of central or only inflorescence axis 2— 4 mm. Calyx lobes widely depressed-ovate to very widely ovate, slightly imbricate. Petals exceeding the calyx lobes, commonly maroon (those from buds from preceding season), occasionally green (those from buds from current season). Fruits 4-7 mm long. Two subspecies, Paxistima myrsinites subsp. myrsinites and P. myrsinites subsp. mexicana, are recognized within this species. The typification of P myrsinites 1s discussed under the subspecies P- myrsinites subsp. myrsinites. Paxistima myrsinites subsp. mexicana is described as new 1A. PAXISTIMA MYRSINITES eae Raf. subsp. MYRSINITES. Ilex? ee Pursh, Fl. Amer. Sept., I. 119. 1814. — Lecroryee: Lewis 5.n., 1806 hotograph!; see eae ep Ww). age ror Nutc., .N. Amer. Pl. 109. 1818. — Type: same as lex? myrsi- he s ie Changed to 5 myrtifoli by Nuttall, and hence the epithet ne is a superfluous n Mypinds myrtifolla var. “ sha’ minor Hooker, Fl. Bor.-Amer. 120-121. 1840. — Type: Apparently considered by Hoaker to pre to. original material of [/ex? myrsinites Pursh. i) — U 241 Myginda SS var. “beta” major Hooker, Fl. Bor.-Amer. 120— 121. 1840. E: Douglas s.n. as annotated by J. Ewan (K s.7., gee Oreophila as (Nutt.) Nutt. ex Torrey & Gray, Fl. N. Am 258 — 259. Pachystima macrophylla Fart, Trans. & Proc. Bot. a Pennsylvania 1:421—422. 1904. — Tyee: Farr 5.n. (PH 37408!, GH ke Pachystima Rrautteri Fart, Ottawa Naturalist 20: ‘108. 1906. — Type: Krautter s.n. (HOLOTYPE: PH 42752!). Pachystima schaefferi Farr, Ottawa Naturalist 20:108. 1906. — Type: Schaffer 512 (HOLOTYPE: PH s.n.!). Paxistima myrtifolia ( ee ) Wheeler, Amer. Midl. Naturalist 29:793 — 794. 1943. Shrub (20-)30 — 100 cm high, the stems sometimes nearly prostrate; adventitious roots may be present. Leaves approximate, 1 — 2 pairs (rarely more) per cm of branch length; blades obovate to oblanceolate, occasional- ly ovate (or elliptic) to lanceolate (or narrowly elliptic), (9-)11— 27(-40) mm long, 4— 10(-15) mm wide; blade margins serrulate to crenulate (occasionally entire), revolute to subrevolute or not (sometimes thickened when not revolute); teeth pointed or rounded, extending from apex to 2/5 to 7/10 (occasionally 4/5) of blade length; blade secondary veins indistinct below (occasionally evident); blade apex obtuse, the apical angle (90°-) 105° — 165°; petioles generally (0.8-)1 — 2(-2.5) mm long. Inflorescences axillary or terminal, averaging 10(3 — 21) per branch, generally composed of (1-)2(-3) flowers each; length of central or only inflorescence axis (1.5-) 2 — 4(-8) mm. Calyx lobes depressed-ovate to very widely depressed ovate, slightly imbricate. Fruits 4— 7 mm long. Typification: No prior type was chosen for Paxrstima myrsinites (1.€. subsp. myrsinites), as confirmed by Wheeler (1943). Two specimens (ol lected by Meriwether Lewis) were mentioned by Pursh (1814) in his description of [/ex? myrsinites, one from “near the Pacific Ocean,” collected November 16, 1805, the other from “on the Rocky-mountain,” collected June 16, 1806. The Lewis and Clark Herbarium at the Academy of Natural Sciences, Philadelphia, contains specimens so designated. An 1805 specimen is also in the herbarium of the Royal Botanic Garden, Kew, England. The Kew specimen is, however, part of a mixed collection (with a Berberis specimen) and is problematic as type material. The 1806 (Rocky Mountain) specimen (PH) seems preferable as the lectotype, and we so designate it. Distribution: Variously known as mountain-lover, Oregon boxwood, myrtle pachistima, myrtle box-leaf and box-leaf, Paxzstima myrsinites subsp. myrsinites is common in the mountain ranges of western North America at altitudes of 600 to 3350 meters. Its range extends from southern British Columbia and Alberta south into Arizona and New — 242 Mexico. The flowers bloom from mid-March to mid-July. This subspecies is quite variable in vegetative morphology. Further investigation may reveal genetic or clinal bases for this polymorphism. Representative specimens: CANADA: Alberta: Watertown Lakes Park, trail co Bertha Lake, 12 Jun 1925, Malte and Watson 289 (\W/TU). British Columbia: Bear Creek Station, Selkirk Mountains, 25 May 1905, Schaffer s.n. (GH, PH, Type of P schaefferi), Bear Creek Station, eastern slope Selkirk Mountains, 20 Aug 1904, Farr s.n.(GH, PH, Isotype of P. macrophylla), Deer Park, Lower Arrow Lake, 4 Jun 1889, Macoun 4058a (MSC); Vancouver Thetis Lake, four mi NW of Victoria, 15 May 1956, Calder, Parmelee and Taylor 16363 (UC). UNITED STATES: Arizona: Apache Co.: Lukachukai Mountains, wooded N slope, | Jun 1950, Clark 15329 (UNM). Cochise Co.: Chiricauhua National Monum ment, Echo Park Trail, 15 Aug 1975, Mason and McManus 3166 (ARIZ). Coconino Co.: Oak Creek Canyon, West Fork, 10 mi N of Sedona, West Fork trail #108, 23 Mar 1988, Navaro s.n. (MU). CALIFORNIA: Del Norte Co.: Shelly Creek Canyon, 3 mi S of Old Monumental, 21 May 1937, Parks and Parks 5646 (DS). Sas Co.: — Summit, 2 mi E of Box Camp, 23 Jun 1942, Tracy 17246 (UC). n Co t. Tamalpais, see! between Laurel Dell and Barth’s Retreat, 16 Mar oan ae ell ie (CAS). Shasta Co.: northern Sierra Nevada, Hatchet Creek, E of Round Mountain, 18 Jul 1930, pe ee (POM). Siskiyou Co.: Black Butte, 15 Jul 1905, Krautter s.n. (PH, Holotype krauttert), Black Butte, [5 Jul 1905, Krautter s.n. (PH Meena of P. krauttert). Yuba : Willow Creek, near Camptonville, 6 Mar 1966, Mott J.n. (CAS). CoLorapo: ee Id Co.: Trappers’ Lake, 30 Jul 1933, Hermann 5503 (GH). Grand Co.: Routt National Forest, Gore Pass on Highway 84, 1 Aug 1962, Porter and Porter 9187 (M C). Gunnison Co.: old town of Gothic, E side of East River, 23 Jun 1952, Barrell 43- - (US). Las Animas Co.: above Whiskey Pass Rd., 6 mi W of Monument Lake campground, 18 Jun 1941, Robbins s.n. (ARIZ). Montesunia Co.: Mesa Verde National Park, rocky canyon below main lodge, 10 Jul 1941, McVangh s.n. (UC). Summit Co.: 8 mi N of elec Blue R. ee 22 Jun 1982, Weber and Wittman a (CM). IDAHo: ae clk Co.: SW slope of Smith 10 jal 1930, Borell s.n. (CAS). Bear Lake Co.: Bear Lake, 1921, ae SM, (DS). Bonner Co. oe mi W of Sand Point, slope ce Clark's Fork River, 14 May 1936, se 2891 (WTU). Clearwater Co.: in brush at summit between Bovill and Elk River, 1949, Cronqutst 5781 (NCSC). Idaho Co.: Lolo Pass, 27 May 1938, Barkley 2417 on Teton Co.: 6 mi W of Driggs, Packsaddle Creek Canyon, | Jul 1968, Muir 5.2. (POM). Montana: Flathead Co.: Rescue Creek and US 2, 28 Jun 1950, Marshall 1176 (MSC). Glacier Co.: Glacier National Park, trail to Mount Brown lookou t, 7 Jul 1939, pea and rie 113(TENN). Powell Co.: 2 mi NW of Woodworth School, Cottonwood ay 1933, Hitchcock 1584 (POM). New Mexico: Catro n Co.: Gila Primitive a 21 May ae Sharp and Orr 332 (PH). Grant Co.: 5 mi N of Pinos oo mountain side above Ree) Creek, 24 Apr 1947, MeVaugh and aa 8051 (GH). yuntains, Karr Canyon, about | mi W of N.M. highway 64 ae 1980, Worthington 6l > (ARIZ). Rio Arriba Co.: Jemez Mountains, San ee Parks, 12 Jun aa Martin, Smith and Schmitt 64-18 (UNM). San Miguel Co.: headwaters of — Rio Las mpas, west of Spring Mountain, 21 Sept 1972, sa 54499 (POM). Taos > 3 mi SE of Taos, Devisadero Peak, 7 Jun 1979, Baker 1033 (NCU). OREGON: Baker ra aoe Wallowa Mountains, Pine Creek, 30 Jun a Jones 7204 (UC). ee Co.: 4 mi N of North Sister Mountains, near McKenzie Pass, 22 Jun 1939, Hitchcock and Sete 4862 (POM). Hood River Co.: Mount Hood National Forest, near Sherwood n pan > n, 243 Forest Sie 13 Aug 1933, Jones 4198 (POM). Lake Co.: Gearhart Mountain region, 3 mi of Finle E ey Corral, 21 Jul 1932, Applegate 7918 (CAS). Josephine Co.: Siskiyou Mountain, phere Ranger on on Sturgis Creek, 5 Aug 1930, Applegate 6597 (CAS). Polk Co.: 4 mi SW of Buell, bank along Mill Creek, | Jul 1930, Peck pOz0E UC). Uran: Box Elder ae Raft River mountains, Clear Creek Canyon, 24 Jun 1947, Preece 644 (UT). Cache Co.: W of Tony Grove Lake, rocky cliffs, 25 Aug 1950, Tiere and Thieret 204 (GH). Kane Co.: whe crite National Park, one half mi E of Rainbow Point, 17 Jun 1957, Buchanan 132 (UT). Salt Lake Co.: top of Clayton Peak, Big Sopa Canyon, 18 Jul 1960, Cottam, nine yee Rowland 160491 (UT, CAS). - Juan Co.: canyon wall opposite Augusta Natural Bridge, 14 Sep 1939, Cutler s.n. (GH a, Co.: Zion National Park, Hidden Canyon, Weights 9772 (UT). WasHINGTON: Chelan Co.: open woods near Merritt, 12 May 1934, Jones 4754 (ARIZ). Columbia Co.: Blue Mountains, stream banks, 23 Jun 1897, ae iM, ae Island Co.: Whidby Island. Goose Rock, 21 May 1933, Thompsson 8940 (C Mount Ranier National Park, trail to Trump Park from Christine Falls, 3 vi ae eva 372 (MU). Okanogan Co.: near summit on Twisp cut-off, 27 May 1932, Fiker 717 (DS). Snohomish Co.: 14 mi N of Seattle. Jun 1892, i SM. res Spokane Co.: Mount Carleton, 21 Jul 1902, Krerger 286 WT). Wyominc: Fremont Co.: along a small creek half way between Lander and South Pass City, 3 Jun 1939, Craig and Craig 3575 (POM). Teton Co.: Teton Pass. 10 Jul 1989, Porter and Porter 7902 (DS) — 1B. PaxistiMa Myrsinires (Pursh) Raf. subsp. MEXICANA Navarro & Blackwell, subsp. nov. Differt a subsp. myrsanites statura parva, foliis Coarctatioribus et parvis. e¢ intlorescentits paucioribus (6) per ramos. Shrub or subshrub 20—45 cm high, the stems sometimes prostrate: adventitious roots may be present. Leaves approximate, 2 (occasionally ) pairs per cm of branch length; blades lanceolate (6-)8 — 12(-15) mm long, G-)4(-5) mm wide; blade margins crenulate (occasionally serrulate, rarely entire), revolute; teeth generally rounded (occasionally pointed), ex- tending from apex to 1/3 to 3/5 (occasionally 3/4) of blade length; blade secondary veins indistinct below; blade ie obtuse, the apical angle generally 90°-135°); petioles (0.8-)I(-2.5) mm_ long. Intlorescences axillary or terminal, averaging 6(3 — 9) per branch, generally composed of 1(-2) flowers each; length of central or only inflorescence axis (1.5-)2 — A(- 5) mm. Calyx lobes widely depressed-ovate to very widely ovate, slightly imbricate. Fruits 4—5S mm long. Type: MEXICO. Coanuita. Municipality of Arteaga, La Siberia, Sierra de la Marta, 27 May 1982, Villarreal 1678 (HoLotypE: MU 134452; isorype: TEX s.n.). Distribution: Paxistima myrsinites subsp. mexicana is apparently restric- ted to mountainous regions of three Mexican states: southeastern Coahuila, southern Nuevo Leon and southwestern Tamaulipas. It grows at altitudes of 2440 to 3500 meters on open hillsides or in forests of pine, fir and oak. The flowers may be found in bloom from late March to mid-July. 244 Representative specimens: MEXICO: Coahuila: municipality of Arteaga, La Siberia, 6 km SE of San Antonio de las Alazanas, 27 May 1982, Villarreal 1678 (MU, TEX, Type of P myrstnites Subsp. mexicana); municipality of Arteaga, Puerto de la Siberia, 10 Oct 1970, Marroquin 1994 (UNL), municipality of Arteaga, Sierra Madre Oriental, 26 Jul 1975, Robert and Passini s.n. (ANSM); 40 mi § of Saltillo, Sierra Madre, Jul 1880, Palmer s.n. (PH); 26 km NW of Fraile, 16 Jul 1941, Stanford, Retherford and Northcraft 454 (CAS); Sierra de la Marta, 17 May 1981, Poole 2324, Hinton and Nixon (TEX). Nuevo Leon: mes ea of Galeana, road to summit of Cerro Potosi, 12.5 mi from 18 de Marzo, 18 1982, Dorr 2270 and Atkins (TEX, ARIZ); municipality of Galeana, canyon below Las Cano: is on Cerro Potosi, 20 Jul 1935 Mueller s.n. (GH); municipality of Galeana, Sierra La Marta, 19 Apr 1981, Hinton 18158 (TEX); municipality of Zaragoza, El Salto, 29 May 1980, Plores O. 5.2. (UNL). Tamaulipas: 20 km NE of Miquithuana, Cerro El Borrado, 2 Apr 1969, Gonzales-Quintero 3855 (MSC); on E and § slopes of summit of Pena Nevada, 19 Jul 1949, Stanford, Lauber and Taylor s.n. (RSA). 2. PAXISTIMA CANBYI Gray, Proc. Amer. Acad. Arts 8:620. 1873. (spelled Pachystima canby1 by Gray — Type: 1869, Canby s.n. (HoLoTyPE: GH!; see typification, below). Shrub or subshrub 10—40 cm high, tending to spread in vegetative clones; older portion of stems prostrate, the upper portion ascending; adventitious roots common on lower stem. Leaves closely approximate, 2—4 pairs (rarely more) per cm of branch length; blades narrowly elliptic to lanceolate, 11 — 22 mm long, 2.3 — 6.2 mm wide; blade margins serru- late to crenulate, strongly revolute; teeth pointed or rounded, extending from apex to 1/3 to 4/5 of blade length; blade secondary veins indistinct below; blade apex obtuse, the apical angle 105° — 120°; petioles (0.5-) I(- 1.1) mm long. Inflorescences axillary or terminal, averaging 4(1—6) per branch, generally composed of 1 — 2(3) flowers each; length of central or only inflorescence axis (1-)2 — 6(-14) mm. Calyx lobes widely depressed- ovate to widely ovate, slightly imbricate. Petals exceeding the calyx lobes, commonly maroon (those from buds from preceding season), occasionally green (those from buds from current season). Fruits 4 mm long, rarely seen. Typification: A specimen at the Gray Herbarium (collected by Canby in 1869) was annotated as the holotype by Vernon Bates in 1984. Asa Gray's 1873 description of Paxistima canbyi states, “Mr. Canby discovered the Alleghenian species in 1868, and obtained flowering specimens upon a second visit to the station in the spring of 1869.” In actuality a small sterile specimen was collected by Canby in 1868 (1858?; cf. Canby in Gray, 1873). However, in regard to collection of specimens by Canby, Gray allu- ded only to those gathered in 1869 (these being flowering specimens, presumably from a single collection), and it was apparently these upon which Gray based his new species. Consequently, the May, 1869 collection 245 by Canby (William Canby s.n.) from Giles County, Virginia is the type collection; the specimen at GH, annotated by Bates, is accepted as the holotype; an isotype is at US Distribution: Paxistima canbyi, variously called Canby’s mountain- lover, cliff-green or rat-stripper, occurs very locally in the Appalachian Mountain region of the eastern United States; it is found on dry to moist, sunny to shaded, northwest to southwest facing, limestone bluffs and ravines in South-central Ohio and Pennsylvania through the Virginias into Kentucky, North Carolina and northern Tennessee. The North Carolina population is at an old nursery site and is considered to have been introdu- ced (Hardin 1963). The presence of P. canbyi in North Carolina was, how- ever, noticed as long ago as 1883 by Chapman, and P canby7 is likely native to North Carolina. Endemic to a small number of areas in these states mentioned above, P. canbyi is listed in Category Two of plants of federal concern, 1.e., more data needed to support listing as threatened or en- dangered (Ohio Division of Natural Areas and Preserves 1988). Paxistima canbyi typically flowers from late March into May, the flowers developing from buds formed during the preceding season. However,a small number of flowers may arise from buds of the current season; these may bloom dur- ing the summer. Representative specimens: UNITED STATES: Kentucky: : Carter Co.: Carter Caves, Devil's Backbone Ridge, 29 May 1986, Navero s.n. (MU); Carter Caves, eee cliff opposite entrance, 29 May 1986, Navaros.n. (MU). Pulaski Co.: Tatesville, S, Lake Cumberland, 10 May 1976, Stephens 5.n. (TENN). On10: Adams Co.: Brush oo Twp., Edge of Appalachia Preserve, 9 Apr 1987, Navaros.n. (MU). Highland Co.: Brush Creek Twp., Fe. Hill Sc. Memorial Park, 1 Apr 1973, Bourdo and Roberts 3294 (OS). NortH CAROLINA: 1874, Canby 5.n. (PH). PENNSYLVANIA: Bedford CO.: Cliff at Lutzville, 6 May 1950, Henry and Buker s.n. (CM); Juniata R. near Lutzville, 6 May 1950, Krowse 97 (CM). TENNESSEE: Hawkins Co.: bluff te South Fork Holston River, Bays Mt. near Laurel Run Gorge, 21 Apr 1984, Somers and Smith s.n. (TENN). WirGinia: Frederick Co.: west of Middletown, above Cedar Creek, 20 Sep 1931, Griscom and Hunnewell s.n. (GH). Giles Co.: May 1869, Canby s.n. ye ao yates s Co.: VMI post, above Maury R., 22 Apr 1963, Guptons.n. (N . Scott : Natural Tunnel, around the rim of tunnel, 17 May 1968, James 9686 fon ae oa near Wytheville, Jun 1875, oe 483 (G — VIRGINIA: Greenbriar Co.: Chocolate Drop, limestone cliff facing cee R., | Aug 1931, MeNe// 5.x. (WVYA). Mercer Co.: mouth of Brush Creek, 4 May 1976 os s.m. (WVA). Mineral Co.: near Keyser, May 1936, Chapman s.n. (W VA) DISTRIBUTION AND GEOFLORISTRIC HISTORY The present distribution of Paxistima in North America (Fig. 2) is prob- ably attributable to its presence in the temperate Arcto-Tertiary forests, and to subsequent geoclimatic restrictions upon these forests, 1.e. orogenic activity, cooling/drying, glaciations. According to Cine: 246 FIG. 2. Geographic distribution of Paxistima in North America. Star = P. canbyi, solid circle = P Bray ry myrsinites subsp. myrsinites, triangle = P. myrsinites subsp. mexicana. 247 (1947), “The Arcto-Tertiary Flora has survived in North America at middle latitudes in two main provinces, an eastern characterized by broad- leafed, deciduous trees, and a western characterized by conifers, broad- leafed evergreens, and broad-leafed deciduous trees and shrubs.” The two species of Paxistima, P. canbyi and P. myrsinites, are indeed presently restric- ted, respectively, to parts of these two regions. Additionally, the pattern and the restricted (localized) nature of the present distribution of P. canby7 have led some (e.g. Transeau 1941) to con- sider this distribution explainable by association with the former north- west-flowing, preglacial Teays River. However, populations generally lie outside the supposed Teays drainage per se (see Steeg 1946, for an account of the Teays drainage). On the other hand, several populations may be cir- cumstantially related to the boundaries of the glacial lake (Lake Tight) formed by ice blockage of the Teays (Wolfe 1942; Braun 1950). The details of the explanation of the distribution of P. canby7 require further elucida- tion. The origin and relationships of Paxistima myrsinites subspecies mexicana are worthy of conjecture. Although most similar to subspecies myrsinites, the variation of subspecies mexicana in the “morphological direction” of P canbyi (Fig. 1) suggests that the Arcto-Tertiary antecedents of present day Paxistima perhaps constituted one transcontinental species complex which later became disjunct (developing more or less morphologically distinct entities) as a consequence of geoclimatic events, such as those mentioned previously. Subspecies mexicana may represent relic populations of the ormer myrsinites-canbyt complex, remaining in a refugium in the mountains of Northeast Mexico; it could also represent a third line of development from an ancestral species. ACKNOWLEDGEMENTS We wish to express our appreciation to the curators of those herbaria mentioned in the Materials and Methods section for loaning specimens utilized in this study. We also wish to thank Dan H. Nicolson for his helpful suggestions regarding orthography and nomenclature during the preparation of this manuscript. This research was funded in part by the Willard Sherman Turrell Herbarium Fund (MU) grants #72 and #85. REFERENCES ANDERBERG, M.R. 1973. Cluster analysis for Applications. Academic Press, New Yor BRAUN, E.L. 1950. Deciduous forests of eastern North America. Hafner Press, New York. 248 CHANEY, R.W. 1947. Tertiary centers and migration routes. Ecol. Monogr. 17:139 — 148. CHAPMAN, A.W. 1883. Flora of the southern United States. Ivison, Blakeman, Taylor and Co., New York. DON, D. 1833. Desc riptions of the new a. ae nee of the class Compositae belong- ing to the floras of Peru, Mexico and Chi . Linn. Soc. London 16:178. DU RIETZ, G. E. ee The ee units “of ee taxonomy. Botanisk Tidsskrift 24:333 — ENDLICHER, S. ne oe Genera plantarum secundum ordines naturales. Beck, Vie ee ICHER, S. 1841. Genera plantarum secundum ordines naturales, Supplement. Beck, Vie FARR, ELM. ‘1904. Notes on some ae British Columbian plants. Trans.& Proc. Bot. oie Pennsylvania 1:417 — FARR, E.M. 1906. Some new ie from the Canadian Rockies and Selkirks. Ottawa See 20:105— 111. GENAUST, H. 1976. Ns ies aii Worterbuch der botanischen Pflanzennamen. Birkhauser Verlag, Basel. GRAY, As 1373. ei ee of new genera and species of plants. Proc. Amer. Acad. Arts 8:620— 631 GREUTER, W. et al.(eds.). 1988. International code of botanical nomenclature. Koeltz Scientific Books, Konigstein, Federal Republic of Germany. HARDIN, J.W. 1963. Pachystima canbyi in Now ere Castanea 28:177 — HOLMGREN, PK., W. KEUKEN and E.K. SCHOFIELD. 1981. Index herbariorum I. Bohn, Schietrenid. and Holkema, Utrecht. HOOKER, W.J. 1840. Flora Boreali-Americana. Henry G. Bohn, London. MEISNER, C.F 1843. Plantarum vascularium genera. Libraria Weidmannia, Leipzig. MERRILL, E.D. 1949. Index Rafinesquianus. Arnold Arboretum, Jamaica Plain, Massachusetts. NUTTALL, T. 1818. The genera of North American plants, and a catalogue of the species, o the Year 1817. Philadelphia OHIO DIVISION OF NATURAL AREAS AND PRESERVES. 1988. Rare native Ohio vascular plants: 1988 — 1989 Status List. Ohio Department of Natural Resources, Cite 13. PIPER, C.V. 1906. Contr. U. S. Natl. Herb. 11. Flora of the State of Washington. Gov- ernment Printing Office, Washington. PURSH, E 1814. Flora americae septentrionalis,]. White, Cochrane, and Company, oO RAFINESQUE, C.S. 1818. Review of Pursh’s flora of North America. Amer. Monthly Mag.& Crit. Rev. I(3):175 RAFINESQUE, C.S. 1819a. Revie of the genera of North American piants and a catalogue of the species in the year 1817 by Thomas Nuttall. Amer. Monthly Mag. & Crit. Rev. IV: 184— 191 RAFINESQUE, C.S. set Remarques, critiques et synonymiques. J. Phys. Chim. Hist. Nat. Arts 89:256— RAFINESQUE, C.S. Sylva Telluriana. Philadelp! SAS Institute, Inc. 1985. SAS user's guide: Statistics, es 5 Edition. Cary, North Carolina. 249 STANDLEY, PC. 1920— 1926. Contr. U.S. Natl. Herb. 23. Trees and shrubs of Mexico. Government Printing Office, Washington. STEEG, K. V. 1946. The Teays river. Ohio J. Sci. 46:297 — 307. TORREY, J. and GRAY, A. 1838. A Flora of North America I. Wiley and Putnam, New York. TRANSEAU, E. N. 1941. Prehistoric factors in the development of the vegetation of Ohio. Ohio J. Sci. 41:207 — 211. UTTAL, L.J. 1986. Once and for all it is Paxistima. Castanea 51:67 — 68. WATSON, S. 1878. Bibliographical index to North American Botany. Smithsonian Misc. Collect., XV. Smithsonian Institution, Washington. WHEELER, L.C. 1943. Ak and orthography of the celastraceous genus “Pachystima” Rafinesque. Amer. Mid]. plist 20 192 192 WOLFE, J.N. 1942. Species ae and a proglacial ees in southern Ohio. Ohio J. Sci. 42:2—11. Book REVIEWS Biackwe_, Witt H. 1990. Poisonous and medicinal plants. 329 pp. Illus. Price unknown. Prentice-Hall, Inc., Englewood Cliffs, NJ 07632. Illustrations are by Thomas J. Cobbe with Chapter 5 (Poiso- nous and Medicinal Fungi) by Martha J. Powell The combination of medicinal and Br ee plants within the same volume ts a natural one stemming from the beginning of man’s knowledge of plants. Since poisons are medicinal and medicines are So. it is merely the dosage and the sensitivity of an individual human being that determines the desired result. This is an excellent book for the classroom, reference, or just interesting reading. WESTERN, DavibD AND Mary Pear. (Editors). 1989. Conservation for the Twenty-first Century. 365 pp. Hardbound. $36.95. Oxford University Press, 2001 Evans Road, Cary, NC 27513. The proceedings of the conference “Conservation 2100: A Fairfield Osborn Symposium” are published tn this volume. Thirty-two contributors have published articles pertaining to the following topics: I. Tomorrow's World; UH. The ae! of Conservation; II]. Conserva- tion Management; IV. Conservation Realities; V. An Agenda for the Future. ae text covers a broad spectrum of facts and ideas ae an ea and global viewpoint. It is recommended for all persons interested in any aspect of our future on this planet. Sipa 14(2):250. 1990. A TAXONOMIC COMPARISON OF ARISTIDA TERNIPES AND ARISTIDA HAMULOSA (GRAMINEAE)! JONATHAN S. TRENT 525 Pinar Del Rio, El Paso, TX 79932, U.S.A. KELLY W. ALLRED Department of Animal and Range Sciences Box 3-1, New Mexico State University Las Cruces, NM 88003, U.S.A. ABSTRACT The morphologic similarity of Aristida ternipes and A. hamulosa was assessed. All 29 measured variables exhibited renee overlap in their ranges, and only eight of the 29 had correlations greater than . Multivariate anes component and discriminant) ceca revealed a lack of sien patterning; only awn lengths discingulshes the taxa. entities are recognized at the varietal level. The epee aes combination A. Go var. hamulosa (Henrard) Trent is made. RESUMEN Se evalus la similitud morfolsgica entre Aristida ternipes y A. hamulosa. Las 29 variables medidas mostraron considerable superposicisn, y solamente ocho de ellos dieron correlaci- ones mayores que 0.50. Un analisis multivariada revels una carencia de patrones feniticos; unicamente la longitud de las aristas sirvis para distinguir los taxa. Las dos entidades son reconocidas a nivel variedad. Se propone la combinacisn A. fernipes vat. hepsi (Henrard) ont Two commonly encountered grasses in the southwestern United States are Aristida ternipes Cav. and A. hamulosa Henr. Both are common on dry, sandy plains and hills of low desert areas, and not infrequent at higher elevations in foothills and on mesa slopes. In general habit the two species are quite similar, with small basal tufts of foliage and large, stiff, widely spreading panicles. They differ most conspicuously in the development of their lateral awns, those of A. ternipes being very short (often hardly notice- able) and those of A. hamulosa being well-developed and obvious. Henrard (1927, p. 221)also called attention to the “curious” tuberculate lemmas of A. hamulosa when he described the species. The difference in awn lengths ‘Journal Article 1391, New Mexico Agr. Exp. Sta., New Mexico Srate University, Las Cruces. This paper represents a portion of a thesis by the first auchor in the Department of Animal & Range Sciences, New Mexico State University. Siwa 14(2):251— 261. 1990. 202 has traditionally placed the two species into separate sections of the genus: A. ternipes in the section Streptachne, and A. hamulosa in the section Aristida (Chaetaria) (Henrard 1929, 1932). However, the two species seem to be more similar than this classification would suggest. Most North American botanists (Hitchcock and Chase 1951; Kearney and Peebles 1969; Beetle 1983) have accepted Hitchcock's (1924) and Henrard’s (1926, 1928) treatment of Aristida ternipes and A. hamulosa as separate entities without evident relationship. Correll and Johnston (1970) suggested that A. hamulosa may be only a form of A. divaricata, but Gould (1951, 1975) called attention to the similarity of A. hamulosa with both A. ternipes and A. divaricata. The purpose of this study was to evaluate the taxonomic relationship of Aristida ternipes and A. hamulosa by assessing the variability in morphologi- cal features and by testing the characters that traditionally have been used to separate them | for characters used in the statistical analysis of Aristida ternipes Taser |. Summar y and A. hamutosa. Character scored Acronym States Culm height CULMHT Continuous Blade width BLADEW Continuous Blade conformation BLADECON O-flat l-some involution Invo Blade pubescence BLADEPUB 0-glabrous scene 2- ee pubescent Collar pubescence COLLPUB 0-glabro l-some ae € 2-strongly eae Ligule length LIGULEL Continuous Panicle length PANL Continuous Longest primary branch length PRIBRNL Continuous Distance to first spikelet SPKLTDIS Continuous First i, ot sngth SECBRNL Continuous Terminal primary branch sngth TERMBRNL Continuous Lateral pedicel length PEDL Continuous Maximum pes of branches per node BRNCHNUM Continuous Panicle branch spreading BRANINDX ratio of spreading (Branching Index) secondary and tertiary branches to the number o primary branches yo) Central awn length CAWNL Continuous Lateral awn length LATA Continuous First glume lenge FSTGLUMI Continuous Second glume length SECGLUML Continuous Glume pubescence GLUMEPUB 0-glabrous l-some pubescence 2-highly pubescent Callus a CALLUSL Continuous Floret length FLORETL Continuous Width of aa at idest point LEMMAW Continuous Ww deh of lemma at narrowest point LEMMAN Continuous Lemma texture LEMMATXT Q-smooth 1-cuberculate 2-scabrous Awn column length AW NCOLL Continuous Awn column twisting COLLTW O-no twisting 1-1 turn 2-2 or 3 turns 3-4 or more turns Anther length ANTHERL Continuous Palea length PALEAL Continuous Elevation of collection ELEV Continuous MATERIALS AND METHODS Field collections of Arsstida ternipes and A. hamulosa were made from populations in Arizona, Colorado, New Mexico, Texas, and Chihuahua, Mexico; emphasis was placed on collecting all forms present in a popula- tion. The field collections were supplemented by herbarium material from throughout the range of the species, including California, Mexico, and Guatemala. From all material gathered, specimens were selected for study that represented the variability present in the two taxa as well as the geo- graphic range of the species. A data set for morphometric analysis was compiled by scoring selected specimens (field and herbarium) for the fea- tures listed in Table 1. Only mature specimens were included in the analy- sis, determined by complete emergence of the panicle from the sheath. A total of 92 individuals were measured. A list of specimens examined may be requested from Allre he DP statistical patkace (Dixon 1981) was used for analysis. In addition to standard, descriptive statistics such as mean, range, standard deviation, and correlation coefficients for all variables, principal components analysis (PCA) was used to assess the morphological similarity or dissimilarity of the specimens (OTUs). Based on a variable by variable correlation matrix, the PCA plotted the OTUs along each component 254 according to its phenetic similarity to each other OTU. Groups, or classifi- cations of the OTUs, suggested by the PCA were then tested by stepwise discriminant analysis (SDA). SDA determined the potential for variables to cause disjunctions between two or more a priori groupings (in this case, those implied by PCA or those specified by a particular variable). A “grouping variable” segregated the OTUs into groups and the analysis determined if these groups were recognizable by the statistical relation- ships of the remaining variables. Output from SDA included the percent- age of OTUs classified “correctly” or “incorrectly,” that is, the percentage corresponding to the a priori groups. A high percentage of correctly classified OTUs indicated that the a priori classification was supported by the other variables. SDA was also used to test the importance or validity of certain variables in creating groups. Lateral awn length was used as the grouping variable, specifying two groups based on a cut-point value of 2.5 mm (those OTUs with lateral awns less than 2.5 mm were assigned to ternipes, those with lateral awns less greater than 2.5 mm were assigned to hamulosa). The SDA then determined if the resultant groups were sup- ported statistically by the remaining variables. RESULTS AND DISCUSSION Morphometric Analysis. Correlation coefficients were calculated for all combinations of characters. All of the correlations greater than 0.50 were with continuous size variables (Table 2), but, the only variables that showed correlations higher than 0.80 were panicle and spikelet features related to specimen size: culm height with panicle length (0.86), primary branch length with panicle length (0.86), central awn length with lateral awn length (0.84), and first with second glume length (0.84). In general, as the size of the specimen increased, the size of the panicle also increased; likewise, the size of spikelet parts tended to increase or decrease in concert. It 1s noteworthy that lemma texture had no high correlations, eventhough Aristida hamulosa had been characterized by its prominent tubercles on the lemma (Henrard 1927). The means and ranges of features with correlations higher than 0.50 were then compared between Aristida ternipes and A. hamulosa (Figure 1). The assignment of OTUs to one of the two taxa was based on lateral awn length because of its traditional importance in identification. OTUs with lateral awns longer than 2.5 mm were assigned to hamulosa, and those with shorter awns to fernipes. With the exception of the awn features, the ranges and standard deviations of every character overlapped extensively. Central awn length had overlapping ranges, but not standard deviations. Lateral awn lengths did not overlap because of the a priori assignment of — 255 Fig. TA. Panicle Features 4060 = are 80 | H 60 f + t as | | t i : H T H T 216 : 5 t f i O l i ] l 1 1 : { pod CH CH| PL PL| B BL] SD SO] SBL SBL Fig. 1B. Gib Gib GeL G2 IG. 1. Range, mean, and I one standard deviation for the correlated features of the data set for Aristida ternipes (T) and A. hanulosa CA). 1A. Panicle features, mee astieget inom. CH =culm height; 3 CAL CAL PL = plant height, BL = primary branch length; SD = distance SBL = secondary branch oan, 1B. Spikelec features, measured in mm. CAL =centra awn lanel LAL = lateral awn length; G1iL=first glume length; G2L = second glume ace 256 Taste 2. Correlation coefficients greater than 0.500 of all variables for Aristida hamulosa and A. ternipes using all OTUs. Acronyms according to Table 1. CULMHT PANL PRIBRNL = CAWNL FSTGLUML SECGLUML PANL 0.856 1.000 = = = = PRIBRNE 0.795 0.856 1.000 = = a SPKLTDIS 0.643 0.635 0.749 = an = SECBRNL = 0.912 = = = LATAW NL = = = 0.83 as = SECGLUML = = = 0.639 0.839 1.000 FLORETL = = = = 0.541 0.504 CALLUSL = = = a a= 0.526 the OTUs based on this feature. However, the range in lateral awn lengths varied continuously from Aamulosa to ternipes. The principal components analysis was conducted using the same set of correlated features. The placement of the OTUs along the first component (PCI) was correlated with over-all size features such as panicle length (0.93), longest primary branch length (0.92), culm height (0.91), and dis- tance to the first spikelet on the branch (0.79). The second component (PCII) revealed differences in spikelet features, including second glume length (0.89), first glume length (0.86), and floret length (0.85). The third component (PCIIL) emphasized lateral awn length (0.91) and central awn length (0.82). The three components accounted for 75 percent of the variability altogether. The phenetic distribution of the OTUs along PCI and PCII, which were size and spikelet components, revealed no discernible separation of taxa, and those plots are not shown here. But a segregation of OTUs was achieved along the third component, based on awn lengths (Figure 2). To test the validity of a partition based on lateral awn length, a stepwise dis- criminant analysis was performed that used this character as the a priori grouping variable but not in calculating the discriminant function. The plot of the OTUs along the canonical variate (Figure 3A) indicated that two groups were distinguished; central awn length was the only variable used in calculation of the discriminant function. However, when both lateral and central awn lengths were removed from the analysis, an extensive in- termingling of the OTUs resulted (Figure 3B), and the discriminant func- tion assigned only 69% of the hamulosa OTUs and 60% of the ternipes OTUs to the “correct” a priori group. The results of the statistical analyses indicated that 1) there was a nearly continuous range of morphologic variation from one taxon to the other, with extensive overlap in the ranges of individual variables; 2) two con- tiguous groups of OTUs were segregated based on awn lengths; and 3) no other basis existed, other than awn lengths, for distinguishing the groups. 257 HAAATL Trt tT eo, ae eae eae a 2 Bos Bee Ba aL, ide At Lf Eiht ht £ 2 eee does gee eae? Cage ae, 0 sdk Ale SE 2. ae ae ae ae ae ak. ae =2 ae 0.0 1.0 20 Pio cs Principal Component FIG. 2. Projection of Aristida ternipes (T) and A. hanulosa (A) OTUs along principal component III. H H H Fig. BSA 7 AOA A T HoH H H H H H T TT H H H H H H H H H H H H eo. oT oF EF OF H H H H H H H H H H H H H H H H H H Eo ho eae “Se HoH H HHH HHHHH AHH ARKH AHH HAA THR T OT OT OT OTT tT T T HT T H T H HoH T HT HH T T H H T HH T T me 3B H T HH HH T H H T H H H H H H H H T T TH T HH HH H H H H H H T HH T HH HH H H H H H H H HH H T HoH HoH HHH H H H H H H H OF <0. 245 HO “O05 00.5 WO Te. 2O -2e Canonical Variable (ow FIG. PUStog ratios of Aristida ternipes (VY) and A. hamulosa GA) OTUs along the canonical vartable of si ae analysis. The grouping variable was lateral awn Jengen on Histogram when only | dc tral lateral awn length was ae from the data set. 3B. Histogram lengths were removed from the data set. 258 Other Observations. Field and herbarium studies yielded other im- portant observations. Noted for the first time for both taxa was the con- sistent occurrence of long, weak hairs at the base of the blade above the ligule. Also characteristic were glabrous collars, an untwisted awn column, and anthers generally longer than 1.2 mm. These features dis- tinguished the bamulosa and ternipes entities from the similar-appearing A. divartcata Willd. and A. havardii Vasey (commonly known as A. barbata Fourn.). Both taxa were found in Texas, New Mexico, and Arizona and throughout most of Mexico. Only the hamulosa entity was found in southern California and southern Colorado, and extended as far south as Honduras, but specimens of ternipes were found from Nicaragua, Costa Rica, the Bahamas, Venezuela, and Columbia, where hamulosa was absent. When sympatric, the two often grew intermingled in the same apparent population and there were no noticeable differences in soil or microsite preferences. The hamulosa taxon has spread to slightly more temperate areas in California and Colorado, and fernipes perhaps represents a more subtropi- cal form. Specimens of hamulosa from California tended to be short in height, with correspondingly short primary panicle branches. The spikelets were also spaced somewhat closer together. The overall effect of these differences was a slightly more congested look to the panicle. California plants could not be distinguished from non-California plants on this basis, however, and numerous small plants with short branches were found within populations from other regions. Aristida ternipes was not found from California. Commonly, branchlets and spikelets were appressed to the axis of the panicle branch (Figure 4A). However, forms with spreading to divaricate branchlets or pedicels were occasionally found in both taxa. This condition was always associated with pulvini in the axils of the branchlets and pedi- cels, causing them to spread outward from their axes (Figure 4B). The ex- pression of the pulvini was measured by the branching index in the morphometric analysis and was not highly correlated with any other fea- ture. Our field observations confirmed this: pulvini seemed to develop arbitrarily in many different populations and both spreading and appressed forms of Aristida ternipes and A. hamulosa were found in the same popula- tion. However, spreading forms transplanted to a greenhouse maintained this feature the following growing season, and pulvini did not appear to be a maturation phenomenon, but were observed in the inflorescences from the time they emerged from the sheath until senescence of the plant. The geographic distribution of the spreading forms was centered in the south- western United States and northern Mexico, with few specimens found 259 FIG. 4. Spreading and appressed inflorescence forms. A. Inflorescence of a specimen of Aristida hamulosa showing the appressed form. B. Inflorescence of a specimen of Aristeda ternipes showing the spreading form. from California or southern Mexico. Spreading forms are likewise found in other species of Aristida, including A. pansa, A. dissita, A. divaritata, and A. havardii. The spreading form of A. hamulosa, in particular, may be con- fused with A. havardii or A. divaricata, but is distinguished by shorter anthers (= 1 mm) and glabrous ligular region in the latter species. Conclusions and Taxonomy. Aristida ternipes and A. hamulosa are near- ly identical morphologically. Apart from the difference in lateral awn length, the two can scarcely be distinguished. Their over-all geographic distributions have considerable overlap, they are found in the same habitats and in intermingling populations, they both display a distinctive pubescence near the ligule, and they share a seemingly arbitrary expression of pulvini in the panicle. A chromosome level of 2n = 44 has been reported for both taxa (Gould 1966, 1968; Stebbins & Love 1941). The recognition of two species based on differences in lateral awn length is unwarranted. Eventhough the two entities can be distinguished only by a single feature, 260 suggesting forma status, we propose recognizing the variation in this complex at the varietal level. This is consistent with treatments of similar variation patterns in other Aristida species and with the widespread lack of sharp boundaries in general between taxa in many North American Aristida (Allred 1984a,b, 1985). Given the priority of A. fermipes in pub- lication date, the correct classification of the ternipes and hamulosa entities would be within the single species A. ternipes with two varieties, var. ternipes and var. hamulosa. The necessary combination for the latter variety is effected below. ARISTIDA TERNIPES Cav. var. HAMULOSA (Henrard) Trent, comb. BastonyM: Aristida hamulosa Henrard, Med. Rijks. Herb. Leiden 54 on. oe, Type: ARIZONA. Tucson, 30 Sep 1894, JW. Towmey s.n. Salient features of the two varieties are compared below: var. fernipes var. hamulosa Lateral awn length )—2.5 mm (2.5)3.5—- 18 mm Central awn length 5-15 mm 10-25 mm Distribution TX, NM, AZ, TX, NM, AZ, CO, Mexico, C. Amer., , Mexico, Guatemala ACKNOWLEDGMENTS We are grateful to the curators of the following herbaria for their generous loans of specimens (acronyms according to Holmgren et al. 1981): AHUC, ANSM, ARIZ, ASU, COLO, ISC, JEPS, LL, NMC, NMCR, NY, OKL, RSA/POM, SD, SMU, SRSC, UNLV, US. Thanks also to two reviewers for their helpful and constructive criticism of the manu- script. David Lee Anderson assisted with the Spanish summary. REFERENCES ALLRED, K.W. 1984a. Morphologic variation and cl: See ation of the North American Aristida purpurea Bae (Gramineac). Brittonia 36(4):3 395. eerie ye ee . Studies in the genus Aristida ates of the southeastern United States. ace et variation in A. purpurascens, A. tennispica, and A. virgata. Rhodora 86(8: 45):73 =. aT 1985. Studes in the Arzstida (Gramineae) of the southeastern United States. I] Mon hone analysis of A. intermedia and A.longespwa. Rhodora 87(850): 137 — 145. BEETLE, A.A. 1983. Las Gramineas de Mexico. Editorial Calypso, Distrito Federal, = CORREL L, D.S. and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Tex DIXON, W.J., ed. 1981. BMDP es software. University of California Press, Berkeley 261 GOULD, EW. 1951. Grasses of the southwestern United States. University of Arizona Press, Tucson. 1966. Chromosome numbers of some Mexican grasses. Can. J. Bot. 44:1683 — 1696. 1968. Chromosome numbers of some Texas grasses. Can. J. Bot. 46:1315 — 1325. 1975. The grasses of Texas. Texas A&M University Press, College Sta- tion, Tex HENRARD, “y T. 1926. A critical revision of the genus Aristida. Vol. I. Med. Rijks Herb. Leiden 54:219 — 220. 1927. A critical revision of the genus Aristida. Vol. Il]. Med. Rijks Herb. Leiden 54A:221— 464. 1928. A critical revision of the genus Arsstida. Vol. III. Med. Rijks Herb. Leiden 54B:630— 631 1929. A oaceaer of the genus Aristida. Vol. I. Med. Rijks Herb. Leiden No. 58 1932. A monograph of the genus Aristida. Vol. Il. Med. Rijks Herb. Leiden No. 58A. HITCHCOCK, A.S. 1924. The North American species of Aristida. Contr. U.S. Natl. Herb. 22:91) 386; and A. CHASE. 195 1. Manual of the grasses of the United States. 2nd ed. U.S.D.A. Misc. Publ. No. 200. Washington, D.C. HOLMGREN, PK. , W. KEUKEN, and E. . ae FIELD. 1981. Index Herbariorum. 7th ed. Bohn, Schelven: & Holkema, KEARNEY, T.H. and R.H. PEEBLES. oo oe University of California Press, Berkeley STE BBINS, res L. and R.M. LOVE. 1941. A cytological study of California forage grasses. Amer. J. Bot. 28:371— 383. CIRCUMSCRIPTION OF AGARISTA BOLIVIENSIS (ERICACEAE) WALTER S. JUDD Department of Botany, 220 Bartram Hall University of Florida Gainesville, FL 32611, U.S.A. PAULA M. HERMANN Departamento de Biologia Universidad Nacional del Sur Per 670 8000 Bahia Blanca, ARGENTINA ABSTRACT Our understanding of the morphological variability of Agarista boliviensis is significantly expanded by an examination of a population in the Sierra de Zapla in the Province of Jujuy in extreme northwestern Argentina. The Sierra de Zapla plants may be distinguished from Bolivian populations of A. dolrviensis by several features relating to the indumentum of ms and leaves, petiole and inflorescence lengths, and leaf margins. The taxonomic status of this population is discussed and a revised description for the species is provided. The pattern of variation of A garista boliviensis is similar to that of the closely related A. excalyptoi- des. The species also is compared with A. paraguayensis, the only other species of Agarista occurring in Argentina Agarista D. Don ex G. Don is a genus of 31 species occurring in both Africa (including Madagascar) and the Americas (Judd 1984; Gonzalez 1989). The genus is most diverse in South America, and is closely related to several genera 1n the Andromedeae (Ericaceae) such as Cratbiodendron W. W. Smith, Lyonta Nutt., and Pzeris D. Don Judd 1979). The morphologi- cal variability of several species is still poorly known, and information regarding one of these, A. bolzviensis (Sleamer) Judd, is reported herein. Agarista boliviensis has been collected in the Sierra de Zapla in the province of Jujuy in extreme northwestern Argentina. It was first reported for the country by Legname (1978) and was listed in the Flora of this province by Cabrera (1983). All other populations of the species are located in central and southern Bolivia. The species occurs in mountainous areas from ca. 1200 to 2500 m altitude. In the Sierra de Zapla of Argentina it occurs 1n moist montane forests and is associated with Podocarpus parlatoret, Alnus acuminata, Eupatorium spp., and Rubus spp. Available herbarium material of A. do/iviensis from the Sierra de Zapla shows several differences from all known specimens of the species collected Sipa 14(2):263 — 266. 1990. 264 in Bolivia, necessitating the following revised species description. As in Judd (1984), the term “pubescent” refers only to the presence of small, more or less whitish, unicellular, nonglandular hairs. DrscripTion: Rhizomatous shrub or small tree to ca. 7 m tall. Twigs glabrous to moderately pubescent, reddish when young, with nonchambe- red to obscurely chambered pith; buds to ca. | mm long. Leaves alternate, reddish on young shoots; blade revolute in bud, + flat at maturity, coriac- eous, ovate, 2—7.5 X 0.9—3.2 cm, base cuneate to rounded and often slightly asymmetric, apex acuminate, margin entire and minutely undu- late to smooth, more or less flat to very slightly revolute at extreme base; adaxial blade surface dark green and lustrous, glabrous or with a very few hairs, especially near margin, but very sparsely to moderately pubescent on midvein; abaxial blade surface glabrous, but very sparsely to moderately pubescent along midvein, with inconspicuous to conspicuous glandular dots along midvein; petiole (3.5-)5— 17 mm long, slender and often flexuous. Inflorescences axillary racemes to 0.5 —4(-6) cm long; rachis moderately pubescent with whitish hairs; pedicels 4 — 9 mm long, sparsely to moderately pubescent; bracteoles 2, opposite to alternate, from basal to within lower 1/3 of pedicel, narrowly triangular to linear (rarely ovate), to ca. 1.7 (rarely 8) mm long; floral bracts to ca. 1.5 mm long. Flowers 5- merous; calyx lobes triangular with acuminate apices, 0.9 — 2 X 0.5— 1.7 mm, abaxial surface glabrous to moderately pubescent, articulated with pedicel, with ca. | — 1.5 mm long projection between calyx and point of articulation; corolla cylindrical, 6-11 X 2—5 mm, abaxially glabrous (or sometimes with a very few unicellular hairs along the veins), white. Filaments 3.5 — 7 mm long; anthers | — 1.2 mm long. Ovary glabrous to moderately pubescent, especially near apex. Capsules subglobose to ovoid, .4—7 mm, placentae subapical; seeds 1.4— 2.6 mm long. Specimens Examined: ARGENTINA. Jujuy: Departamento Capital, Cerro Zapla, Mina 9 de Octubre, Villamil 2936 (BBB, FLAS, NY); ibid., amil et al. 4311 (BBB, FLAS); Departamento Capital, Sierra de Zapla, Burkart et al. 30549 (FLAS [frag.], bee Sierra de Zapla, Mina 9 de Octubre, es ipak et al. 32009 (SD. BOLIVIA. Chuquisaca: Sucre, Alto de Aguas Blancas, eae 249 ( oh Cochabamba: Rosal, below pumping sta- tion, Brooke $702 ¢ E NY). hare: , San Pedro, Pasopaya, Anonymous 3400 (GH). Santa Cruz: Tres Cruces, Herzog 1634a L). Tarija: lomas peladas, Alto de las Canas, Troll 359 (B, M); Camino de Emborozu, La Minors Turpe et al. 4777 (BAA). The very close relationship of the recent collections from the Sierra de Zapla in northwestern Argentina to those from central and southern Bolivia, t.e., typical Agarista boliviensis, is seen in the fact that plants from both regions share several characters: absence of multicellular gland- headed hairs; ovate leaves that are more or less flat at maturity, with slender 265 and at least sometimes slightly flexuous petioles and acuminate apices; often short inflorescence axes that are moderately pubescent (with whitish hairs); white flowers with short calyx lobes; and capsules with subapical placentae Judd 1984). The Sierra de Zapla population is the southernmost of the species. As is often the case in isolated peripherial populations (Mayr 1969), it is somewhat distinctive morphologically. Plants from this popu- lation usually can be differentiated from Bolivian plants by their sparsely to moderately pubescent twigs (vs. glabrous to sparsely pubescent); leaves with the midvein more or less moderately pubescent (vs. only very sparsely pubescent); petioles 3.5 — 10 mm long and not or only slightly flexuous (vs. 6— 17 mm long and frequently flexuous); leaf margins entire and smooth to obscurely undulate (vs. usually entire and minutely undulate, but rarely only obscurely undulate); and inflorescences 0.5 — 4(-6) cm long (vs. 0.5 — 2.5 cm long). Some of the flowers on the Sierra de Zapla plants also have longer filaments (to 7 mm) than those seen in flowers of Bolivian plants (to 4.5 mm). The Sierra de Zapla plants have been illustrated by Cabrera (1983) and a typical Bolivian plant of A. bol/viensis was pictured in Judd (1984). Initially, we considered giving varictal rank to this distinctive popula- tion of Agarista boliviensis in the Sierra de Zapla. However, additional study of available material indicatec eon on that formal taxonomic recognition is un- warranted due to the degree of overlap in the presumed diagnostic characters, and because an extremely similar pattern of variation is shown by the closely related A. eacalyptoides (Chamisso & Schlechtendal) G. Don (see Judd 1984). Agarista encalyptoides is a widely distributed species of southern Brazil that shows variation in stem and leaf pubescence, degree of undulation of leaf margin, length and amount of flexuousness of the petiole, and inflores- cence length Judd 1984). It is, thus, not too surprising that additional collections of A. boliviensis have revealed extensive variability in these same features. Agarista eucalyptoides can easily be distinguished from A. bolzvien- sis by its ovate to oblong leaves and the indumentum of its inflorescence axes, 1.e., densely covered with ferrugineous hairs in A. eacalyptoides in contrast to moderately pubescent with whitish hairs in A. bolzviensis Judd 1984). The only other species of Agarista occurring in Argentina is A. paraguay- ensts (Sleuamer) Judd. This species grows in northeastern Argentina in Mis- iones province as well as several localitics in Paraguay Judd 1984). Agarista boliviensis differs from A. paraguayensis in several features: longer and occasionally slightly flexuous petioles, consistently acuminate leaf apices, leaves always lacking a dense indumentum on abaxial surface, con- eon 266 sistent absence of multicellular gland-headed hairs, often shorter racemes, subapical placentae, and longer seeds. Although superficially similar, the two taxa probably are not closely related. Noteworthy in this regard, is the acenta position in the two species, t.e., subapical in A. — difference in p bolrviensis and more or less central in A. paraguayensis. ACKNOWLEDGMENTS The authors thank the curators of the herbaria from which specimens have been borrowed for this study. We also thank O. Ahumada and A. Rothman for their assistance during the second author's field trip to the Sierra de Zapla, and the Departamento de Biologia (UNS) for financial aid for that trip. REFERENCES CABRERA, A. L. 1983. Ericaceae, Pp. 2— 13, in A. L. Cabrera (ed.) Flora de la Provincia de ee part 8. Coleccién Crentifica del INTA, Buenos Aires GONZALEZ V., L. M. 1989. Hallazgo de una nueva especie de Avavisia (Ericaceae) en oe ‘Mexico, Acta Boranica Mexicana 5: 13-17. JUDD, S. 1979. Generic relationships in the Andromedeae (Ericaceae). J. Arnold Arbor. 60: 477 —903. 1984. A taxonomic revision of the American species of Agarista (Ericaceae). J. Arnold Arbor. 65: 255 — 342. LEGNAME, P. 1978. Una nueva especie de Gomidesia y tres nuevas citas para la flora Argentina. Lilloa 35: 79 MAYR, E. 1969. eee of ne Zoology. McGraw-Hill Book Co., New York. A NEW STATUS FOR QUERCUS SHUMARDII VAR. ACERIFOLIA (FAGACEAE) NICK STOY NOFF Glenbard East High School Lombard, IL 60148, U.S.A. WILLIAM J. HESS The Morton Arboretum Lisle, IL 60532, U.S.A. ABSTRACT Quercus shumardit Buck. var. acerifolia Palmer is elevated to species Q. acerifolia. It 1s restricted to the north-facing bluffs of Magazine Mountrain, Logan County, Arkansas. A comparison of certain morphological features with Q. shamardii sensu lato is made. Quercus shumardit Buckl. var. acerifolia Palmer (Maple-leaf Oak) was described in 1927 from specimens that E. J. Palmer collected in 1923 from Magazine Mountain, Arkansas. In the interim, no additional localities for this taxon have been discovered. It was included in the Report on Endange- red and Threatened Plant Species of the United States (United States Fish and Wildlife Service 1975) in the category of Threatened species, and more recently in Category 2 (possible listing may be appropriate, but sufficient data not available to support listing now) of the Federal Register of En- dangered and Threatened Wildlife and Plants (1985) by the United States Fish and Wildlife Service. The single known locality for the Maple-leaf Oak occurs on land within the Ozark National Forest. The Ozark National Forest has issued a special use permit to Arkansas Department of Parks and Tourism for development of a state park on Magazine Mountain. Ownership of the land in the even- tuality of park development will remain with the United States Forest Service. An Environmental Impact Study (EIS) : determine the feasibility of park development has been initiated and will be completed in 1991. Significant elements of flora and fauna on the mountain will be important factors in the selection of alternatives for park development in the EIS. Quercus shumardit var. acerifolia is clearly related to typical Q. shumardi1. Quercus shumardii sensu lato has also included other varieties such as schneckii (Britt.) Sarg. and texana Buckley sensv Trelease (= Q. buckley: Nixon and Dorr). We have visited the type locality for var. schnecki/) along the bottom- land of the Wabash River in [linois. The key morphological feature Sipa 14(2):267 — 271. 1990. 268 separating var. shumardi from vat. schnecki7 is the shape of the acorn cupule (saucer-shape = var. shumardir, turbinate = var. schneckii). Trees with both types of acorn cupules occur at this locality. At this time, because the key morphological characteristic separating the two varieties and bottomland habitat (= var. shvmardii) versus upland habitat (= var. schnecki7) do not hold up, we do not recognize a distinction between these two taxa. Two other taxa, Q. texana sensu Dorr and Nixon (1985) (= Q. nuttallii Palmer) and possibly QO. georgiana M.A. Curtis make a part of the Q. shamardii complex. It is not the purpose of this paper to address the relationships of these taxa, although we are studying them for a later report. Stoynoff and Hess have accumulated data to warrant raising Quercus shumardii vat. acerifolia to species status. Petzold and Kirchner (1864) were the first to use Q. acerifolia, but as a synonym of Q. rubra and without a description. Consequently, it was not a valid publication and Q. acerifolia is not a later homonym and available for the oak from Magazine Mountain. We are still in the process of gathering and analyzing data (especially from seedling stock), which will form the basis of comparative studies. Because of the impending EIS reviews, we believe it is important to propose the change in status now, rather than waiting for full confirmation. This may well be the most rare species of oak known and deserves very special preservation efforts and status. | QUERCUS ACERIFOLIA (Palmer) Stoynoff & Hess, stat. nov. — Basionym: Quercus shumardit Buckl. var. acerifolia Palmer, J. Arnold Arbor. 8:24—55. 1927, non cas ca Petzold & Kirchner, nom. nud. Arbor. Muscav. p. 656. 1864. Tyee: UNITED STATES. Arkansas: Logan Co.: rocky Gans cop of ee Mountain. 8 Oct 1924, Palmer 26434 UsoTyPE: ore MOR!). Table | is a comparison of certain morphological features of Quercus aceri- folia with its closest relative Q. shumardii. Maple-leaf Oak is a shrub or small tree to five (occasionally to 15) m tall. Typically, several ascending stems originate near the base at or below ground level. The leaves are broa- der than long and quite distinct from typical Q. shumardii leaves, which are longer than broad (Fig. 1). It has acorns (nuts only) that rarely are more than 17 mm long, 1/4 to 1/3 smaller than those of Q. shumardit (Fig. 2). Quercus acerifolia grows on the north-facing bluffs of Magazine Mountain at an elevation of 2600 feet. There are two populations, one at Brown Spr- ing and the other between 1.3 and 2.0 km west along the cliffs. They occur li tO rifolia and Q. shun ad hi —E. Q. acerifolia by re 7 oe off rom agazine Mountain, oe in © soe ae —J.Q.5 has Clay , Hess se Stoynoff 6433 i. *0. SSee, a la: 6500 _ She ea ec nia, Hess 0514. UL. jaiea Co. “illinois, a and aie 6436. J. ic Co., Arkansas, Hess al Stoynoff 440. 269 270 Taste 1. Comparisons of certain morphological features of the Quercus shumardii complex. Feature Q. acertfolta Q. shumardit Habit Shrub or small tree, 3-6 Tree, 15 — 30 (-40) m tall, (-15) m tall, multiple stems single stem Branch (yr 1) color Brown Brown pubescence Stellate, glabrescent Stellace, glabrescent Leaf habit Wider than long Longer than wide length 7— 14 cm 8— 17 (-20) cm width 8— 15 (-18.5) cm 6— 15 (-16) cm length/width x 0.81 (7 = 39) 1.22 (9 = 83) # lobes 5 (-7 5 —7 (-9) Bud color a dark brown, lower scales Apex brown, lower scales tan, eae margins hyaline margins pubescence Gi tab Glabrous Acorn cupule depth (2-) 3-5 (- ° (4-) 6-12 mm width (12.4-) 14 at a) mm (16.5-) 18 — 26 (-28) mm nu length 13— 17 (-17.5) mm (16-) 18 — 26 (-29) mm Distribution Logan Co., a from oy sto F TX, er KS. mostly within 75 m of the 65 m high bluffs on level ground or, more commonly, on gentle slopes (in what would have been a savannah-like zone) and along the rocky rim. ACKNOWLEDGEMENTS We wish to thank Marion T. Hall for his critical review of this manu- script and support of this study. Richard Jensen, Kevin Nixon, and Gary Tucker made several suggestions and improvements to the early drafts and we are indebted to them. We are indebted to Paul Manos for providing us with a photocopy from the library at the Bailey Hortorium, Cornell University, of the reference by Petzold and Kirchner. Nancy Hart-Stieber, staff artist at the Morton Arboretum, consulted on the illustrations and Elaine Fairbanks drew the leaf and acorn outlines. We thank the United States Fish and Wildlife Service for partial support of the field work under a Cooperative Agreement No. 14-16-004-89-940. REFERENCES DOR : L. J. and K. C. NIXON. 1985. a. 7 the oak (Quercus) taxa described by S. we kley (1809 — 1884). Taxon 34:211— PAL me R, 1. J. 1927. On Nuttall’s trail tirouah an J. Arnold Arbor. 8:24—55. 271 (y vi8ee. Se Nes ras Sa a 4 Y V MN F G H | Zi | J . Acorn nut and cupule outlines of Quercus acertfolta and Q. ). shumardit. A-E. Q. eee gelled . van der Linden and Hess from Logan Co., Arkansas. —J.Q. shumardi. FOG. | Woods, Wabash Co., Illinois, Hess and Stoyno/f 6503. HH. Clay , linois, Hess and Stoynoff 5.1. Shenandoah Co., Visine Hess O514. J. Misstssippt Co. Eee Hess and Stoynoff 0509. PETZOLD, E. C. and G. KIRCHNER. 1864. Arb. Muscav. 830 pp. UNITED STATES FISH AND WILDLIFE E SERVICE. 1975. Federal register of threatened or endangered fauna or ae review of status of vascular plants and determination of “critical habicat”. Vol. 40. No. 127 Federal register of endangered and threatened wildlife and plants; review of plant taxa for listing as endangered or threatened species. Part IV. Vol. O. No. 188. ANNOTATED CHECKLIST OF NEW MEXICAN CONVOLVULACEAE DANIEL E AUSTIN' Department of Botany Arizona State University Tempe, AZ 85287, U.S.A. ABSTRACT pecimens examined in 14 herbaria resul lred in the identification of 25 species of Con- aoe for the state of New Mexico. The list includes three species not recorded in the recent state flora, and makes nomenclatural changes in five others to bring them in accord with current literature. A lectotype is chosen for 1. mexicana. RESUMEN Un revision de las muestras de catorce herbarios ha resultado en la identificacion de veinticinco especies de Convolvulaceas para cl estado de New Mexico. La lista incluye tres especies nuevemente reportados en el estado, y se realiza cambios de nomenclatura en cinco de las otras para ponerlas en acuerdo con la literatura moderna. Se selecciona un lectotipo para Ipomoea mexicana. Although an updated flora of New Mexico was recently published (Martin and Hutchins 1981), there have been subsequent additions to the state (McDonald 1984, Spellenberg et al. 1986). More species are added as a result of studies of southwestern members of the family (Austin 1990b; in prep.). In addition, some of the names used by Martin and Hutchins (1981) are correcte In the following list, distribution data are provided for species on a county-by-county basis, and comments are made concerning the biogeo- graphic relationships of each in the southwestern United States. Basically, the flora consists of species derived from three sources: the Great Plains, Mexico and Meso-America, and introduced weeds This list includes three species not included by Martin and Hutchins (1981), i.e., Ipomoea dumetorum, 1. plummerae and 1. pubescens, and changes five names used by the latter authors. Twenty-five species are now documented for the state. This number is similar to Arizona (30 species, Austin, unpubl.) but small compared with those found in Texas (58 species, Correll and Johnston 1970). ele inent address, Department of Biological Sctences, F lorida Atlantic University, Boca Raton, FL 3431 Sipa 14(2):273 — 286. 1990. 274 KEY TO GENERA sags bases obtuse to acute. Styles 2; stigmas 2, globose; leaves elliptic to lanceolate or ovate- lanceolate; flowers salverform, 0.5 —0.65 cm long ..............000-. Cressa 2. Styles 2; stigmas 4, linear to club-shaped; iene ovate to almost linear; lowers rotate, funnelform or salverform, 5—22 mm long ........... Evolvulus |. Leaf bases truncate, cordate to hastate. 3. Leaves reniform; flowers mostly green, less than | cm wide ......... Dichondra 3. Leaves variable, but not reniform; flowers white or colored other than green, mostly over 1 cm wide 4. Flowers white, with or Pion tinges of lavender to pink on limb. 5. Calyx usually enclosed by 2 foliaceous bracts (not in C. longipes corolla funnelform, 3—6 cm long; stigmas oblong, flattened .. Ca/ystegia 5. ie not saat the bracts scalelike; corolla either campanu- late, eee funnelform, funnelform or salverform, (0.5-)1 — Me teas SUDU IAG scare ate eoe bene eens Convolvulus 4. Flowers lavender, blue, red or white with a purple to purple-red NOSE Ay cs shana wd vat on ates ws ue Eb oa Reed be wae Cae es [pomoea KEY TO CALYSTEGIA : Leaf bases markedly 2-angled; calyx 15-30 mm long ...... C. sik ? angulata . Leaf bases cordate to subsagittate, the lobes rounded; calyx 10 — 12 POOR Nowe ats + fied eee sae de Huang eee ace tle as wc erie ae hauesios C. macount CALYSTEGIA R. Brown HeEpcr BINDWEED 1. CALYSTEGIA MACOUNH (Greene) Brummitt, Ann. Missouri Bot. Gard. 52: 215. 1965. — Tyee: CANADA. SASKATCHEWAN: Assiniboia, Milk River, Aug 1895, Macounn (not seen). Calystegia interior House, Bull. Torrey Bot. Club 32: 140. 1905. — Type: COLO- R ca. Fe. Collins, 19 Jun 1896, Crandell 1625 (NY!). Apparently rare in New Mexico, this basically Great Plains species ex- tends south into this state. Although the taxonomy of our native taxa is complex, the group has been discussed by Brummitt (1980). vecimen examined. San Miguel Co.: Las Vegas, Soldiers Camp, 14 Jun 1927, Bro. Agios 18720 (US). 2. CALysTeGia sepium (L.) R.Br. or ANGULATA Brummitt, Kew Bull. 35(2):328. 1980. — Tyee: IDAHO. Canyon Co.: Macbride 318 (NY!). Calystegia sepium (L.) R. Br. var. ee (Brummitt) N. aes in A. Cronquist et al., Intermountain Fl. Vasc. Pl. Intermountain West, U.S.A. 4:77 1984. This North American subspecies reaches its limits in the southwestern United States. Numerous people have misinterpeted this taxon, and the recent Utah flora (Welsh et al. 1987) records it under Calystegia sepinm with the incorrect statement that it is an introduced European plant. These Zio plants are easily confused with the Great Plains taxon C. sylvatica spp. fraterniflora (Mackenzie & Bush) Brummitt, as was done by Tryon (1939), Correll and Correll (1972) and Lehr (1978). — specimens examined. Colfax Co.: Clarke 16131 (UNM). Dona Ana 19 Jul 1902, ee n. (ARIZ, NMC); Wooton & Standley 3353 (ARIZ, NMC). Rio Arch Co.: Jul | Newberry s.n. (US). San Juan Co.: Standley 7031 (US). San Miguel . Jul 1881, ee Sn. (NY). EY TO CONVOLVULUS 1. Leaves almost as broad as long; calyx 3 — 5 mm long; perennials from deep creeping root, forming late patches i. sawun dee saw ae eee eee tae C. arvensis : Leaves usually much longer than broad; calyx 6 — 12 mm long; perennials rom taproot, sometimes divided at apex but nor forming large, eis patches —_ SPopads a YE ae ae a esis eee Skene oe one eee eee Thouars 1. CGHitans CONVOLVULUS L. BiInpweep 3. CONVOLVULUS ARVENSIS L., Sp. Pl. 153. 1753. — Type: SWEDEN: speci- men 218.1 (LINN, microfiche!). This European introduction has become one of the most widely distribu- ted members of the family in North America. It is a problem weed in cotton and corn fields. Representative specimens examined. Bernalillo Co.: 16 Jul 1945, Reed s.n. (UNM). Catron Co.: in 1964, James s.v. (UNM). Chaves Co.: Bohrer 1975 (ARIZ). Colfax Co.: Jun 1979, Higgins and Campbell s.n. (UNM). ee Co.: | Oct 1907, eee sn. (NMC Dona Ana Co.: Anderson 13 (NMC). Eddy Co.: 18 May 1940, Hershey s.n. (NMC). ea : Hess 2065 (ARIZ). Guadalupe Co. : San 146 (ARIZ). Hidalgo Castetter oe (UNM). Lea Co.: Pearce 2675 (ARIZ). Lincoln Co.: Hutchins 3505 (UNM). McKinley Co.: Ne/son 7341 (UNM). Otero Co.: 24 May 1970, Todsen s.n. (NMC). Quay Co.: Waldrop 37 (UNM). Rio Arriba Co.: Baker 530 (ARIZ). Roosevelt Co.: Castetter 10720 (UNM). San Juan Co.: Levin 408 (AR IZ). San Miguel Co.: 20 Jul 1965, Broeske SM, eee Sandoval Co.: Ne/son 7342 (UNM). Santa Fe Co.: Bartlett 63 (NMC). Sie Co.: Metcalfe 1186 (NMC, NY, UC). Soccoro Co.: Moeller 277 (NMC). Taos Co.: ter ee (UNM). Torrance one Bedker 1042 (UNM I : : a et - ). Valencia Co.: Riffle 1216 (UNM). 4. CONVOLVULUS EQUITANS Bentham, Pl. Hartweg. 16. 1839. — Tver: MEXICO: Hartweg 98 UX? not seen). Convolvilus incanus sensu auct., non Vahl. This tropical American species reaches its northern limit in Arizona Utah, New Mexico and Texas. Representative specimens examined: Catron Co.: Mu/ford 516 (NY). Chaves Co.: Earle & Earle 248 (NMC, NY, UC). Colfax Co.: Griffiths 55 37 (US). De Baca Co.: Dunn 1955 (UNM). Dona Ana Co.: 18 May 1936, Hershey s.n. (NMC). Eddy Co.: 3 Aug 1909, W ahs s.n. (NMC). Grant Co.: 22 Jun 1906, Wooton 5.x. (NMC). Guadalupe Co.: Clark 7344 (UNM). Harding Co.: Ward ut al, 81-244 (NMC). Hidalgo Co.: Cazzer 405 (ASU); Spellenbere & Spellenberg 3825 (ASU, TEX, NMC, NY). Lincoln Co.: Locke et al. G1 1-40 276 ee Luna Co.: Hershey 2043 (NMC). Otero Co.: Fletcher and Hageren 600 (UNM). : Castetter 10690 (UNM). San Miguel Co.: 1899, Cockere/l & Porter 5.n. (NMC). : Plowman and Kilham AP295 (GH). Sierra Co.: Spellenbere & Todsen 2539 23 Sep 1907, Hanson s.n. ua hae Co. (NMC, NY). Soccoro Co.: Fleetwood 10 (NMC). Union Co.: (NMC). Ward (1984) recorded the chromosome numbers of this species, based on his Harding County collection, as n = CRESSA — ALKALI WEED 5. Cressa TRUXILLENSIS Humboldc, Bonpland and Kunth, Nov. Gen Spe Pl. 33. o 19. — Tyee: PERU: Trujillo, Humboldt 4 ee 3727 (micrOficHE: B!; isorype: F!). Cressa cretica L. var. truxilensis GH.B.& K.) C hoisy in DeCandolle, Prod o: 140. 184 Cressa depressa Goodding, Bot. Gaz. 37:58. 1904. — Typr: NEVADA: Goodding 726 (UC!) Cressa tnsularis House, Bull. Torrey Bot. Club 33:315. 1906. — Type: MEXICO: Revillagigedo Isls, Barkelew 252 (US!, UC!). Cressa erecta Rydberg, Bull. Torrey Bot. Club 40:466. 1913. — Type: UTAH: Garrett 870 (NY!) Cresia minima Heller, Muhlenbergia 8:140. 1913. — Type: NEVADA: Heller and Ken- nedy 80034 (NY!). Cressa truxillensis H.B.& K. var. minima (Heller) Munz, Aliso » 1958 Cressa pumila Heller, Muhlenbergia 8:142. tab. 17. 1913. nomen nudum. Cressa vallicola Heller, Muhlenbergia 8: 140. tab. 17. 1913. — Type: CALIFORNIA: Heller 8936a UC!). Cressa truxiflensis H.B.& K. var. vallicola (Heller) Munz, Aliso 4:96. 1958. Throughout the North American range of this species (e and s California and se Oregon, e to Utah, w parts of Texas and Oklahoma) there is con- siderable variation that appears to be of minor taxonomic importance (cf. Austin 1990b). No New Mexican populations are thought to be worthy of a varietal name at this time. For example, plants which are erect have been ate referred to C. truxillensis; those which are prostrate are referred to depressa Goodding. Both of these growth forms have been found grow- ing in the same stand (Arizona: Yuma Co.: Austin and Austin 7586 ASU). Although the plants are locally common in coastal Sonora, they are infre quent to rare in Arizona, New Mexico and Texas. pees specimens ae Bernalillo Co.: i and — 7361 (UNM). Chaves Co.: Waterfall 4313 (ARIZ). Dona Ana Co.: 12 Jun 1892, Wooton s.n. (NMC) Eddy Co.: Castetter 10683 rte Otero Co.: 16 ae 1936, Ps som. (NMC), Soccoro Co.: Castetter s.n. (UNM. L0G619) — KEY TO DICHONDRA |. Plants with appressed, whitish or canescent pubescence; pedicels 4-6 mm long, recurved near their attachment to the stolon.................. D. argentea iT |. Plants not whitish or canescent; pedicels 5 — 13(-26) mm long, recurved NAG ESOAIV ES fama: 4 cde ike bia Oe a ae oa oe ee halle D. brachypoda DICHONDRA Forster PENNYWworRT 6. DicHONDRA ARGENTEA Willd., Hort. Berol. 297. t. 81.1806. — Type: COLOMBIA: Tolima near Honda, Bonpland (B?). These plants often grow on southwestern-facing rocky ridges in Dona Ana and Luna Counties. Although plants may be locally common, the species is infrequent in the state. The species occurs in New Mexico, Texas, and was found once in Arizona in 1931 (Harrison 8256 ARIZ). In Mexico it occurs from Chihuahua south to Chiapas; also found in Central and South America. epresentative specimens studied: De Baca Co.: 23 Oct 1904, Wooton s.n. (NMC); 25 7 1894, Wooton s.n. (NMC). Dona Ana Co.: Austin & Austin 7637 (ASU). Grant Co.: Knight 2725 (UNM). Harding Co.: Wooton 5.2. (UNM 18050). Luna Co.: Goodding 3189 (NMC). 7. DICHONDRA BRACHYPODA Wooton and Standley, Contr. U.S. Nat. Herb. 16:160. 1913. — Tver: NEW MEXICO. Dona Ana Co > Organ Mountains, Filmore Canyon, 23 Sep 1906, Wooton & Standley s.n. (US!). This species of the Mexican-U.S. border is known from Arizona, New Mexico and Texas. In Mexico it has been found from Chihuahua to Oaxaca. Representative specimens studied: De Baca Co.: 1890, : ooton s.n. (US). Eddy Co. Jul 1909, Maes n. (NMC). Grant Co.: Barneby 2541 . Hidalgo Co.: ne ee es 5318 (NMC, NY); aaa G Salleabens 6318 aie NY). Sierra Co.: Metcalfe 377 (GH, nes NY — KEY TO EVOLVULUS mau developed, me or shorter than the ae cave 2. Sepals densely pilose, 2—2.5 mm long; corolla — 10mm wide; leaves elliptic, ovate or sons to lanceolate; stems with appressed- pilose and long aie EL ICO OMICS iy encores een Glen Ans ene eas E. alsinoides 2. Sepals pilose to tomentose, 3 — . mm long; corolla (10-)12 — 22 mm wide; leaves lanceolate to Nigar amee late. stems appressed pilose to tomentose, rarely with spreading trichomes.............. 00000000 E. laetus 1. Peduncle absent or extremely short, always fe than subtending leaves. 3. Sepals lanceolate to narrow-lanceolate, 4-5 mm_ long, spreading PCC eae Cu enaes ans eee eee ea oneae eee eave weees Li. nuttallianns 3. Sepals oblong-lanceolate, 3—5 mm long, appressed-pilose .......... E. sericeus EVOLVULUS 1: 8. Evo_vuLus aAtsinoibes L. var. ANGUSTIFOLIA Torr., Bot. Mex. Bound. 150. 1858. — Tyee: TEXAS. Brewster Co.: near the Grand Canyon of the Rio Grande, August, Parry (not found in CM, GH, ISC, MO, NY, PH, US or YU). on 273 ace —— L. var. acapulcensis pee ) van Sige Meded. Bot. Mus. b. Riks Univ. Utrecht 14:34. 1934. — Type: MEXICO. Guerrero: near le Willdenow 6128 (B). — This species is pantropical, and has been divided into a large number of varieties. The variety that occurs in Arizona, New Mexico, Texas and Mexico is var. angustifolia Torrey (cf. Austin 1990a). Nearby in Texas ts the var. hirticaulis Torrey. While this Sonoran Desert variety seems to be rare in New Mexico, it is frequent in southern Arizona. Representative specimens studied: Dona Ana Co.: Todsen 700802-3 (NMC). Hidalgo Co.: Castetter s.n. (UNM 16459). Luna Co.: Barneby 2485 (NY) 9. Evotvutus Larrus Gray, Proc. Amer. Acad. Arts 17:228. 1882. — Tyrer: ARIZONA: 1881, Pringle (F!, GH!, US!). Evolvulus arizonicus Gray var. /aetws (Gray) van Ooststroom, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 14:76. 1934 Evolonlus avizontenus Gray, Syn. Fl. N. Amer. 2, 1:218.1886. — Type: MEXICO. SONORA: sandy prairies, Sep 1857, Thurber 1023 (GH!); see Austin (1990a) on complexities of typification, Martin & Hutchins (1981: 1557) separated these two named varieties on the basis of stem and leaf pubescence: short and appressed in var. arizonicus, and both short-appressed and long and spreading in var. /aetus. These traits do not allow separation of the named taxa across the geographic range of the species (cf. Austin 1990a). Representative specimens studied: a Ana Co.: 19 Jul 1901, Wooton s.n. (NMC). Grant Co.: Moore 103 (ARIZ). aie C 0.: Spellenbere & Repass ve (NMC LO. EvotvuLus NUTTALLIANUS Roem. and Schult., Syst. Veg. 6:198. 1820. — Tyr: not seen. volt ulus pilosus Nute., Gen. N. Amer. Pl. 1:174. 1818, nom. tllegit. — Type: on the banks of the Missourt, Nvfia// (not found). Evolt sas oreophilus — Greene, eafl. ot. Observ. and — Crit. 1:15]. 3— 1906. — Typr: NEW MEXICO. Sterra Co.: Metcalfe 1228 (NMC!, NY!, Uc! US). Perry (1939) originally pointed out that the Roemer & Schultes name has priority over the Nuttall name. This is a Great Plains species that reaches its limits in the southwestern United States. Xepresentative specimens studied: Chaves Co.: ee 9155 (NY). Colfax Co.: Standley 6291 (US). Dona Ana Co.: Wooton 7 128 NMC . Eddy Co.: 1 Aug 1909, W VOLUN SR. Mc ). Grant Co.: Menaliesa 4. (UNM 18728). een Co.: Tichaikowsky 3 ( ee Co.: Spellenbery et al. 6041 (NMC, NY). Hidalgo Co.: Moeller & a son 4 rae C). Lea Co.: Marley et al. 1458 (UNM). Lincoln Co.: Gordon and Dunn S82 (UNM). Wee Co.: Worthington 13709 (NY). Otero Co.: Fletcher and Hageren 644 (UNM). Quay Co.: collector unknown 7358 (UNM), Eggleston 20121 (GH). Roosevelt FS 279 Co.: 17 Aug 1909, Wooton s.n. (NMC). San eee Co.: Loswre 350 (ARIZ). San Miguel Co.: Broeke MOOG (UNM). Sandoval Co.: Castetter 7364 (UNM). Santa Fe Co.: McKinley S4 (UNM). Sierra Co.: Todsen 55267 (NMC). nee Co.: Herrick 711 (NMC). Union Co.: Bartlett 234 (NMC) Some specimens of this species and E. sericevs are difficult to separate. Sepal pubescence and shape will usually allow their separation. II. Bae SERICEUS ae Prodr. Veg. Ind. Occ. 55. 1788. — Type: JAMAICA: Swartz (M, Evolvulus wilcoxiana House, Bull. Torrey Bot. Club 33:315. 1906. — Tver: ARI- /, A: Wilcox 96 (US!). Evolvulus serweus var. discolor (Benth.) Gray, Syn. Fl. N. Amer 2, 1: 436. 1886. — Tver: MEXICO: aus Lagos (Jalisco) and Aguascalientes (Aguascali- entes), Hartwee 20 (K, L, Martin & Hutchins (1981: 1557) separate these two taxa on the single basis of leaf pubescence: upper surface pubescent in var. sericeous; upper leaf surface glabrate and green in var. dsco/or. Both forms may be found within the same population of the plants, thus only one taxon seems worthy of recognition. (cf. Austin 1982). This its a tropical American species that reaches its northern limits in the United States, tn Arizona, New Mexico, Texas, Georgia and also Florida. Representative specimens studied: Catron Co.: Shelton 115 (NY). Curry Co.: Clark 5.1. (UNM 5990). Dona Ana Co.: 29 Aug 1894, Wooton s.n. (NMC). Eddy Co.: Marley 5.1. (GH, UNM 74839). Grant Co.: Metcalfe 100 (GH, NMC, NY, UC) ). Hidalgo Co.: Morr 109 (NMC). Luna Co.: Castetter 10692 (UNM). Ore Co. Meents & Moir 20 (NMC) Roosevelt Co.: Castetter 10693 (UNM). San Miguel Co.: Cockerel/ s.r. in 1899 ( NMC). Sierra Co.: Metcalfe 1259 (NMC, NY). Soccoro Co.: se 7719 (NMC). KEY TO /POMOEA l. ia res pedatisect, less often sagittate. 2. Flowers salverform, 3.5 — 10 cm long, nocturnal or diurnal... 2.2... I. tenuiloba 2. Flowers funnelformn. mostly less than 3 cm long, diurnal. 3. Sepals hirsuce; corolla 2.5 —3 cm long ....... 0.000.000 c ee eee I. leptotoma 2 Sy = > i 3. Sepals glabrous or muricate; corolla | — 3 cm long. A. Calyx olabrous; corolla. = 1,5 Cm 1OnGes wns cade aen agen d |. costellata A 2 4. Calyx muricate-tuberculate; corolla 2— 3 cm long. 5. Sepals 5-6 mm long; peduncle plus pedicel about 5 — 10 (-14) mm long; tuber elongate... .....0000..0...0 0000. L. capillacea 5. Sepals 7 —9 mm long; peduncles plus pedicels about 14 — 18 (-22) mm long; tuber globose to subglobose ........... 1. plummerae 1. Leaves simple to lobed or toothed, often cordate. Oi PEREGE MGR OS heater af ingen csavtorah 2. & dare eta eo oe See eee L. leptophylla 6. Climbing to eae vines 7. Corollas 2— 2.6 ¢ slonkt scarlet, orange or yellow ............ Ll. crtstulata 7. Corollas 4-15 cm long, lavender to white or purple. 280 8. Pedicels and peduncles glabrous or with appressed small tri- chomes. 0. Sepals tianewar wus wea ches oe seeon eos wn hes |. cardiophylla Os Seals Ota adecassueeniaddieoseteuis teu eseee sss 1. dumetorum . Pedicels and peduncles with spreading, ascending or reflexed i oo es apices acute to obtus Sepals 8 - 15 mm lone. ovate- eye to elliptic and founded at he fe corolla 2.5 —4.3 (-5.0) cm ea . 1. purpurea 11. Sepals 15-28 mm long, ovate-attenuate to lanceolat attenuate and truncate at the base; corolla 4-8 cm aa ge eer een oe ee ee ee oe eee I. pubescens 10. Sepal apices acuminate to long-acuminate 12. Sepals 9— 12 mm long, ore eter: and not con- spicuously dilated at the base; corolla 1.6 — 2.5 cm long sunlit mate tern eye manatee © GA eae cdl Wie g Bone ee eee anes I. barbatisepala 12. Sepals 12 — 28 mm long, lanceolate to ovate-lanceolate, conspicuously ioe at the base; corolla 2— 10 cm long. 13. Sepals 12 — m long, lanceolate; corolla 2.0 — 3.7 (4 4.5) cm oe AMMUAl s deaeuna cei as Sees I. hederacea 13. Sepals 15 — 28 mm ae ovate-lanceolate to narrowly lanceolate; corolla 6— 10 cm long; perennial .... 1. /indheimert IPOMOEA LL. Morninc GLory 12. [POMOEA BARBATISEPALA Gray, Syn. Fl. N. Amer. 2, 1:212. 6. — Type: TEXAS: Wright 507 (GH!, US!). Some specimens are difficult to separate from the closely allied J. hederacea Jacq. The species, however, was originally endemic to the southeastern U.S., el. barbatisepala is a Mexican species on the margin of its range in . New Mexico and Texas. Representative specimen studied: Eddy Co.: Clark s.n. (UNM 4877). Luna Co.: Todsen 164 (NMC). 13. Tpomora capittacea (H.B. & K.) G. Don, Gen. Syst. 4:267 1838. — Typr: COLOMBIA: Bonpland (microfiche!). Ipomoea muricata Cav., Icones Pl. 5:52. pl. 478. f. 2. 1794, non L. (1763), non Jacq. 789 This is an American species ranging from Arizona and New Mexico through Mexico and Central America to South America. Representative specimens studied: Catron Co.: Fletcher 820 (UNM). Grant Co.: Rusby 301 (NY). Lincoln Co.: Earle & Earle 492 (NY). Sierra Co.: Knight 2199 (UNM). 14. IPOMOFA CARDIOPHYLLA Gray, Syn. Fl. N. Amer. 2, 1:213. 1886. — Typr: TEXAS: Wright 1314 (GH!). 281 The type was collected in Texas (Hudspeth Co., Hueco Mts., E of El Paso. 13 Oct 1849) where it was rediscovered by McDonald (1982). This species was most recently relocated by Ms. Katie Skaggs, naturalist on the A. B. Cox Nature Conservancy land in the Organ Mountains near Las Cruces. The species is easily confused with I. parasitica (H.B.K.) G. Don and I, tricolor Cav. of Mexico, Meso-America and South America. Representative specimens studied: Dona Ana Co.: 28 Sep 1902, Gaee n. (NM Sep 1980, Worthington 6655 (TEX), 11 pe 1980, 6746 (TEX); 23 Oct 1975, Von L ie (UNM). Grant Co.: Zimmerman and Zimmerman 2006 (SNM fide ee 1982: 261). 1S. IPOMOEA COSTELLATA Tors ‘ai Mex. Bound. 149. 1859. — Tyee: TEXAS: Wright 505 (GH!, US Ipomoea futilis A. Nelson, Univ. Wyoming Publ. Sci. 1(3):65. 1924. — Tyrer: ARI- ZONA: Hanson 1016 (RS) This annual desert species ts similar to and undoubtedly related to J. leptotoma. It occurs in Arizona, New Mexico and Mexico (Baja California, Chihuahua, and Sonora, south to Chiapas and Veracruz). Representative specimens studied: ey Co meee 515 (UNM). Catron Co.: ake 9151 (UNM). Chaves Co.: Earle & Earle 331 (NMC, NY). noe Pte 1902, Wooton s.n. (ARIZ, NMC). . see ‘Spelenber etal, 8270 (N a Co.: Spellenberg & Spellenbery 3852 (NMC, NY, TEX). Lincoln oe rae 2590 (UNM). Luna Co.: W oC 13742 (NY). = Miguel Co.: Hi// 12258 (GH). Soccoro Co.: Metcalfe 766 (ARIZ, C). 16. Ipomoea crisrutara H. Hall., Med. Rijksherb. Leiden 46:20. 1922. — Tyee: MEXICO: based on syntypes including Bowrgean 1061 ne DC!). A nom. nov. for Quamoclit gracilis H. Hall, Bull. Herb. Boiss. 7:416. 1899. [pomoea coccinea auct., non L. Martin & Hutchins (1981: 1560), Wooten & Standley (1915), Tides- trom & Kittell (1942), Kearney & Peebles (195 1) and Shreve & Wiggins (1964) have applied two incorrect names to these populations: [. coccinea var. coccinea and I. coccinea var. hederifolia. In fact, these populations represent Ipomoea cristulata, a largely Mexican (Baja California, Chihuahua, and Sonora south to Michoacan and San Luis Potosi) xerophyte that extends northward into Arizona, New Mexico, Texas and into the Great Plains. [pomoea cristulata 1s easily distinguished from the ornet two taxa, which are best considered species, by its sepals which are 3 — 4 mm long, and (when in fruit) by reflexed peduncles. The other two species have smaller sepals and erect fruits. [pomoea coccinea L. 1s a southeastern United States endemic; [. hederifolia 1s a tropical American species now widely spread and natura- lized in the wet tropics of the world. 282 Ward (1984) reported the Hidalgo County collection with a chromo- some number of n = — Representative specimens studied: ys Co.: Hutchins 9843 (UNM). Dona Ana Co.: 16 Aug 1895, Wooton s.n. (NMC). t Co.: Beckworth 150 Vea Hidalgo Co.: Spellenberg 3834 (NMC). Lincoln Co. : Y er 1899, Turner s.n. Rio Arriba Co. : Castetter 100686 (UNM). San Miguel Co.: Standley 5272 (NMC). ee 1 Co.: Yarnell 128 (UNM). Santa Fe Co.: Bartlett 65 (NMC). Sierra Co.: 27 Jul 1904, Metcalfe s.n. (NMC). Soccoro Co.: Metcalfe 825 (NMC). Torrance Co.: Fletcher 5612 (UNM). Valencia Co.: McCallum 619 (UNM). 2 17. lPomMoEA DUMETORUM Willd. ex Roem. & Schult., Syst. Veg. 4:789. 1819. — Tyee: COLOMBIA or ECUADOR: without locality (B). Martin & Hutchins (1981) do not include this in their flora. This Mexican, Meso-American and South American species was identified and relocated by McDonald (1982, 1984) in New Mexico and Texas. McDonald (1984) cites specimens in addition to those listed here. Dona Ana Co.: a Mes., McDonald 140 TEX, not seen). Lincoln Co.: White Mountains, alt. 7400 ft, 25 Aug 1907, Wooton & Standley s.n. (NMC, US), White Mts., ale. 2500 ia (MO, not s Because the species has been so rarely collected in the U.S.A., additional specimens will be cited. TEXAS. Jeff Davis Co.: Mt. Livermore, alt. 2700 m., Warnock 23008 (SR, not seen); Davis Mts., Madera Canyon, near Livermore, Hinckley s.n. (ARIZ). 18. [POMOEA HEDERACEA Jacq., Collect. Bot. 1:124. 1786. — Teer: Based on Dillenius, Hort. Elth. t. 80, fig. 92 (plate selected as lectotype by Verdcourt, 1957). [pomoea desertorum House, Ann. N.Y. Acad. Sci. 18:203. 1908. — Type: ARIZONA: Thornber 29 (ARIZ!, NY!) Lpomoea hirsutula authors, pro parte, non Jacq. f. (1811). Martin & Hutchins (1981: 1560) separate both their 1. hederacea and I. hirsutula from 1. purpurea on the basis of leaf lobing. If the plants have en- tire leaves they are placed in I. purpurea. \f there are leaf lobes, they key to either of the other two species. Leaf lobing will not separate these plants under any circumstances (Elmore 1986); only characteristics of the sepals will separate them. The correct citation is 1. hederacea Jacq. because Jacquin proposed the name as a new species, not a transfer of a Linnaean name (cf. Austin 1986a). While the species is a common weed in cotton fields in Arizona, it may be rare in New Mexico. At least, it is seldom collected since | found only two collections. Specimens studied: Hidalgo Co.: 20 Aug 1955, Castetter 11350 (UNM). Luna Co.: 30 Aug 1895, Mutlford 1088 (NY). 283 19. [pOMOEA LepTOPHYLLA ‘Torrey in Fremont, First Rept. 94. 1845. — Tver: forks of the upper Platte to Laramie River, Fremont (HOLOTYPE: US!; PROBABLE ISOTYPE: NY!). [ consider this Great Plains species an indicator of where prairies former- ly existed in New Mexico. It reaches its southwestern limit in New Mexico. Representative specimens studied: Chaves Co.: Wagner and bance fe (UNM). epres Colfax Co.: Griffiths 2323 (US). Curry Co.: 18 Aug 1909, Hae sin, (N . De Baca .: Higgins 8659 (NY). Dona Ana Co.: Castetter 7375 (UNM). Eddy Co. ce & Repass 5255 (NMC). Harding Co.: Van Devender 84-377 ee Hidalgo Co.: herb. Chap- man s.n. (NY). Quay Co.: 29 Jul 1942, Suges s.n. (N Rio Arriba ae Perdue 6128 (U Sica: Co.: Secor 61 (TEX). San mies ; Co (TEX). Sandoval 2 Sep 1936, Hershey s.n. (NMC). Sar : Hitchcock et al. pan Torrance Co.: ee 7372 (UNM). Union Co.: sere 350 (NMC). 20. [POMOEA LEPTOTOMA Torr., Bot. Mex. Bound. 150. 1859. — Tver: MEXICO. Sonora: Thurber 977 (GH!). Lponoea leptotoma var. wootonts Kelso, Rhodora 39:151. 1937. — Type: ARIZONA: 10 > 1914, Wooton s.n. (US!). This northern Mexican species is frequent in parts of Arizona, but appar- ently rare in New Mexico. Specimens studied: Curry Co.: Whitehouse 5.n. (TEX). Hidalgo Co.: Castetter 9509 (UNM 21. IPOMORA LINDHEIMERI Gray, Syn. Fl. N. Amer. 2, 1:210. 1886. — Tver: TEXAS: Wright 508 (GH!, US!). This Mexican-U.S. border species is known from Texas, New Mexico and Arizona as well as Coahuila and Chihuahua in Mexico. oe specimens studied: Dona ss Co.: 19 Sep 1976, Todsen s.n. (NMC). Eddy Co.: 4 Aug 1905, Wooton s.n. (NMC); Starr & Sie ron ae Otero Co.: Gordon and Norris 552 (UNM). County Pialoen: Wrieht 1612 (NY 22. IPOMOEA PLUMMERAE Gray, Syn. Fl. N. Amer. 2, 1: suppl. 434. 6. — Tyre: ARIZONA: Lemmon 2839 (GH!) Ipomoea cuneifolia’ Gray, Proc. Amer. Acad. Arts 19:90. 1883, non Meisner (1869). — Type: ARIZC » Lemmon 2859 (F!, GH!, US!). Ipomoea evregia House, Torreya 6:124. 1906, nom. nov. for 1. cunetfolia Gray. The species was not recorded for the state by Wooton and Standley (1915), nor Martin & Hutchins (1981), although it was included with a query in Tidestrom and Kittell (1941). This is a Mexican (Coahuila, Chihuahua, Sonora) species reaching its northern limits in Arizona and New Mexico. 284 Representative specimens studied: Catron Co.: Fletcher 2762 (UNM). Grant Co.: Spellenberg et al. 5864 (NMC). Lincoln Co.: 5 Aug 1897, Wooton s.n. (NMC), Wooton 627 (NY). Otero Co.: 8 Aug 1899, Wooton s.n. (NMC). San Miguel Co.: in 1899, Cockerell & Porter s.n. (NMC). Sierra Co.: 24 Aug 1969, Todsen s.n. (NMC). Soccoro Co.: Metcalfe 271 (GH, NMC). 23. IPOMOEA PUBESCENS Lam., Encycl. Meth. Bot. 1:265. 1791. — Type: America, collector unknown (K!). Ipomoea heterophylla Orcega, Hort. Matr. Dec. 1:9. 1800. — Type: MEXICO: not seen. Ipomoea lindheimeri: Gray var. subintegra House, Ann. N.Y. Acad. Sci. 182196, 1908. — Type: ARIZONA: Lemmon 2835 (GH!, UC!). This species was not included by Martin and Hutchins (1981) although they had specimens of it misidentified as 1. /indhermerr in the UNM herbarium. This is a widespread American species that reaches its northern limit in the southwestern United States. Representative specimens studied: Dona Ana Co.: Knight 3415 (UNM). Eddy Bailey 721 (US). Hidalgo Co.: collector unknown 7367 (UNM). Luna Co.: ese Spellenberg 6620 (NMC [TPOMOEA PURPUREA (L.) Roth, Bot. Abh. 27. 1787. — Tyee:U.S.A. Dillenius, Hort. Elth. ¢. 84, fig. 97. 1732 (Lecrorype: chosen by Verdcourt 1963!). Ipomoea hirsutula Jacquin f., Eclog. Pl. Rar. 1:63. t. 44. 1811. — Type: no specimen found, (LECroTyPE: the ae chosen _ ere 1990). Ipomoea mexicana Gray, Sy ~N. Amer. 2(1): 210. 1886. — Type: NEW MEXICO AND ARIZONA. SS, on cae Gace no lectotype has been chosen (House 1908) ), the pene is here designat { —52.° C. Wright ee (LECTOI : GH); on same sheet 1s: hens a. > Fr. Huachuca, 1882. Lemmon 2838 (GH!). Since both collec- tions were probably on the sheet when Gray published the binomial, he surely examined both. He cited, however, only the collection by Wright; thus, 1t ts chosen as lectotype. Gray also cited a collection by Fendler which is in the GH as a sheet containing two collections: New Mexico. Plantae Nova- pe 1847. Fendler 062 (GH! Arizona. w/o loc. 1878, Dr. Load 152-A (GH!). Gray ale cited a collection by Thurber which has not been located in GH. —_ The key in Martin & Hutchins will cause the user to place the entire- leaved specimens of 1. purpurea here and the lobed-leaved specimens into J. hirsutula, here they are treated synonymously. This species is now pantropt- cal because of cultivation, but it was undoubtedly originally Mexican. It occurs in Arizona, New Mexico, and Texas and has been introduced and/or escaped in the Great Plains, the southeastern United States and the north- eastern United States. Representative specimens studied: Bernalillo Co.: Wagner 519 (UNM). Catron Co. : Hutchins 9121 (UNM). Chaves Co.: Earle 256 (TEX, NY). Dona Ana Co.: Worthington — 285 6635 ae Eddy Co.: Higgins 9233 (NY). Grant Co.: Barkley 14710 (TEX). Guada- 1 ‘0.: Tichaikowsky 387 (ARIZ). posers ae a and Martin 5768 (UNM). Tape Co.: Spellenberg & Spellenbere 3905 (A NMC, NY). Lincoln Co.: Wooton & eae 3631 (NMC). Luna Co.: Spellenberg & ie 622 27 (NMC, NY). - tero Co.: lard & Soreng 81-534 (NMC). San Miguel Co.: Standley 49406 ( NMC, NY). Sandoval - Dixon 4-348 (UNM). Santa Fe Co.: Bartlett 64 (NMC). Soccoro Co.: : Aug 1900, Wooton sm. (NMC). Torrance Co.: Bedkey 1542 (UNM). 25. [IPOMOEA TENUILOBA Torr., Bot. Mex. Bound. 148. 1859. — Tver: TEXAS: Bigelow (US!). Ipomoea lemmont Gray, Proc. Amer. Acad. Sci. 19:20. 1883. — Type: ARIZONA: Lemmon 2840 (GH!, US!). aoe tenniloba Torrey var. lemmonit (Gray) Yatskievych & Mason, Madrono 31:102. 1984. Since both varieties and intermediates occur in New Mexico, and they have been completely discussed by Yatskievych & Mason (1984), they wil not be discussed in detail here. _ Representative specimens studied: Eddy Co.: 2 Aug 1909, Wooton s.n. (NMC). Grant Co.: Wagner 3444 (UNM). Hidalgo Co.: 15 Sep 1980, Todsen 5.x. (NMC) EXCLUDED SPECIES se langle Benth. inge given by Martin & Hutchins (1981: 1562) includes the range of 1. shumardiana (Torrey) Shinners, a Great Plains species that is distinct. There is one specimen of 1. /ep- tophylla (Winhoff 508 ASU) misidentified as 1. longifolia. Ipomoea longifolia has not on verified for any part of the United States except southeastern Arizona (cf. Austin 1986b). ee pubescens Lind]. re species has been collected once (Santa Fe Co.: 10 Jun 1925, Bro. Benedict 128 US) a a garden. It was undoubtedly cultivated as the species is cultivated farther north in the Great Plains Ipomoea alba L There is a specimen collected in 1949 in feaan as (Castetter 7334 UNM). Although the sheet does not indicate that the specimen was cultivated, this was almost certainly the case. The species typically grows in swamps and other oak farther south in the tropics. Indeed, the species is now pantropical in the wet tropics because it has been introduced and cultivated from the New World. ACKNOWLEDGMENTS Thanks are extended to curators of herbaria at A, ASC, ASU, ARIZ, CAS, DES, GH, NMC, NY, UNM, TEX, UC and US. This research was carried out while the author was on sabbatical leave at Arizona State University. REFERENCES AUSTIN, D. E 1982. oe eels In: Z. Luces de Febres & J. A. Steyermark, eds. Flora de Venezucla. Vol. 286 —CSC«dL' BG. NNomee clare of thee Ipomoea nil complex (Convolvulaceae). Taxon 35:355— 358 a sa _ 198%6b. Moth pollinated Ipomoea longifolia (Convolvulaceae). Desert Plants 8(1):1 16. 1990a. Comments on southwestern United States Evolvulus L. and Tpomoed L. iGoaeatalics: Madrono 37(2): 124-132 1990b. Annotated checklist of Arizona Convolvulaceae. (in prepara- tion). —_______.. in prep. Convolvulaceae In: Vascular plants of Arizona. Univ. of Arizona Press, Tucson, BRUMMITT, R. K. 1980. Further new names in the genus Ca/ystegia (Convolvulaceae). Kew Bull. 35(2):327 — 334. CORRELL, D. S. and H. B. CORRELL. 1972. Aquatic and wetland plants of south- western United States. Environmental Protection Agency, Washington, D.C. and M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas. eee Research Foundation, Renn ,C.D. 5 ELMOR ode of esol and inheritance of leaf shape in Ipomoea Be + BAIT 359). HOUSE, H 1908. The North American species of Ipomoea. Ann. New York Acad. Sci. 18:181— a KEARNEY, T. H., R. H. PEEBLES, and collaborators. 1951. Arizona flora. Univ. Calif. Press, Berkele LEHR, J. H. 1978. aes of the flora of Arizona. Desert Botanical Garden, Phoenix. MARTIN, W. C. and C. R. HUTCHINS. 1981. A flora of New Mexico, Vol. 2, J. Cramer, Verlag, German MCDONALD, J. A. 1982. Fc aia of the Ipomoea tricolor complex (Con- volvulaceae). Ph.D. dissertation, University of Texas, Austin. —™".s«d'92 4 Pies dhmoran Pe ain an amphi-tropical disjunct morning story i in the southwest U.S. Sida — a ERRY, L. PE 939. cies pees an invalid name. nae 37109. SHREVE, E i I. L. WIGGINS, 1964. Vegetation a flora of the Sonoran Desert, Vol. 2. Stanford ae Stanford. SPELLENBERG, R., R. WORTHINGTON, P KNIGHT a R. FLETCHER. 1986. Additions to the flora of New Mexico. Sida 11(4):455 — TIDESTROM, I. and Sister T. the L. 1941. Flora of oe and New Mexico. Catholic University of America s, Washington, ee R.M. Jr. 1939. The ee and of C. sepium. Rhodora 415 — 423, waa. D. E. 1984. Chromosome counts from New Mexico and Mexico. Phytologia 36°95 60, WELSH, S. L., N. D. ATWOOD, L. C. HIGGINS and $. GOODRICH i 1957. Utah bes Cex Basin Nat. Memoir No. 9. Brigham Young Unive WOOTON, E. O. and P C. STANDLEY. 1915. Flora of New Mexico. a U.S. Natl. Herb. ee 14-519. YATSKIEVYCH, G. & C. T. MASON, Jr. 1984. A taxonomic study of leh a Torrey (C eo: with notes on related species. Madrono 3 1(2):1 DELETIONS AND RESTORATIONS IN THE FLORA OF SOUTHERN FLORIDA DANIEL B. WARD Department of Botany, University oe Florida Gainesville, FL 32611, U.S FRANK C. CRAIGHEAD _ yp 2 4 Naples, FL 33940, U.S.A. ABSTRACT The species listed in the Annotated Checklist of the Vascular Plants of Collier, Dade, and Monroe counties, Florida (1965) as occurring in the three southernmost counties of Florida have been compared with those species included in A Flora of Tropical Florida (1971, 197 76) and with the known flora of che area. Corrections are made by deletion of 62 species and six families reported to occur in South Florida, restoration of 18 species and one family once reported but later omitted, and assignment of 23 names that should have been included in synonymy in the later publication. The vascular flora of Florida is a fascinating one, varied and exotic beyond that of any other state in eastern North America. The abundance and novelty of this flora, and the economic and aesthetic interest in it by a rapidly expanding human population, has made welcome those too-few efforts to describe or merely to enumerate its plants. It is inevitable, given the demand for studies of the state’s flora, the paucity of experienced floristic botanists in the area, and the pressures upon them to make their information available, that preliminary listings and tentative identifications will be placed in print. Lack of time either in the field or in the herbarium, lack of access to literature or authoritatively named specimens, or lack of adequate understanding of the biological realities that keep plant distribution from being a random and wholly un- predictable event, all have doubtless contributed to the publication of errors. Such errors, once unequivocally placed in print, gain a life of their own, being copied and recopicd with ever increasing verisimilitude, and are suppressed only with great difficulty. No writer dealing with technical minutiae can be free of all error, and in most circumstances later commentators are perhaps best advised to make corrections gently by stating the facts accurately without specific mention of aberrant views. We do feel an obligation, however, where the general topic (plant identification and distribution in the state of Florida) 1s so 1n- timately tied to our professional experience and knowledge, and particu- larly where one of us by implication is responsible for the statements made, Sipa 14(2):287 — 304. 1990. 288 to document as clearly as we can, errors in this field with the hope that such documentation will inhibit their repetition in later publications. In 1965 the Fairchild Tropical Garden and the University of Miami Press released an Annotated Checklist of the Vascular Plants of Collier, Dade, and Monroe counties, Florida. Its authorship was given as Dr. Olga Lakela, University of South Florida, Tampa, and Dr. Frank C. Craighead, of the Everglades National Park and Fairchild Tropical Garden. This listing was welcomed in southern Florida and served as a standardized tabulation of the flora of these three counties until superceded by Robert W. Long & Olga Lakela’s A Flora of Tropical Florida, University of Miami Press, 1971. The Annotated Checklist contained 1,470 species of ferns, gymnosperms, monocotyledons, and dicotyledons, and for most gave the habitats, the counties, and in many cases the exact areas from which specimens had supposedly been collected or reported. Documentation was stated to be based on the herbaria of the Everglades National Park, the University of Miami, the University of South Florida, and on previously published records. In what remains as an inexplicable misunderstanding, the present second author's name was listed, following that of Dr. Lakela, as co-author. He was not. This publication was not his doing, and he did not request nor anticipate that his name be so credited. He did give of his time and inform- ation to Dr. Lakela during her trips to southern Florida, and he did give her free access to the herbarium of the Everglades National Park, of which he was then curator and to which he had largely contributed. It must be attri- buted to the generosity of Dr. Lakela that she so acknowledged this help. But it has left him in the uncomfortable position of being considered responsible for errors that he had no part in making, and even further of being aware that his tentative identifications, never intended to be presen- ted as definitive, are the source at least in part of statements in the Annota- ted Checklist that are patently false and denigrate the scholarship of its authors. | ‘My triend and co-author, Dr. Frank Cooper Craighead, died 15 May 1982. After his retirement as a U.S.D.A. entomologist, “Craig” became the unofficial bur high] y respected botanist of the Everglades National Park. His enthusiastic and perceptive field studies in this second career produced a number of botanical publications eae Onchids and Other Air Plants of the Everglades National Park (1963) and Trees of South Florida (19 Craig was not a man é a temper. It was at his urging chat I began this compilation in the early 1970s. Encouragement was also received from George Avery, a meticulous collector and field O a. oserver of ike flora of southern Florida. ve with the loss dicondt 1 death of those persons immediately | William Long, 21 July 1976; eae i Korhonen Lakela, 17 May 1980; George Newton Avery, ca. 12 July 1983), other tasks c preceder Now, with a resurgence of interest in the ei a Ponds and with state-wide floristic projects actively underway in Fol thassee, Gainesville, Tampa, and Miami, it seems time to make these notes available. — D.B. 289 The inadequacies of this preliminary annotated listing were apparent, and the authors of A Flora of Tropical Florida clearly attempted exculpation eon by disregarding this earlier production. Nowhere in the 962 pages of their 1971 book, neither in the list of “selected” references nor at any other appropriate point, is there mention that six years earlier one of them had authored a treatment of the plant species of the identical geographic area. But this earlier annotated listing cannot so simply be ignored. Copies of it abound, and are regularly cited by authors interested in the flora of southern Florida (Al-Shehbaz 1985; Austin 1980; Miasek 1978; Miller 197 1a, 1971b; Poppleton et al. 1977; Rogers 1984, 1985; Spongberg 1978: Webster 1967; etc.). Further, a peculiar stylistic feature of the P/ora has left the earlier Annotated Checklist still its essential companion. This ts the practice of A Flora of Tropical Florida, apparently unique in recent North American local floras, of omitting almost all exact statements of distribution in the treated area for almost all species. The Annotated Check- list, although its oo information may not be detailed to the degree desired, at least goes well beyond the larger book in specifying the counties from which collections have been seen or reports cited. This disregard of the earlier Annotated Checklist by the authors of A Flora of ‘Tropical Plorida has left in a botanical limbo the names that appeared in the listing but not in the later book. A contemporary reviewer (Gillis 1973) noted that such names exist and suggested that it is an obligation of floristic writers to account for previous names recorded (and previously un- challenged) for the area they treat. We support this policy most strongly. It seems particularly appropriate that an author who is aware of an error in his own work be the one to call attention to his earlier erroneous statement, thus most effectively remov- ing doubt as to its invalidity. But when a previous author has not taken the opportunity to do so, the mantle of responsibility falls more broadly on the botanical community, and those with information that might prevent error by a still later generation of writers would themselves appear obligated to make correction. It is in this spirit that we have undertaken the present task. The following listing is intended to be comprehensive only of the 103 names that appear in the 1965 Annotated Checklist that cannot be accounted for (in one way or another) in the 1971 A Flora of Tropical Florida or in its essentially identical 1976 “new edition.” Additional names used in the Annotated Checklist, perhaps two to three times as many as in the following list, are not used for species in the later publication but are recorded as synonyms under a name accepted in the I/ora or are otherwise un- ambiguously traceable 290) The vanished names may be accounted for in one of three ways. First, and most numerous, are those species reported on the basis of mis- identifications, where the error appears to have been detected and the species was correctly deleted from the later Flora. Sixty-two such species have been noted by us, and had the F/ora contained the customary account- ing of excluded species, these names would have been disposed of in that publication.’ These names constitute somewhat over 4.2% of the species enumerated by the Annotated Checklist. They include the sole representatives in South Florida (here defined, as in the two publications under discussion, as limi- ted to Collier, Dade, and Monroe counties) of eighteen genera and six families (Araliaceae, Marsileaceae, Nyssaceae, Punicaceae, Sparganiaceae, Zosteraceae). It must be made clear that additional species are included in the Annota- ted Checklist which we do not believe occur in South Florida. Our own un- derstanding of their ranges casts immediate doubt on the inclusion of such species as Cakile edentula, Carya floridana, Hypericum galioides, Juniperus silicicola, Oxalis stricta (= O. cymosa), Polygonum perstcarta, Prunella vulgaris, Sabatia campanulata, and Vinca minor, some of which we are not aware occur in Florida, much less in the southernmost counties covered by the Flora. But each of these names is included in the F/org (some with qualifications), and thus their tabulation falls outside the present scope of our study.’ ‘Teas worth comment that there is no standardized w ay of handling species that are ee for the { area under study but are believed inappropriate for inclusion. Three ese pe th diversity of treatment. Deam (Flora of Indiana 1940) prea his 707 excluded species into a single appendix. Steyerm: is (Flora of Missouri ies 3) es his aueoas species at the end of each genus. Voss (Michivan Flora 1972, 1985) inserted hts E cext of the species with which they were most related, 1 che names recorded in the Annotated Checklist, the presence of a name in A Flora of Tropical I lines a not senna | reflect the presence of that species in the flora of southern Florida. Persons Who LIV As only cursory review of this strange cee cannot eae. the tee number ot species listed therein that actually are unknown in South Florida, cithe as modern- di ay au ior he following names, ee with the names listed in the above text, are offered as a partial ed in A Flora of Tropical Florida that appear not to be known in the flora of tabulation of species incluc South Florida: Acanthospermum aes Amaranthus ee Amoracia aquatica, Asimina spectosa, \iplentum platynenron, Bacopa ieee Bignonta capreolata, Brassica haber. ie longi. Chloris ae: eee (Heterotheca) floridana, Eleocharis vivipara, Eragrostis simplex (= E. eminent), Flavert floridana, na ings one ENUM semper Gentiana pennelliana, Juncus ee ees indica, Lev, hloa filtf » Linarta floridana, Liquidambar styraciflua, Lobelia flovidana, ee te ae nae ee fle: i Ja sylvatica (8.1.), Oncidium bahamense, Oxalis violacea, Petalostemon eae iia es Pi puree fmbriata, Poly us AU ieee Rhexia nashis, Rhynch ip Vain lat. Salsola kali lea} Ser aria gl WHed, Silene antirvh Stip oides, Suaed Tradescantia ohiensis, Tragiola(G la) pilosa, Typha angustifolia, Weleiana scandens, Lannichellia palustris. 29:1 A second category of vanished names is composed of the species that were reported in the Avnotated Checklist but were not retained by the Flora even though the species is known in South Florida or may reasonably be assumed to have existed there in the recent past. Considerable latitude 1s required in the assignment of a species to a flora, for at the poorly documented end of the scale one or more specimens may exist yet there be little doubt that the species is no longer present in a living state. Such quasi-components of a flora are perhaps best handled in the form of a note, with the species left unnumbered or in some way or other given a secondary or tentative status. We have found eighteen species and one family (Zingiberaceae) which we believe to be proper inclusions in the flora of South Florida chat should not have been dropped by the Flora. Six of these, for nomenclatural or other reasons, require names different from those used in the Annotated Checklist. Finally, a third category of vanished names is simply a tabulation of syn- onyms that the F/ora has failed to assign to an accepted species. These names are retained in the present listing since the inconsequentiality of this correction is of course not apparent to the person attempting to trace a name from the Annotated Checklist to the appropriate treatment in the Flora. We have considered twenty-three names worthy of comment. We have listed these vanished names in alphabetic order, rather than in the sequence originally used, as an aid to rapid checking. Data as to habitat and range given in the Annotated Checklist are here repeated in quotes, €x- cept for those names that should have been treated as synonyms by the Flora. The use of “C,” “D,” or “M” in these quoted passages is the code used in the Annotated Checklist in reporting species as present in the counties of Collier, Dade, and Monroe. Habitat and range data as given for those species that we believe should be deleted from the flora of South h Florida are of course considered erroneous. 22 LIST OF CORRECTIONS ADIANTUM CAPILLUS-VENERIS. “Moist hammocks, solution holes, CDM.” This fern is not known south of Hernando County (Correll 1938; Wherry 1964; FLAS; USF). DELETE SPECIES. ALOcaAsiaA INDICA. “Homestead, D.” This (or A. macrorrhiza (L.) G. Don) is perhaps cultivated as a rare novelty, but is not known by us as an escape. DELETE SPECIES and the genus Alocasia. ALOE VERA. This name does not appear as a synonym under Adve barbadensis Mill. by which name the species is treated in the Flora (p. 281). Dates of publication of these two names, however, indicate the correct name is A/ve vera (L.) Burm. f.' RALIA SPINOSA. “Hammocks, D.” The Fora (p. 659) qualified the earlier report with “apparently noc well established in south Fla.” No Dade County specimens have been located, and the species apparently does not extend south of Polk County (FLAS, USF). DELETE SPECIES, the genus Arafia, and the Araliaceae, BRACHIARIA PLANTAGINEA. “Moist ground, D.” This Brazilian grass has now escaped at several places in Florida, but no collections have been seen south of Palm Beach County (PLAS). Brachiaria subquadripara (Vrin.) Hitchc. is a frequent escape in South Florida and was treated in the F/ora (p. 168); it was not recorded in the Annotated Checklist, and the presumption is that the earlier name was based on a misidentification. DELETE SPECIES. /AKILE LANCEOLATA, “Coastal beaches, DM.” Our unc erstanding of Cakile is that it is represented in South Florida by two entities, both subspecies of C. lanceolata: ssp. lance- dlata, and ssp. fustformis (Greene) Rodman (Rodman 1974: FLAS: USE). The Annotated Checklist has these two (the second in the form of an unpublished combination attributed to J.P. Patman), but has also C. edentula (Bigel.) Hook., a northern species that we have not seen south of St. Johns and St. Lucie counties (Rodman 1974: FLAS). The Flora (p. 431) recorded C. edentula (with a note that it is only “presumably” in its area), and C. fusiformis; the F/ora should have retained both C. lanceolata and C, fusiformis, either as two species or as lesser entities of C. lanceolata, the prior name. RESTORE Ca&ile lanceolata (Willd.) O. E. Schulz. CAPSICUM FRUTESCENS. “Hammocks, CDM.” Recent workers generally creat this species as distinct from Capsicum annuum L. (cf. Smith & Heiser 1951: Heiser & Pickersgill 1969; contra, Shinners 1956). The common native species in South Florida is C. /rutescens, and was correctly so recorded in the Annotated Checklist. Occasional plants of C. annuum L. var, aviculare (Dierb.) D'Arcy & Eshbaugh (1973) occur, perhaps as escapes, throughout Florida and could appropriately have been included in the checklist; the Flora (p. 759) reported only this species, as C. annuum var. minimum (Mill.) Heiser. RESTORE Capsicum frutescens a Cassia CORYMBOSA. “Homestead, D.” This shrub is restricted in cultivation to central and north Florida Usely 1975). We do not believe it escapes even within this northern range. DELETE SPECIES. ‘This plant was first described by Linnacus in 1753, as Adve perfoliata var. vera. Linnacus’ varietal epithet was transferred co specific level by Burman in 1768, while Miller, also in 1768, independently } | J described the species as A. barbadensty. Following the analysis of Reynolds (1966) that the name A/ve I & 5 y harbadensis Mill. anctedated Alve vera (L.) Burm. f., Mill preci a ‘rs name has obtained wide use. But more ise dating of the relevant publications (Stafleu 1967) indicates that the appropriate portion of Burman’s Flora Indica appeared “Mar-Apr (ante 6),” while Miller's The Gardeners Dictionary was pub- lished 16 April. A/ve vera would thus appear to be the prior name by a minimum of ten days (Newton 79). 1979 ie Pa, CAYAPONIA RACEMOSA. “Hammocks, Everglade Keys, D.” This habitat and location data would appear taken directly from Small (1933) which in turn is based upon early collections from Dade County (Small & Carter 792 in 1903, NY; Small & Wilson 1593 in 1904, NY; Small & Carter 2722 in 1906, NY). Although the F/ora cautioned (p. 816), “No recent collections of this species from south Fla. have been seen, and it may no longer be in our area,” 1t was found again in 1976 (Castellow Hammock Park, Dade County, Avery 486, FLAS, FTG). Small’s identification, however, was in error, and has been corrected b P. Wunderlin. RESTORE (replace with) Cayaponta americana (Lam.) Cogn. in DC. CEANOTHUS AMERICANUS. “Drier sites - C.” We have not seen this species south of Polk County, and Brizicky (1964b) was not willing to extend its range south of “northern Florida.” The Flora (p. 582) recorded the species only as a note, remarking that it “...may occur locally in our area. However, we have seen no specimens from out Fla.” DELETE SPECIES and the genus Ceanothus. JHAMAESYCE CHIOGENES. Burch (1965) has considered this name as synonymous with C. blodgettii (Engelm. ex Hitche.) Small. CHAMAESYCE CORDIFOLIA. “Sand dunes, CDM.” Correctly interpreted, this is a northern species. Burch (1965) did not record it south of Highlands and Lee counties. DELETE SPECIES CHAMAESYCE GEMELLA. Burch (1965) treated this as synonymous with C. opthalmica (Pers.) Burch. C AESYCE GLOMERIFERA. Burch (1965) considered this as synonymous with C. byper- rcifolta (L.) Millsp. CHAMAESYCE MATHEWSII. Burch (1965) included this with C. maculata (L.) Small. CHAMAESYCE MOSIERI. Burch (1965) treated this within C. garberi (Engelm. ex Chapm.) Small. CHRYSOPSIS MICROCEPHALA. “CDM.” We fully agree with Small (1933), Dress (1975), and Semple et al. (1980) that Chrysopsis, Pityopsis, and Heterotheca merit separate gencric recognition. Dress (1953) included this entity within the typical variety of Pyfyopsrs jon (Chrysopsis) graminifolia, assigning it a range in Florida south only to Bradforc counties, while Semple & Bowers (1985) interpreted it as within var. fenmifolia which they extend to southern Florida. Although the F/ore (p. 855) could perhaps have placed this in and Flagler synonymy under its Heterotheca graminifolia var.graminifolia, one might best avoid use of the name. DELETE SPECIES CHRYSOPSIS NERVOSA. “Pineland, CDM.” Although viewed by Semple & Bowers (1985) as a variety of Prtyopsts graminifolia, we support Dress (1975) in retaining P. nervosa at specific rank. This species is common in South Florida and is perhaps what the Flora (p. 855) intended ae its Heterotheca graminifolia var. tracyi. RI ‘STORE (replace with) Pityopsis nervosa (Willd.) I ZIENFUEGOSIA HETEROPHYLLA. This species was excluded from Florida (Fryxell 1969), our plant now being known as C. yacatanensis Millsp. The Flora (p. 593) treated these species correctly but did not clearly indicate the previous assumption that they were identical. CLERODENDRON FRAGRANS. The plant intended by the Annotated Checklist is widely cultivated in Florida and occasionally escapes. We are in agreement with Moldenke (1980) that this name should be placed in the synonymy of Clerodendrum philip} Schauer, by the Flora (p 7) Crusia FLAVA. “Not seen recently, hammocks, Key West, M.” Wood & Adams (1976) have pointed out the reports of this tropical species for Florida are unsubstantiated, the specimens so labeled being C. rosea Jacq. DELETE SPECIES. 294 Cy PERUS INFLEXUS. Horvat (1941) and other workers have considered this a synonym of Cyperus aristatus Rotth. CYPERUS PSEUDOVEGETUS. related to C. virens Michx. They were well understood by McGivney (1938 seen collections south of northern Florida. The present report should probably be referred to both common in South Florida and “Low ground, CD.” This species is one of several closely We have not either C. distinctus Steud. or C. surimamensis Rottb., a cited by the Flora. DELETE SPECIES. SMODIUM CILIARE. “Cutler Ridge, D.” This sae is predominately northern and is rare a of Alachua County. It is, however, in Dade C pan as documented by recent collections (Atwater in 1958, FLAS; Avery 486 i in 1968, FLAS). ae not have been deleted the Flora. RESTORE Desmodinm care hl ex Will DIUM LINEATUM. “Homestead, s species 1s largely tom with only a few sellections seen by us south of Alac ie County. In Dade County we know of it only as a collection from a “scarified lot, Homestead” (Hawkins 41 in 1927, FLAS). We have no reason to believe that this specimen was the source for the above report, but it provides sufficient verification. RESTORE ae /ineatum DC. DIGITARIA DIVERSIFLORA. “Olc and roadsides, CDM.” Swallen (1963) has dis- tinguished this tropical species, ee in Dade and Monroe counties, from D. ciliaris (Retz.) Koel. (=D. ascendens (ABK.) Henr.). Swallen’s name, however, was not the earliest. RESTORE (replace with) Digitaria nae Roem. & Schult. ex Loud. Dicirarta firirorMis. “D.” This species is largely northern; we have seen no collections from peninsular Fl iorida, South Florida collections are probably to be referred to D. villosa (Walt.) Pers. DELETE SPECIES DoticHos Hose. “Agr. Exp. Sta. Homestead, D.” This species is perhaps better known as Vigna hosei (Craib) Back. It has been introduced into Florida on an Pama basis, as a possible ground cover. It 1s not as n to escape. DELETE SPE ELEOCHARIS ALBIDA. “Wet soils, This aan species 1s ne n in Dade County, with several recent collections (Gis es in 1971, FLAS; Avery 1196 in a. FLAS). It should not have been deleted by the Flora. RE STORE Eleocharis albida Tor ELEOCHARIS EQUISETOIDES. “Wet gladeland, solution holes.” This species is not known south of Lake County (Ward & Leigh 1975). South Florida ealledtions probably should be referred to Eleocharis interstincta (Vahl) R. & S.; this species was not reported in the Annotated Checklist but correctly does appear in the Flora (p. 219). DELETE SPECIES ELEPHANTOPUS TOMENTOSUS. “Pineland dryer sites, CDM.” James (1959) and Ward (1975) have described this species as not extending closer to South Florida than Leon and Wakulla counties. The only South Florida representative of this genus is E. e/atus Bertol. The earlier error was corrected, but not explained, in the Flora (p. 877). DELETE SPECIES. ERLANGEA INCANA. “D.” The report of this species is from Moldenke (1944). He flatly stated it to be in cultivation. We do not know otherwise. DELETE SPECIES and the genus Erlangea. FIMBRISTYLIS HARPER. Ward (1968) and Kral (1971) have treated this name as inclu- ded with Fimbristylis caroliniana (Lam.) Fern. It should have been so indicated by the Flora (p. FURCRAEA MACROPHYLLA. ~D.” The Flora (p. 290) treated this species only as a note, remarking it “may persist” from cultivation. It is rarely if ever cultivated and there appear to be no reports, nor documenting specimens, of its persistence. We see no need to retain such an insubstantial waar even as a note. DELETE SPECIES GALACTIA BRACHYPO “Miami, D.” This name ts based on A. W. Chapman collec- 295 tions from the vicinity of the Apalachicola River, northwestern Florida. Although Rogers (1949) maintained it as a species, it seems more probably an aberrant form of the northern Galactia erecta (Walt.) Vail. A Dade County collection cited by Rogers (dry rocky soul, Miami, Hood 71863 in 1912, FLAS) is apparently an atypical G. volwbilrs (L.) Britt. This last species may be what was intended by the above — DELETE SPECIES. GREWIA POPULIFOLIA. “Fantastic gardens, Sout iami, D.” This species is now becoming frequently cultivated in South and Central Florida but 1s not known to escape. DELETE SPECIES and the genus Grewia. HyYMENOCALLIS CAYMANENSIS. Recent authors (Adams 1972; Correll & Correll 1982) consistently place this name in synonymy under Hymenocallis latifolia (Mill.) Roem JATROPHA MANIHOT. “CDM.” Rogers (1963) has treated this plant, the manioc or cassava, as Manthot esculenta Crantz. The Flora (p. 536) accepted this name but failed to give its synonym. This frost-sensitive species is very sparingly grown on the Florida keys and in Dade County, but we do not believe it persists outside of cultivation. The only basis for its inclusion by Small (1933) and the Flora appears to be a 1904 collection by J. K. Small reported by Webster (1967), from what was likely a cultivated source. DELETE SPECIES (and Manihot —, LANCHOE CRENA “Waste places — CDM.” The Flora was of two minds as to the inclusion of this species. a was treated as a note (p. 438), with the statement, “in disturbed sites and hammocks....no specimens, however, have been seen recently.” This is not one of the more vigorous species, and we are unaware that it ever escapes. DELETE SPECIES. ALLSTROEMIA INTERMEDIA. “Florida Keys, M.” Porter (1969) assigned this name to the synonymy of Ka/lstroemia paras Norton, a species not known to occur in Florida. Our representative of this genus is K. maxima (L.) Hook. & Arn. DELETE SPECIES. LACHNANTHES TINCTORIA. This name should have appeared in the F/ora as a synonym of oe hada (Lam.) Dandy. It does appear in place of the correct name in the egend f Flora’s plate of the species (p. 292). NGUAS SPECIOSA. “Cult. — D.” This species is betcer known as A/pinia zerumbet. It was collected in Addison Hammock, Dade County, as early as 1915 (FLAS) and has since been found repeatedly as an escape in the south and central parts of the state. RESTORE (replace with) A/pinia zerumbet (Pers.) Burtt & R. M. Sm., and Zingiberaceae. INIMA. “Canals, D.” Daubs (1965) recognized a species under this name, although he used it for plants occurring no closer to Florida than Texas. His specimens, however, conform to Lemna minor L., a species rare in Florida and unknown south of Glades County (Landolt 1986). We suspect the Annotated Checklist may have had the very similar L. obscura (Austin) Daubs, which is common. DELETE SPECIES LEMNA VALDIVIANA. “Stagnant pools, C.” Peninsular Florida is appropriate for che reported range of this species. D. W. Hall has informed us he has seen collections of it from Collier and Dade counties, and E. Landolt has provided us an unpublished map showing its presence in Collier County. RESTORE Lemna valdiviana Phi I. LIMNOBIUM SPONGIA. ‘Shallow waters, C.” This dis aquatic is rare south of Polk and St. Johns counties, but we have seen a specimen from Collier County (Atwater M-137 in 1959, FLAS). RESTORE Limnobiam es (Bosc) S ud. LOCHNERA MINOR. “Wasteland - The Flora (p. 701) retreated to a note that this species (Vinca minor L.)“...is w ne y ae and has been collected as an escape in Palm Beach County. [t may Ae occur locally 1n our area.” Even this more modest distribu- tion does not accord with our observation that this northern eee cannot be cultivated successfully in peninsular Florida. DELETE SPECIES (and Vinca minor) and the genus Lochnera. 296 LORINSERIA AREOLATA. “Open hammocks, CDM.” We have not seen specimens of this ern south of Glades County, although Correll (1938) reported a Lee County collection. JELETE SPECIES and the genus Lorinseria. LubDWIGIA INTERMEDIA. This combination, as published in the Annotated Checklist, is — rh illegitimate. Its basionym, [snardia imtermedia Small & Alexander, was treated by Munz (1965) with Ludwigia repens Forse. LupwiGiA PALUSTRIS. “Fresh water, CDM.” We have seen this species south to Charlotte and Lee counties (FLAS), but noc farther. It ts & e confused with Ludwigia repens Forst., which is common in southern Florida. DELETE SPECIES. LUDWIGIA SPATHULIFOLIA. “Low ground, D.” ae a. is related to L. curtissi Chapm. with which it has hesitantly been combined by a recent monographer (Peng 1989). Since its type locality is near Perrine, Dade County, the name should have been addressed by the F/ora. Pending a further judgment as to its status, RESTORE Ludwigia spathulifolta Small MACADAMIA TERNIFOLIA. “Spice and Fruit Park, Homestead D." This tree ts occasional - ly cultivated in Florida, buc does not spies DELETE SPECIES and the genus Macadamia. MAMMEA AMERICANA. “Canal edge, Tamiami Trail, D.” This tree is infrequently cultiva- ted, and is tender. The Flora (p. 609) believed it “probably is not established.” Its report as an escape was based upon Moldenke (1944). Without further indication of its persistence, we believe it best excluded from our flora. DELETE SPECIES and the genus Mammed. Marsitea vestita. “D.” Old reports of this fern ally from “Orange Beach,” Dade County, were based on a collection (Underwood 66 in 1891, PH) from Orange Bend, Lake County (Ward & Hall 1976). In this century it has been known in Florida only in Franklin, Hillsborough, Sarasota, and Seminole counties. DELETE SPECIES, the genus Marsilea, and the Marsileaccac. Nyssa SYLVATICA VAR. BIFLORA. “Swamps, C.” We have not seen the swamp cupelo (for which we prefer N. bif/lora Walt.) south of Glades County. (The typical black tupelo does not extend south of Alachua County.) There appears to be no previous report from Collier County. Monachino & Leonard (1959) called attention to a specimen labeled as from Lignum Vitae Key, Monroe County (Small & Britton in 1919, NY), but H. K. Rickert (pers. comm., 1966) was unable to relocate the specimen. The Flora (p. 648) noted the Lignum Vitae Key collection, but remarked, “No recent collections ite oo en seen, and its occurrence in our area is doubtful.” DELETE SPECIES, the genus Nyssa, and the Nyssaccae. ics of specimens attributed to Lignum Vitae Key and distributed to the University of South Florida and perhaps elsewhere 1s unquestionably in gross error as to its origin. In each case the label ts a standard printed form, headed “New York Botanical Garden, with the cooperation of Mr. Charles Deering, Exploration of the Florida Keys, Tropical Florida.” The labels further bear in print, “Hammock, Lignum Vitae Key, Monroe County. Collec- tors J. K. Small, N. L. Britton, December 13, 1919.” Further data, in blue ink, is in the hand of J. K. Small. In each case the known range of the species mounted on the sheet and named on the label ts completely at variance with the stated source on Lignum Vitae Key. Two striking examples of this mis-labeled series of specimens are Geatiana pennelliana Fern. and Janens trigonocarpus Steud. The gentian was reported in the Plora (p. 697) without details but with the suggestion that the species oS no longer be present in the area. The basis for chis Ses was an excellent sheet (USF 17005) of six flowering plants labeled in Small’s hand, “Gentiana” and correctly ee ie R. W. Long as Gentiana leigh ye That species, however, is a tightly restricted pine flatwoods endemic known only ane six counties in West Florida (Wakulla to Walton) and well doc ae by Causes 2) (1941) and Pringle (1967). It is difficult to believe it could survive as a disjunct native or be adventive in the tropical hammock of Lignum Vitae Key. oe Juncus trigonocarpus was reported by the Plora (p. 280) as “Hammock, Lignum e Key...rare,” the report being based on a sheet (USF) bearing three plants, with the same printed heading and hand-labeled “Juncus.” This species belongs to the same acid-soil coastal plain flora as the Gentiana, it is known in Florida only from west of Franklin County (FLAS, FSU, USF), alchough northward it extends into the Carolinas. Again, the habitat of this species is grossly difference from that of Lignum Vitae Key. The full history of what appears to be a set of spurious 1919 Lignum Vitae Key collec- tions has not been traced. Since Small did not refer to these would-be striking range exten- a, sions in his later publications, it is clear that he did not accept them as the labels would now appear to read. The USF specimens were obtained by Mr. George Cooley from the New York Botanical Garden, in unmounted form, in the early 1960s. They would appear to have been part of a “rich collection,” referred to by Monachino & Leonard (1959) that for many years had lain in storage at NY, but no information is at hand as to how many other pe labeled specimens were acquired by USE if any, or how widely dispersed they yay be in other herbaria. It is apparent that species with less sharply restricted North Florida ranges may more readily pass as acceptably small non-disjunct range extensions. At the least, a caveat ts in order as to the acceptance of South Florida ranges based on 1919 Small & Britton collections from Lignum Vitae Key. OENOTHERA MOLLISSIMA. “Drier soils, CDM.” Small (1933) reported this South American species (as Ratmannia mollissima) to occur trom Florida to Texas, a claim that has been disregarded by North American workers (Munz 1965; Correll & Johnston 1970). The Florida plants, at ne are not of that species, but have been suggested by PH. Raven (pers. comm., 1978) to represent Ocnothera humifusa or O. humifusa — O. laciniata hybrids. DELETE SPECIES Oryza SATIVA. “Anhinga Trail, Taylor Slough, D.” A perennial, awned rice, “Red Rice,” now commonly treated as distinct from Oryza sativa L., has been known for some years to be well established in the vicinity of Taylor Slough, Beene ladles National Park, and is represented by specimens (Atwater GS-174 in 1959, FLAS; Craighead in 1961, FLAS). RESTORE (replace with) Oryza raufipogon — ft. PANICUM CONDENSUM. This species was obscurely recorded by the Flora (p. 189) variety of Panicum agrostoides Spreng., a species better known (Voss 1966) as PB. rigidulum Bosc ex _ 1S) CUM CONSANGUINEUM. “Pinelands, CIM.” Freckmann (1967) retained this en- tity at cae level of species and oe it south co central Florida. Specimens we have seen support this range. It is very close to, and in fact appears to intergrade with, P. angastifolrum E., a frequent South Florida species. DELETE SPECIES. PANICUM GEMINATUM. This distinctive grass was retained in the Flora (p. 176) as Paspa- fidium gemmatum (Forssk.) Stapf in Prain, but without indicative synonymy. Gould (1968) and other recent workers have supported this generic eae CUM LONGIFOLIUM. “Low ground, wet pincland, ” This name is now usually placed in synonymy under Paarenm rigidulum Bosc ex Nees. a es this grass is somewhat ageressive and is widespread to che north, we have not scen it south of Okeechobee County. DELETE SPECIES. PASPALUM DIFFORME. “Pinelands, "We know this plant (as a ies of P. flor- danum Michx. var. floridanum) south ae to Flagler anc Marion counties. Specimens bear- ing this name from Dade County have been misidentified Passa floridanum vat. glabratum Engelm. ex Vasey (= P. gigantenm Baldw. « ex Vasey). DELETE SPECIES ‘ 298 PHYSALIS TURBINATA. “Pineland, D.” Even after che work of Menzel (195 1) and Water- fall (1958, 1968) we remain in doubt as to the correct names of the Florida species of Physalis section Pubescentes. More commentary is needed than is appropriate here. Our present interpretation is that the Flora (p. 754) was correct in describing this species as hairy and with a fruiting calyx up to 4.cm. long; many specimens bearing this name are the near-glabrous, smaller-fruited P cordata Mill. (= P. pubescens L. var glabra (Michx.) Water- all). We have, however, seen no specimens of true PB turbinata Medic. from Florida. DELETE SPECIES PIPER OTOPHYLLUM. “Mangrove belt, eae CDM.” Adams (1972) considered this name synonymous with Piper fadyenti C. DC. in DC., a montane endemi c of Jamaica. It disappeared frot m the Flora, apparently eae as a gross misidentification. DELETE SPECIES and the genus Peper. NSETTIA DENTATA. “Pinelands, D.” Neither Burch (1966) nor we are able to cite collections of this species from Florida. DELETE SPECIES. POLYGONELLA MYRIOPHYLLA. “Sand scrub, C.” The Flora (p. 375) did not refer to the earlier Collier County report, but noted this species “has been found in Dade County.” Horton (1963) knew it only as an endemic of central Florida (Highlands County and north- ward), and we have been unable to locate documenting specimens south of that area. DELETE SPECIES. POTAMOGETON FLUITANS. “Long Pine Key, DC.” This name is now usually considered a synonym of P otamogeton nodosus Poir. in Lam. which in our exper western Florida. Confusion with the widespread P. illinoensis DELETE SPECIES. PUNICA GRANATUM. “Waste places, old fields - D.” Although this shrub, as Small (1933) stated, does persist around old homesites, it has only very sparingly been cultivated in southern Florida and apparently has never been documented there as an escape. DELETE SPECIES, the genus Pwnica, and the Punicaceae RHACOMA CROSSOPETALUM. jos e is not found south of ee is suggested. This name should properly have been listed by the Flora (p. 368) as a synonym under aaa rhacoma Crantz (Brizicky 1964a). RHAPIDOPHYLLUM HYSTRIX. “D.” We believe this palm does not range south of Hardee ane Highlands counties (F LAS. USF). DELETE SPECIES and the genus Rhapidophyllum. yO RHYNCHOSPORA EDISONIANA. Gale (1944) treated this as Rhynchospora microcarpa Baldw. ex Gray HY NCHOSPORA FERNALDI. “Coastal beaches, C.” Gale (1944) reported this species only south to Lee County, and the Flora (p. 231) deleted it ee reference to equivocal statement in the Annotated Checklist. However, collections from Collier County do eXISC Craighead | in 1956, FLAS; Sandy excavation, Marco Island. Atwater in 1958, FLAS). RESTORE ee cca Gale. RHYNCHOSPO ymous with » the un- =~ lades, D.” This coastal plain species is admitted to the flora of Florida aay on the Bach . a few northern collections (Leon Co., FSU; Duval Co., FLAS). It is unknown in the peninsula. DELETE SPECIES. RUELLIA NUDIFLORA. “Pinelands, D.” Long was a student of Ruellia but did not discuss the distribution of this species. It apparently is restricted to Texas and northern Mexico and had not previously been reported for Florida. It was omitted, without com- ment, by the Flora (p. 7 ) DELETE SPECIE SAGITTARIA FALC “C.” The nearest location at which this species is known to occur is in Franklin County, western F] tonids i rie DELETE SPECIES SALVIA MUTABILIS. We assume tl mutabilis (A. Rich.) Briq. is a fapsus calami for the common H yptis 299 SCHOENOLIRION ALBIFLORUM. Although this name may originally have been applied to the more northern Schoenolirion croceum (Michx.) Wood (Sherman 1969), it has long been used for S, e/fiotti ee under which the fora os 283) might have placed it in synonymy. SCHRANKIA ANGUSTISILIQUA. Probably only Schrankia — (Dryand.) Macbr. occurs 1n ae South Florida area (Isely 1973; FLAS; contra, Beard 1964, who cited Broward and Dade County specimens as $. wncinata Willd.). Isely (1973) ee S. angustisiliqua to be a “phase” of S. microphylla, within which the Flora (p. 454) should have placed this name in synonymy SCLERIA SETACEA. Core (1936) originally held this taxon distinct from Scleria reticularis Michx., but Fairey (1967), working under Core’s direction, treated it as var. pubescens Britt. The — name nen have been given by the Flora (p. 237) as a synonym. oe The type of this taxon is a collection from Miami (Pollard a C allins aoa in "1898, Y), and thus the name should not have been disregarded by the Flora (p. 298). esc - plant can be interpreted to fall within a rather inclusive interpretation of the East Coast and Bahamian nee arenicola Bicknell, the South Florida population appears to merit specific rank (Ward & Gillis 1975). RESTORE S$ —~ yrinchinm miamiense Bicknell. SMILAX LANCEOLATA. “Hammocks, CDM.” This name is a synonym of Smilax smallii Morong. The ees Pte is not known south of Highlands County (Duncan 1967; FLAS) DELETE 5 IGRU oe soul, CDM.” Although this Eurasian ae has been repor- ted in Dade County ‘O Arcy 1974), recent workers (Heiser et al. 1979; Ogg er al. 1981; Schilling 1981) place all related South Florida materials in S. americanum Mill. (including S, eel Jacq.) or S. pseadogracile Heiser. DELETE SPECIES. SPAR 1UM AMERICANUM. “Swampy shores, CDM.” This distinctive plant is not known in Florida south of Highlands County, either by Beal (1960) or by us (FLAS, FSU, FTG, USF). DELETE SPECIES, the genus Sparganium, and the Spargantaceae. SPARTINA CYNOSUROIDES. “Saltwater beaches, low tidal pee CDM.” We have seen no records of this grass south of Duval C a DELE’ TE SPECIE SPERMACOCE TENUIOR. “Pinelands, CDM.” Long (1970) to treat Spermacoce flori- dana as a variety of S. tenuior L. In the Flora (p. 806) the typical variety was excluded from South Florida. This conforms with our Se a alchough the differences between these two taxa seem of specific magnitude. ADD S. floridana Urban’ to the South FI ee flora, and DELETE SPECIES en above STENOCARPUS SINUATUS. oral ci es - D.” central and southern Florida, o there is no evidence that it escapes. DELETE SPECIES This tree is cultivated occasionally in and the genus oe STYLISMA AQUA earing floor of hammock, D.” This species was attributed to South Florida By Gi are see a . noted the reported range to southern Florida had been based upon misidentified specimens, mostly of Stylisma villosa (Nash) House, a a s pas was not a south of the Apalachicola River in West Florida. DELETE $ aoe DENTATA. “Moist hammocks, CD.” We have scen specimens of this species from Collier County (Scw// in 1937, FLAS; Avery 2071] in 1969, FLAS), as well as a oe > Spermacoce floridana Urban dates from 15 Aug 1913 (Symbolae Autillanae 7:550, published on this date according to Carroll & Sutton 1965), while S. £eyensés Small appears to be later. The preface of Small, F aes of the Florida Keys, 1913, was dated 11 Aug 1913, but the publication was recorded in the 28 Nov 1914 Index to American Botanical | eed Torrey Bot. Club 41:575). Since the Index was 914. ae am of the Florida Keys would seem not to have been available until after Oct | 300 “volunteer in slat house,” Dade Co. (FLAS). It had been deleted by the F/ora. RESTORE eee dentata (Forssk.) E. St. John. THELYPTERIS PATENS. “Moist hammocks, CDM.” Neither Wherry (1964), Smich (197 1b this species beyond the single 1905 Dade County collection discussed by R. St. John (in Small 1938). We concur with the Flora (p. LOL) that this species is highly unlikely co on ~nor recent field botanists in South Florida have been able to expand state records of remain a member of our flora. DELETE SPECIES. THELYPTERIS SERRA. “D.” St. John (in Small 1938), in his report of this species for the state, nee that the original specimens from Dade County were not wholly typical of the species as known in the West Indies. Wherry (1964) and Smith (197 la) have pointed out i indeed, the Florida collections so named fall within the morphological limits of Thelypterts angescens (Link) Munz & Johnst. DELETE SPECIES. THELY PTERIS TETRAGONA. “Moist a CD.” This species is not known south of Marion and Hernando counties (FLAS, USF). DELETE SPECIES. THELYPTERIS ULIGINOSA. The Flora : 100) omitted this name as a synonym of The/y- on pleris torvestana “— ) Alston. THermopsis MOLLIS. “Pineland, hammocks, CDM.” Wilbur (1963) reported this piedmont cz south only co northern Georgia. No conflicting specimens have been a. DELETE SPECIES a nd sii genus Thermopsis. ONIA SCABERRIMA. “Sandy pineland, CDM.” Jones (1964) failed to find this taxon, which he treated as ey ene Michx. var scaberrima (Nutt.) Gray, outside of South Carolina and eastern Georgia. DELETE SPEC A FLORIDANA. “Margins, hammocks, CD.” This legume, alchough common in 7S northern and ee oa apparently does not occur south of Desoto County (FLAS, USF). DELETE SPL VIGNA UNGUICULATA. “Waste places, CD.” This plant is frequently seco hin we know it as an escape only on Sanibel Island, Lee County (FLAS). DELETE SP WALTHERIA AMERICANA. The Flora (p. 604) failed to give this name as a synonym a Waltherta indica L. AREA CUNEIFOLIA. “D.” Channell & hee (1964) have reemphasized the earlier in- secies as known only from Liberty and Gadsden counties, West > terpretation of th Florida. Dade cn material would be the related but distinct Warea carters Small. DELETE SPECIES WASHINGTONIA fiLIFERA. “CDM.” The vast majority of Florida individuals of this genus are the gracefully slender Washingtonia robusta Wendl. Washingtonia filifera is cultivated only with rarity in Florida, and we are unable cto find evidence that it escapes. DELETE SPECIES. WEBSTERIA SUBMERSA. “Submersed aquatic, C.” This plant is better known as Websterra ihe (Poir.) Hooper (= Scerpas confervoides Poir. in Lam.). It ts rare in Florida, and we en no specimens from south of Lake and Highlands counties. Inclusion of this species in South Florida is based upon specimens from Collier County so annotated by K AS. FSU, USE ~ Svenson (FLAS, FSU, USF); the plants, however, are submersed forms of the common Eleocharis baldwinti (Yorr.) C nen ae LETE SPECIES and che genus Websteria. WoLFFIA COLUMBIANA. “Canals, D.” The Flora (p. 254) omitted this genus. Yet this species is frequent in Florida and D. a Hall, recently a student of the Lemnaceae, reports to us that he has seen a Dade County collection (Stimpson 738, FSU). RESTORE Wolffia columbiana Karst RIS BALDWINIANA. “C.” Kral (1966) did not find this species south of Marion County. We are unable to oa him. DELETE SPECIES. ZAMIA INTEGRIFOLIA. “Pinelands, CDM.” The Flora (p. 108) reported the common 301 Florida species of this genus to be Zamia pumila L., a name initially applied to plants from Hispaniola but extended by Eckenwalder (1980) to all members of the genus in the West Indies and Florida. The Florida representatives, though undoubtedly “founder effect” selec- tions from this Caribbean complex, seem sufficiently uniform to merit taxonomic recogni- tion (Hardin 1971; Ward 1979). Aiton’s Zamia imtegrifolta, sans earlier, 1s oo SSTORE rally superfluous aad ie meee RE i (replace with) Z. floridana A. OSTERA MARINA. “CDM.” This marine species ranges en Beaufort, North oe (Den Hartog 1970), north into the arctic seas. Ie is completely unknown in Florida. DELETE SPECIES, the genus Zostera, and Zosteraceae. ACKNOWLEDGMENTS We should like to thank John Beckner, Helen B. Correll, William J. Dress, David W. Hall, Patricia K. Holmgren, David L. Martin, John Popenoe, Warren H. Wagner, and Richard P. 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Gard. 53: 5 — 376. CARROLL, E. & $. SUTTON. 1965. A cumulative index to the nine volumes of the Symbolae Antillanae Arnold Arbor., Jamaica Plain, Mass. 272 pp. CHANNELL, R. _W. JAMES. 1964. Nomenclatural and taxonomic corrections in Warea (C ee. Ror 66:18 — 26. CLAUSEN, R. T. 1941. Studies in the Gentianaceae: Gentiana, section Pnenmonanthe, ew en Bull. Torrey Bot. Club 68:660 — 663 CORE 1936. The American species of Scleria. Brittonia 2: i 105. coe. a S. 1938. A ane check-list of Florida ferns and fern allies. Amer. Fern J. Pet 1646-34, 0110 CORRELL, D.S. & H. B. C a 1, 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz. 1692 pp. 302 CORRELL, D. 5. & M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Found., Renner. 1881 pp. D'ARCY, W. G. 1974. Rhitin: and its close relatives in Florida. Ann. Missouri Bot. Gard. 61:819 — 867 D'ARCY, W. G - H. ESHBAUGH. 1973. The name for the common bird pepper. Phytologia oe oi DAUBS. bo: 1965. A monograph of Lemnaceae. 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JAMES, C. W. 1959. The status and distribution of Elephantopus elatus. Rhodora 3 sh S. B. 1964. seme - the narrow-leaved Vernonia of the southeastern United . Rhodora 66:382 mie R. 1966, Xyris ae of the continental United States and Canada. Sida 1971. A treatment of Absildgaardia, Bulbostylis and Fimbristylis (C pees ae) for North America. Sida 4:57 — 227, LAKELA, O. & E C. CRAIGHEAD. 1965. Annotated checklist of the vascular p = ants of 303 Collier, Dade, and Monroe counties, Florida. Fairchild Tropical Garden & Univ. of Miami Press, Coral Gables. 95 pp. LANDOLT, E. 1986. eee investigations in the family of duckweeds (Lemnaceae). The family of Lemnaccae — a sae aa study. Vol. 1. Veroff. Geobot. Inst. ETH Stiftung Rubel Zurich. 71: 6 LONG, : W. 1970. Additions and nomenclatural a in the flora of southern Florida = Rhodora 72:17 — 46. LONG, . W. & O. LAKELA. 1971 [“new edition,” 1976]. A Flora of Tropical Florida. Univ. of Miami Press, Coral Gables. 962 McGIVNEY, M. V. DeP. 1938. A revision of - yperas} subgenus Excyperus founc United States. Catholic Gary of Amer. biol. ser. no. 26. 74 pp. eae M. Y. 1951. Cycoraxonomy and genetics of PAysalis. Proc. Amer. Phil. Soc. 95:1 183. MIASE K. M. A. a ae and local fern floras of the United States, suppl. Il. Amer. Fern J. oe 109 — | MILLE .G. 197 . ak genera of the Urticaceae in the southeastern United States. J. Acola ne 52:40 — 68. 97 |b. The Polygalaceae in the southeastern United States. J. Arnold Arbor. 52:267 — 284. MOLDENKE, H. N. 1944. A contribution to our knowledge of the wild and cultivated flora of Florida — 1. Amer. Midl. Naturalist 32:529— 590 1980. A sixth summary of the Verbenaceae, Avicenniaceae, Stilbaceae, (oie in the Chloanthaceae, Symphoremaceae, Nyctanthaceae, and Eriocaulaceae of the world as to valid taxa, Se ee distribution and synonymy. Plainfield,N. J. 629 pp MONACHINO, J. & LEONARD. 1959. A new species of Justicia om Florida. Sy Giga UNZ, P A. 1965, ee N. Amer. Flora 5:1 — 2 MYINT. ear Revision of the genus Stylavme ( Convolvu eee, Brittonia 18:97 — 117. NEWTON, L. E. 1979. In defence of the name Alve vera. Cact. Succ. J. Gr. Brit. Al: i OGG, A.G., B. _ ROGERS & E. E. SCHILLING. Characterization of black nightshade (Solanum se and related species in the United States. J. Weed Sci. Soc. 29:27 — 32. NG, C. I. 1989. The systematics and aaa of Ladwigia sect. Microcarpinm eee Ann. Missouri Bor. oe 6:221— 302 POPPLETON, J. E., A. G. SHUEY & H. C. “SWE ET, 1977. Neer of Florida’s east coast: a checklist of the vascular plants. Florida Sci. 40:362 — 389. PORTER, D. 1969. The genus Kallstroemia eet en Contr. Gray Herb. 198-41 = [53, san eee J. S. 1967. ous of Gentiana, section Prenmonanthae, in eastern North crica. ees 19:1 Gi ,G. W. he The Aloes of Tropical Africa and Madagascar. Trust. Aloes a Far Mbaban — Siem 1974 syste and evolution of the genus Ca&s/e (Cruciferae). Contr. Gray ne ‘ 205: = ae RS, . 7. 1949. a. genus Galactia in the United States. Ph. D. thesis, Duke Univ. — oa D. J. 1963. > a of Manihot esculenta Crantz and related species. Bull. Torrey Bot. an 90:43 — ROGERS, G. K. 1984. an Zingiberales (Cannaceae, Marantaceae, and Zingiberaceae) in the ean United States. J. Arnold Arbor. 65:5 —55 304 1985. The fee of the Phytolaccaceae in the southeastern United States. J. oo Arbor. 66:1 —3 SEMPLE, J. C . BLOK, & P. Scie 1980. Menpholeeical, anatomical, habit, and hi ibicat ee among the goldenaster genera Chrysopsis, Heterotheca, and Pityop- sis (C ue 7 ey Canad. J. Bor. 58:147 — 163. SEMPLE, J. C. & E D. BOWERS. 1985. A revision of the goldenaster genus Pityopsis Nutt. (Compositae: Astereae). Univ. Waterloo Biol. Ser. 29:1 — 34. SHERMAN, H. L. 1969. A systematic study of the genus Schoenolirion (Liliaceae). Ph. D. thesis, oe Univ. 129 pp a SHINNERS, L. H. 1956. ‘Technical names for the cultivated Capsicum peppers. Baileya SMALL, J. — 1933. Manual of the southeastern flora. New York. 1554 pp. —__________—.. 1938. Ferns of the southeastern states. Lancaster, Pa. 517 pp SMITH, A. R. 197 la. a he Thelypteris normalis complex in the southeastern United States. Amer. Fern J. 61 32 SJ SMITH, A. R. 197 Ib. Systema of the neotropical species of Thelypteris section Cyclo- sorus. Univ. Calif. Publ. 59:1— 143. SMITH, PG. & C. B. HEIS 1951. Taxonomic and genetic es on the cultivated peppers, C pied annuum 4 aaa C. frutescens L. Amer. J. Bot. 38:362 — 368 SPONGBERG, S. A. 1978. The a of Crassulaceae in the southeastern nited States. J. Arnold Arbor. 59:197 —2 STAFLEU, E A. 1967. Taxonomic ae Int. Bureau Plant Taxonomy and Nomencla- ture, Utrecht. STEYERMARK, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames. 1725 2 SWALLEN, J. R. 1963. New species of Digitaria and Trichachne. toes 65:355 — VOSS, E. G. 1966. Nomenclatural notes on monocots. Rhodora 68:435 — 463. VOSS, E. G. 1972. Michigan flora, Pare I. Cranbrook Institute of Science. 488 pp. ——___________. 1985. Michigan flora, Part II. Cranbrook Institute of Science. 724 p WARD, D. B. 1968. Contributions to the flora of Florida Castanea 33: 123— 135. 4, Fimbristylis Cae ee) Contributions to the flora of Florida — 7, Elephantopus, Psendo- elephantopus (C barnes: Castanea 40:213 — 217. ed. 1979. Rare eerie plants of Florida, vol. 5: Plants. Univ. resses of Fla., Gainesville. 175 pp. B. & D. W. HALL. 1976. Re-introduction of Marsilea vestita into Florida. Amer. Fern J. 66:113— 115. WARD, D. B. & W. T. GILLIS. 1975. The Sisyrinchium of the Bahamas. Phytologia 4S Pp WARD, D. B. & E. M. LEIGH. 1975. Contributions to the flora of Florida — 8, E/- eocharis (Cyperaceae). Castanea 40:16 — 36 WATERFALL, U. T. 1958. A taxonomic study of the genus PhAysalis in North America north of Mexico. Rhoden 60152173, a 12968. eee in United States Physalis. Rhodora 70:574 — 576. WEBSTER, G. L. 1967. The genera of Euphorbiaceae in the southeastern United States. J. Arnold her 48:303 — 361, 363 — 430. WHERRY, E. T. 1964. The southern fern guide. Doubleday & Co., Garden City. 349 pp. WILBUR, L. 1963. The Leguminous plants of North Carolina. North Carolina Agr. Exp. Sta. tech. bull. 151. 294 WOOD, C. E. & P ADAMS. 1976. The genera of Guttiferae (Clusiaceae) in the southeastern United States. J. Arnold Arbor. 57:74 —90. DOCUMENTED CHROMOSOME NUMBERS 1990: 1. MISCELLANEOUS NORTH AMERICAN VASCULAR PLANTS BRUCE D. PARFITT, DONALD J. PINKAVA, DEBBIE RICKEL, DAN FILLIPI, BETH EGGERS Department of Botany, Arizona State University Tempe, AZ 85287-1601, U.S.A DAVID J. KEIL Department of Biology California Polytechnic State University San Luis Obispo, CA 93402, U.S.A. The following meiotic chromosome counts are documented by specimens deposited in Herbarium of Arizona State Untversity (ASU). Previously uncounted taxa are represented by an asterisk (*). A double asterisk (**) indicates a new number for the species. METHODS Flower buds were collected in developmental series from plants growing in native habitats. Buds were killed and fixed in chloroform, ethanol, and glacial acetic acid (6:3:1 v/v) or ethanol l and glacial acetic acid (3:1 aa. transferred to 70% ethanol after 24 hrs, and refrigerated. Anthers were squashed in iron/acetocarmine and mounted in Hoyer's medium (Beeks Ly): Percentage positive pollen stainability was determined for the three Penstemon individuals for which chromosome counts were obtained. Pollen was taken from closed anthers of herbarium specimens and stained in ant- line-blue lactophenol for 48 hrs (Maneval 1936). A minimum of 500 pollen grains per individual were scored. Percentage of staining, normal- size grains is indicated in parentheses after the chromosome number tn the results below. ANACARDIACEAE *Ruus AROMATICA Aiton var. PILOSISSIMA (Engl.) Shinners. n= 15. — Arizona. Yavapai Co.: NW side of Prescott, Arrowhead Dr, Parfitt 3898 & Roberts. Stipa 14(2):305 — 308. 1990. 306 APOCYNACEAE AMSONIA TOMENTOSA Torr. & Frem. vat. TOMENTOSA. 7 ll. — Arizona. Mohave Co.: jct. of Signal Rd & Alamo Rd NNW of Alamo Lake, Parfitt 4181 & Christy. *CYCLADENIA HUMILUS Benth. 7 = 7. — Uran. Grand Co.: Castle Valley, red clay slopes below Parriot Mesa, Anderson 86-51. ASTERACEAE BERLANDIERA LYRATA A. Gray var. MONOCEPHALA B. L. Turner. 7 = 15. CHiHuAHUA. Rte 14, near La Junta, Pinkava 13334, 13339, McGill, Reeves G Nash. CHAENACTIS STEVIOIDES Hook. & Arn. 7 = 5. — ArIzZONA. Apache Co.: Navajo Reservation, Hwy 12 ca. 5.7 mi NW of jet with Hwy 13, just N of Tsedadhotsosi (a tiny mesa), Reeves 8273 & Parfitt. *CIRSIUM DRUMMOND Torr. & Gray X C. wHEELERI (A.Gray) Petrak. » = 16. — Arizona. Apache Co.: Alpine Campground, 5 mi N of Alpine, Landrum 5249 & Landrum. *HELENIUM ARIZONICUM Blake. » = 15 + 1B. — Arizona. Coconino Co.: AZ 260, near turnoff to Willow Springs Lake, TLIN R14E S31, Parfitt 3845 & Rickel. MACHAEFRANTHERA GRINDELIOIDES (Nutt.) Shinners. 2 = 4. Nevaba. White Pine Co.: 38 mi W of Ely Cemetery, US 50, Parfitt 3817 & Roberts. MACHAERANTHERA ASTEROIDES (Torr.) Greene var. GLANDULOSA B.L.Turner. » = 4. — Arizona. Yavapai Co.: Antelope Hills Golf Course, US 89, Prescott, P7//2p7 5, STEPHANOMERIA EXIGUA Nutt. var. EXIGUA. » = 8. — Baja CaLmorNIA Norte. 22.9 mi S$ of San Vicente, Pinkava 11110, McGill, Hensel, & Maclntyre. STEPHANOMERIA PAUCIFLORA (Torr.) A. Nels. 7 8. — ARIZONA. Pinal Co.: 37.4 mt NW of Oracle Junction, ee 10993, Lehto & Hensel. BORAGINCEAE **LITHOSPERMUM INCISUM Lehm. » = 14. — Arizona. Apache Co.: Navajo Reservation. SW slopes of Chuska Mts., ca. 3.2 mi NE of Tsaile, 36° 19’ 20" N, 109° 10’ 10” W, Reeves 8302 & Parfitt. BRASSICACEAE *PENNELLIA LONGIFOLIA (Benth.) Rollins. 2 = 8. — ARIZONA. 307 Apache Co.: ca. 6 air mi E of Mt Baldy Peak, TON R27E S14, Parfitt 3852 & Rickel. CACTACEAE OPUNTIA REPENS Bello. 7 = 11. — Puerto Rico. ca. 4.4 mi SE of Boqueron along PR 303, Keil 16512. CUCURBITACEAE *MARAH GILENSIS Greene. 7 = 15. — Arizona. Maricopa Co.: ca. 2.5 mi N of Sunflower, TON RYE S4, Parfitt 3731 & Bricker. ERICACEAE * ARCTOSTAPHYLOS PRINGLE Parry. 2 = 13. — ARIZONA. Yavapai Co.: 1.1 mi E of Crown King, Parfitt 3740, Bricker G& Eggers. FABACEAE *PsORALEA MEPHITICA S. Wats. 2 = |1. — Arizona. Maricopa Co.: ca. 11 mi S of Sunflower, T4N R8E S82, Parfitt 3732 & Bricker. Vicia pULCHELLA Kunth. » = 7. — Arizona. Apache Co.: White Mtns, 2.7 mi NE of jct Forest Service Rds 117 & I 17a, near Carnero Lake, T8N R27E SO, Parfitt 3874 & Rickel. LAMIACEAE MENTHA ARVENSIS L. var. vILLOSA (Benth.) S. R. Stewart.» = 48. — Arizona. Coconino Co.: AZ 260, near turnoff to Willow Springs Lake, T1IIN R14E $31, Parfitt 3848 & Rickel. RUTACEAE *THAMNOSMA MONTANA Torr. & Gray. 7 = LO. — Catirornia. San Bernardino Co.: N side of Clark Mt, TI7N R13E S15, Parfitt 3586 G Baker. SCROPHULARIACEAE *KECKIELLA ANTIRRHINOIDES (Benth.) Straw ssp. MICROPHYLLA (Gray) Straw. 7 = 8. — Arizona. Maricopa Co.: 12.7 mi S of Sunflower, Parfitt 3725 & Bricker. OrtHocareus tuTEUs Nutt. 7 = 14. — Arizona. Apache Co.: ca. 6 air miles E of Mt. Baldy Peak, TON R27E S14, Parfitt 3854 & Rickel. PENSTEMON EATONU Gray ssp. EATONIL. 2 = 8(97.4%). — ARIZONA. Maricopa Co.: cult., 924 W. Watson Dr, Tempe, Parfitt 3003. PENSTEMON PSEUDOSPECTABILIS Jones var. CONNATIFOLIUS (A. Nels.) 308 Keck. 7 = 8 (60.0%). — Arizona. Maricopa Co.: cult., 924 W. Watson Dr, Tempe, Parfitt 3601. *PENSTEMON EATONIL SSp. EATONIL X P) pSEUDOSPECTABILIS var. CONNATIFOLIUS (FI hybrid, a volunteer resulting from natural pollina- tion). 7 = 8 (36.2%). — ARIZONA. Maricopa Co.: cult., 924 W. Watson Dr, Tempe, Parfitt 3602. ROSACEAE Correction. The following was erroneously reported as Rhus ovata (Parfitt et al, L985 PRUNUs ILLICIFOLIA (Nutt.) Walp. 27 = 15. California. San Diego Co.: 117.2 mt W of Yuma, AZ, at jct. of CA 94 & I-8, Gallagher 82-9. REFERENCES BEEKS, R. M. i Improvements in the squash technique for plant chromosomes. Aliso le - 13 MANEV, Ee = ee Lactophenol preparations. Stain Technol. 11:9. PARFIT i. D., Baker and M. - ee 1985. IOPB chromosome number reports. oa Taxon 35:162 — NOTES A NOTE CONCERNING THE TYPIFICATION OF TWO PLANTS DESCRIBED FROM TEXAS — Ina recent revision of the genus Ca//irhoe Nuttall (Malvaceae) (Dorr 1990), a paratype (Lindheimer 681) was chosen as the lectotype for C. /eiocarpa R. Martin since the holotype (C. D. Marsh s.n.), which had been deposited at NA, was missing and presumed lost. However, the holotype has been found since at US (D. Nicolson, pers. comm.). Evidently, in 1953 the herbaceous material at NA was transferred to US and the Marsh specimen was misfiled under Ma/va L. The revision of Callirhoe (Dorr 1990, p. 40) should be corrected as follows: CALLIRHOE LEIOCARPA R. Martin, J. Wash. Acad. Sci. 28(3):108. 1938 aes ae — Tyee: UNITED STATES. Texas: aes . o.: Spofford, 4 Apr 08, C. D. Marsh s.n. (HOLOTYPE: US-2133209 ex NA-8099!). When Dorr and Barnett (1986) sought to clarify the identity of Nephropetalum Robinson & Greenman (Sterculiaceae) they concluded that the genus was synonymous with Ayewza L. (Sterculiaceae) and that the sole species, N. pringlez Robinson and Greenman, was conspecific with A. limitavis Cristobal. They were unable to locate the type of N. pringlez, which presumably was deposited at GH. Consequently they designated an isotype at WT as lectotype. After Dorr & Barnett (1986) published this lectotype, C. Crist6bal informed the curator of GH that the holotype of N. pringler was filed under Trimmfetta L. (Tiltaceae) (W. Kittredge, pers. comm.). Crist6bal concurred with our taxonomic assessment and N. pringler remains a synonym of A. /imitaris. However, the typification of the former name Is now: NeEPHROPETALUM PRINGLEL Robinson & Greenman, Bot. Gaz. (Craw- fordsville) 22: 168. 1896. — Twer: UNITED STATES. Texas. Hidalgo Co.: Hidalgo, woodlands, 3 Aug 1888, Pringle 2272 (HOLOTYPE: GH!; isoryee: VWT!). — Laurence J. Dorr, New York Botanical Garden, Bronx, NY 10458-5126, uti REFERENCES DORR, L. J. 1990. A revision of the North American genus Ca/firhoe (Malvaceae). Mem. New York Bor. Gard. 56:1—75. DORR, L. J. and L. C. BARNETT. 1986. The identity of Nephropetalum (Sterculiaceae). Taxon 35:163 — 164 Sipa 14(2):309. 1990. 310 MONTIA_ LINEARIS (PORTULACACEAE), NEW TO MISSIS- SIPPI — While examining a shipment of exchange specimens that the junior author had sent to MICH in 1989, the senior author and A. A. Reznicek (MICH) noticed that two specimens determined as Claytonia virginica looked odd. In a later communication, the junior author noted that he also thought that the specimens looked a little strange for C. virg7- nica, but thought chat their appearance might be attributed to the use of herbicides along roadsides where they were found. The presence of white rather than dark-veined pink petals ruled out C. virginica immediately. The combination of fibrous roots and alternate caul- ine leaves excluded the plants from the genus C/aytonia, suggesting instead that they belonged in the genus Montia. Comparison with the MICH hold- ings of Monta revealed that the Mississippi collections were specimens of Montia linearis (Dougl. ex Hook.) Greene, narrow-leaved montia, a native western species found at lower elevations from British Columbia south to California and eastward to southwestern Saskatchewan, Montana, and Utah (Hitchcock & Cronquist 1973, Scoggan 1978). These determina- tions were subsequently confirmed by Walter A. Kelley of Mesa State College, Grand Junction, Colorado. Montia linearis is known from the following collections: MISSISSIPPI: Coahoma Co.: between highway ane old railroad tracks, along US 49W between Tutwiler and Rome, T25N,R3W,S36,SE”%, 4 May 1988, Bryson 7483 (SWSL) G 7572 (MICH, SWSL). Panola Co.: wet area N of MS Hwy 6, 11.4 mi W of Batesville, T9S,RIOW,S20, 4 May 1988, Bryson 7553 (SWSL). Quitman Co.: wet area along MS Hwy 3, Sof Marks, 4 May 1988, Bryson 7537 (SWSL). Sunflower Co.: wet area along US 49W, S mi N of Drew, T23N,R3W,S4, NEY, 4 May 1988, Bryson 7544 (SWSL). Tallahatchie Co.: wet area, W side of US 49W, 1 mi S of Tutwiler, T25N,R2W,S31,NW%4, 4 May 1988, Bryson 7458 IBE, SWSL); roadside park, jct. of US 49E & 49W, S of Tutwiler, 4 May 1988, Bryson 7500 (MICH, SWSL); 11 Apr 1990, Bryson G Newton 8775 & 8783 (CLEMS, DUR, FLAS, GA, IBE, MESA, MICH, MMNS, MO, NCU, NLU, NY, OSC, SMU, SWSL, TAES, TENN, UARK, UC, VDB, VSC, and WTU). This is the first report of the genus Motia in the Southeastern United States. Its occurrence along roadsides in five counties strongly suggests an introduction from an unknown source, possibly the first such occurrence outside of its native range. It may have arrived via a railroad source since all but the Panola County site are within 4 mi of Illinois Central Gulf track- age abandoned in the 1980 The largest of these Sees is at the roadside park just south of Tutwiler. The plants here occur in open areas or under widely scattered Quercus nigra and Q. phellos on poorly drained, heavy clay and/or on well Sipa 14(2):310. 1990. oa al drained, sandy, loam soils. Within the park, M. /zmearis is colonial much like populations of C/aytonia, ranging from a few plants to several thousands over areas of less than one square meter or up to 30 m° in size. Since we have no information on when the species may have first arrived, it is likely that either other specimens may exist in herbaria, likewise assumed to be Claytonia virginica, or that it has been overlooked. From the habitat range of these collections, M. linearis may be expected in other locations within the Mississipp1-Yazoo Delta Region along roadsides or in cemeteries, roadside parks, and even lawns. — Richard K. Rabeler, Univer- sity of Michigan Herbarium, North University Building, Ann Arbor, MI 48109-1057, U.S.A. and Charles T. Bryson, USDA-ARS, Southern Weed Sci- ence Laboratory, Stoneville, MS 38776, U.S.A. REFERENCES HITCHCOCK, C.L. and A. CRONQUIST. 1973. Flora of the Pacific Northwest. Univer- sity of Washington Press, Seattle. xix + 730 pp. SCOGGAN, H.J. 1978. The flora of Canada, Part 3 - oo ees co Violaceae). Natl. Mus. Canada Natl. Mus. Nat. Sci., Publ. oo -1115 CAREX COMOSA (CYPERACEAE), NEW TO MISSIS- SIPPI. — Carex comosa Boott is a large, conspicious sedge that inhabits low, wet, non-acid soils (Mackenzie 1935). It occurs from southeastern Canada to Minnesota, generally southward to central peninsular Florida and west to Texas, and in the west from California to Washington, east- ward to Idaho (Steyermark 1968, Godfrey and Wooten 1979). It is dis- junct in central Mexico (Hermann 1974). However, the nearest known record to Mississippi is from Crittenden County, Arkansas which ts adja- cent to and across the Mississippi River from Mississippi and Tennessee (Smith 1978). Despite searches for this species, especially by the senior author during the past decade, C. comosa had not been found within the boundaries of Mississippt. While on a collecting trip, C. comosa was found growing at an oxbow lake in Coahoma County located in the Yazoo-Mississippi Delta Region. Similar oxbow lakes are scattered throughout the Yazoo-Mississippi Delta Region. Specimens collected: MISSISSIPPI. Coahoma Co.: 5.6 mi (9 km) N on US 49 from its jct. with MS 3, E side of US 49, Home Cypress Hunting C a Lake, NE Dublin; E of US 49 Sec. 34, T26N, R3W, 15 May ra Bryson YS58 & & G. Jones ( ctb, private reg of Charles T. Bryson, se , IBE, MICH, srs NLU. SWSL, TENN, UARK, VDB, VSC, WARM), . Es 4719 & Bryson (ASTC, SMU, SWT, TAES, TEX, US). Sipa 14(2):311. 1990. 212 The habitat is an open oxbow lake (cypress slough) with scattered bald cypress trees (laxodium distichum) growing in the lake. The area adjacent to the oxbow ts cleared farmland with brownish loamy clay soil, alluvium (Qa) of the Tutwiler formation within the Delta Region (Holocene, Quaternary) (Bicker 1969). Carex comosa was found frequently growing on bald cypress stumps and logs in close association with C. decomposita, and less frequently growing along the bank’s edge. Other associated taxa were Ludwigia sp., Bidens sp., and Lemna sp. Due to the large population size of more than 50 clumps and the large cespitose clumps, the authors believe that C. comosa has been established at this location for many years and is not a recent introduction. Subsequent searches in potential habitat in Coahoma County and surrounding counties in northwestern Mississippi by the senior author during 1990 failed to lo- cate additional populations of C. comosa. This record is approximately 62 miles (LOO km) southeast from the nearest station in Arkansas. We thank Gretchen D. Jones for her assistance in the field and for edito- rial comments. We also acknowledge Richard Carter (WSC), David Castaner (WARM), A. A. Reznicek (MICH), and J. K. Wipff (TAES) for helpful suggestions. — Charles I. Bryson, USDA, ARS, Southern Weed Sci- ence Laboratory (SWSL), Stoneville, MS 38776, U.S.A. and Stanley D. Jones, M. Tracy Herbarinm (TAES), Department of Range Science, Texas AGM University, College Station, TX 77843, U.S.A. io REFERENCES BICKER, A. R., JR. 1969. Compiler. Mississippi Geological Survey. Mercury maps Inc. Jackson, MS. GOL JEREY, R. K. AND W. WOOTEN. 1979. Aquatic and wetland plants of southeastern United Cee Monocotyledons. University of Georgia Press, Athens, 712 meee i J. 1974. Manual of the genus Carex in Mexico and Central America. U.S Dept. hai ee - 467. 219 pp. MACKENZ 35. Cyperaceae. North American Flora 18 (part 7). New York ee a. S, p. oe 33, SMITH, E. B. 1978. An atlas and annotated list of vascular plants of Arkansas. University of Arkansas, Fayerceville, 592 pp. STEYERMARK, J. A. 1963. Flora of Missouri. The lowa State University Press, Ames, p. 373-74. Sipa 14(2):312. 1990. 313 CHLORIS INFLATA (POACEAE) NEW TO LOUISIANA — Recent collections from Louisiana have yielded a specimen of Chloris inflata Link. espite an extensive distribution throughout the tropical and subtropical regions of the Old WORLD and New World, there have been few collec- tions of Chloris inflata from the continental United States. Prior to the collection reported herein the only confirmed collections of this taxon were from Bexar County, Texas (WA. Si/vews 6413, US) and Cameron and Hidalgo counties in extreme southern Texas (Correll and Johnston 1970; Gould 1975). A range map dot shown for Mississippi in a paper by Ander- son (1974) could not be verified. Allen (1980) did not report it for Louisiana. Considering the widespread distribution of this species in the Caribbean and Bahamas, it is highly likely that this taxon will be found elsewhere in the southern United States in the near future. The collection data are: LOUISIANA. Orleans Parish: on the shoulder between Leake Ave. and the tracks of the Illinois Central RR, opposite the U.S. Army Corps of Engineers facility, 7 Aug 1989, John Macgregor 892 (UNO) — John R. Macgregor, Department of Biological Sciences, University of New Orleans, New Orleans, LA 70148, U.S.A. and Charles M. Allen, Division ue Sciences, Loutsiana State University at Eunice, Eunice, LA 70535, U.S. REFERENCES ALLEN, C.M. rie Grasses of Loutsiana. The University of Southwestern Louisiana, Lafayette. 358 ANDERSON, D. bE. 1974. a of the genus Ch/oris. Brigham Young Univ. Sci. Bull. Biol. Ser. 19(2):1 aes L, D.S. and M.C. i INSTON. 1970. Manual of the vascular plants of Texas. xas Research Foundation, Renner, Texas. 1881 pp. boa. EW. 1975. The grasses of Texas. Texas A&M University Press, College Station. 653 pp. Stipa 14(2):313. 1990. (Continued from front cover) A taxonomic comparison of Artstida ternipes and Aristida hamulosa (Gramineae). Jonathan S. Trent and Kelly W. Allred. Circumscription of Agarista bolivtensis (Ericaceae). Walter 8. Judd and Paula M. Hermann. A new status for Quercus shumardii vat. acerifolia (Fagaceae). Nick Stoynoff and William J. Hess. Annotated checklist of New Mexican Convolvulaceae. Daniel F. Austin. Deletions and restorations in the flora of southern Florida. Daniel B. Ward and Frank C. Craighead. Documented chromosome numbers 1990: 1. Miscellaneous North American vascular plants. Bruce D. Parfitt, Donald J. Pinkava, Debbie Rickel, Dan Fillipi, Beth Eggers, and David J. Keil. ZL 263 267 Zf> Zor 305 NOTES. A note concerning the typification of two plants described from Texas. 309 — Montia linearis (Portulacaceae), new to Mississippi. 310 — Carex comosa A ae w to Mississippi. 311 — Chloris inflata (Poaceae) new to Louisiana. 3 Book reviews 168, 188, 214, 230, 250, CONTRIBUTIONS S IDA to Botany VOLUME 14 NUMBER 3 AUGUST 1991 CONTENTS Plantae alpinae novae Mexicanae: Sedum chrysicaulum (Crassulaceae). J. Andrew McDonald. A natural intergeneric hybrid in the x = 6 group of the Astereae (Asteraceae). Ronald L. Hartman and Meredith A. Lane Morphological characters as indicators of rubber content in Guayule (Parthenium argentatum — Compositae). M.A. Foster, S.E. Gabel, T.8. Ward, L.G. Kleine, PK. McCann and Jaroy Moore. Classification and systematics of eastern North American Vitis L. (Vitaceae) north of Mexico. Michael O. Moore. Hard grass (Sclerochloa dura, Poaceae) in the United States. David M. Brandenburg, James R. Estes and John W. Thieret. Seedling morphology in Clematis (Ranunculaceae) and its taxonomic implications. Frederick B. Essig. A new species of Conradina (Lamiaceae) from northeastern peninsular Florida. Robert Kral and Robert B. McCartney. Observations on Fryxellia pygmaea (Malvaceae). Paul A. Fryxell and Jesus Valdés R. Biocidal Stda (Malvaceae). Charles L. Burandt, Jr. The genus Hosta Tratt. (Liliaceae) in Korea. Myong Gi Chung and Jong Won Kim. Common names for vascular plants: guidelines for use and application. John T! Kartesz and John W. Thieret. (Continued on back cover) 315 ao) 339 369 O77 US ISSN 0036-1488 S1ipA, CONTRIBUTIONS TO BOTANY Founded by Lloyd H. Shinners, 1962 Publisher Wm. E Mahler Botanical Research Institute of Texas, Inc. 509 Pecan Street Fort Worth, Texas 75102 Editor Assoctate Editor Barney L. Lipscomb John W. Thieret Botanical Research Institute Northern Kentucky University of Texas, Inc. Highland Heights, Kentucky 41076 509 Pecan Street Fort Worth, Texas 76102 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 3, pages 315 — 524. Copyright 1991 by Wm. E Mahler PLANTAE ALPINAE NOVAE MEXICANAE: SEDUM CHRYSICAULUM (CRASSULACEAE) J. ANDREW McDONALD Department of Botany University of Texas at Austin Austin, TX 78713, U.S.A. ABSTRACT Recent explorations of the alpine-subalpine floras > northeastern Mexico have revealed several new species of Sedum. The most widespread of these, here described as S. chrysi- caulum, approaches most closely S. parvum ssp. a but is easily ee from the latter by persistent, pale yellow-green, dorsally sulcate leaves, petals 7.5 mm long, and a preference for habitats above timberline. RESUMEN joe Exploraciones recientes de las floras sess en el nordeste de México han dado luz a varias especies nuevas de Sedum. La espe mas ampliamente distribuida, S. chrysi caulum, aqui descrita, asemeja aS. parvum ssp. nanifolium, con la cual se distingue por hojas persistentes, verde-amarillentas palidas, sulcadas en la superficie adaxial, pétalos 5.0 — 7.5 mm de largo, y por prefencia de habitats arriba de los limites arboreos. SEDUM CHRYSICAULUM McDonald, sp. nov. (Fig. 1) A — ee Hemsl. ssp. vanifolio (Frod.) Clausen caulibus infernis herbaceis foliis el- li orsaliter sulcatis 5 — 6 mm longis petalis 5 — 7 mm longis et folliculis 4-5 mm pane: differt. Herbs perennial, cespitose, 3—7 cm tall, 2—20 cm wide, glabrous. Stems branching from base, ascending, 5-9 cm long, | — 2 mm in diam at maturity, yellow-green, glabrous, smooth; internodes somewhat longer in basal portions, 0.5 —9.0 mm long; nodes occasionally rooting; roots fibrous. Leaves simple, sessile, narrowly ellipsoid, dorsally sulcate (flattened upon drying), 5 — 7 (-10) mm long, 2— 3 mm wide, light green or rarely orangish, with occasional red spots, glabrous. Inflorescence in terminal, congested cincinni; flowers 2—8; sepals 5, broadly lanceolate, mm long, 1.5—2.0 mm wide at base, pale green, coriaceous, glabrous; petals 5, erect, narrowly elliptic or lanceolate, 5 — 8 mm long, 2.0 —2.5 mm wide medially, ca. 2 mm wide at base, yellow with occa- sional red-pigmented dots or longitudinal striations, glabrous; stamens 10, 5-6 mm long, terete, attenuate, ca. 0.5 mm wide in diam at base, filiform at apex, yellow; anthers ovoid, basally cordate, yellow; carpels 5, 6—8 mm long, 1.2— 1.5 mm wide at base, yellow, glabrous. Fruit con- Sipa 14(3):315 — 319. 1991. iy IN: h, i! WW "a iM A ct Aly ae a VK cs FIG. 1. Illustration of Sed, eaf; note the red seri he red striations on icanlum. A. Growth habit. B. Mature follicle. C. Mature stem anc the stem, and the red punctae on the blade. 317 sisting of 5 erect, free follicles, narrowly ellipsoid, 7 —9 mm long, ca. 2 mm wide at base, often red-pigmented, glabrous, suture ca. 5 mm long, the style persistent, forming an attenuate apex ca. 2 mm long; seeds ca. 10, narrowly ellipsoid, 0.7 — 1.0 mm long, ca. 0.4 mm wide, brown, glossy, densely papillate. Type: MEXICO. Nurvo Leon: Mpio. Rayones, summit of Sierra La Marta, eastern ridge-top in ee vegetation, ca. 3650 m, 24 Aug 1989, McDonald & Mayfield 2556 (HOLOTYPE: > ISOTY PES Gea NY, UAT). — examined: MEXICO. Coahuila: a Arteaga, summit of Sierra La Viga, 3700 m, 22 Aug 1986, aaa 2099 (TEX), 24 Oct 1984, McDonald & Gomez 1157 (TEX, UAT); Mpto. pues as Suegenie of Sterra ise Marta, 3600 — 3700 m, 31 Aug 1980, Hinton et a 17977 (TEX); eee 1980, Hinton et al. 17919 (TEX); 25 Oct 1984, McDonald & Gomez 1242 (TEX); 22 Aug i McDonald 2136 (YEX). Nuevo Leon: Mpio. Galeana, summit of Cerro Potosi, 3700 19 Jun 1966, Gilbert 9 (TEX); 23 Aug 1984, Lavin 4787 (TEX); 14 Oct 1970, Hinton a 17303 (TEX); Mpio. Zaragoza, Sierra Pena Nevada, N of Picacho de San Onofre, 30 Nov 1984, McDonald & Gomez 1298 (MEXU, TEX, UAT) Three species of Sedum in the alpine-subalpine vegetation of north- eastern Mexico have yellow flowers and relatively short leaves (<6 mm), suggesting close relationships with S. parvum Hemsl. (sensu lato, Clausen 1978, 1979, 1981). Beaman & Andresen (1966) adopted the latter epithet for all of the Sedum collections from the alpine flora of Cerro Potosi, Nuevo Leon, despite the morphological and ecological distinctiveness of these populations in relation to the type population of S$. parvum Hemsl. ssp. parvum from warm and semiarid regions near the city of San Luts Potosi (Clausen 1979). All three of the above-mentioned alpine stoneworts appear to be undescribed (McDonald 1990), but two must await formal recogni- tion pending a critical study of the entire complex (Nesom, 1n prep). The most distinctive and widespread of these, occurring in all seven timberline refugia studied by McDonald (1990; Fig. 2), most closely approaches in morphology the low-elevational S. parvum Hemsl. ssp. nanifolium (Frod.) R. Clausen. Sedum chrysicaulum, here described, shares the following fea- tures with S. parvum ssp. nanifolinm: smooth stems, flattened and/or sulcate leaves that often bear distinctive red markings, and yellow flowers. Sedum parvum ssp. nanifolium is distinguishable from S. chrysicaulum, however, by green or reddish, basally lignescent stems that often branch distally, flattened leaves, ovate to broadly elliptical, 2.5 — 3.5 mm long, generally deciduous at the base of stems and congested in apical regions of the stems, petals 4 — 5 mm long, and follicles 2— 3 mm long. This and other subspe- cies of S. parvum recognized by Clausen (1978, 1979, 1981) generally occur in relatively arid, chaparral vegetation or pine-oak associations of northeastern Mexico and western Texas (Clausen 1978). 318 COAHUILA SAN LUIS POTOSI FIG. 2. Known distribution of §. chrysicanlum. TEXAS a1? In contrast, S. chrysicanlum has herbaceous, light yellow-green stems that branch mostly from the base, leaves narrowly elliptic, dorsally sulcate (a feature commonly obscured by desiccation), 5 — 6 mm long, persistent and dispersed relatively evenly along stems, petals 5.0 — 7.5 mm long and follicles 4—5 mm long (Fig. 1). Sedum chrysicaulum generally occurs from 3400 — 3700 m in timberline and alpine vegetation of northeastern Mexico (Coahuila, Nuevo Leon, Tamaulipas; Fig. 2) as an associate of sub- alpine vegetation dominated by the genera Arracacia Bancr., Castilleja Mutis, Erys¢mum L., Penstemon Schmid., and Senecio L., and stunted 1n- dividuals of Pinus culminicola Andresen & Beaman and Pinus hartwegit Benth. (McDonald 1990). Plants of S. chrysicaulum flower and fruit from June to October. ACKNOWLEDGEMENTS The author gratefully acknowledges support from the World Nature Association that financed an expedition to collect the type specimen. Dr. Guy Nesom shared his views on the relationships of the new taxon, and suggested the specific epithet. REFERENCES BEAMAN, J. H. & J. W. ANDRESEN. 1966. The vegetation, cee and phytogeog- raphy of the summit of Cerro Potosi, Mexico. Amer. fa foe eee) CLAUSEN, 1978. Sedum, seven Mexican perennial species. aH Torrey Bot. Club 105:214— 22 3) jesse 1979. ike in six areas of the Mexican cordilleran plateau. Bull. Torrey Bor. Club 10 16:205 — 1981. Virion of species of Sedum of the Mexican cordilleran plateau. Arnold Printing Corp., Ithaca, New York. McDONALD, J. A. ne “The alpine-subalpine flora of northeastern Mexico. Sida 14:21- 18 eS Bp Bp ace of the ks subalpine eae flora of north- eastern ves in P Ramamoorthy, J. Fa, R. Bye, & A. eds.), Biodiversity of Mexico: its origin a distribution. Oxford Press, New Yorke. BOOK REVIEWS Mayes, VERNON O. AND BARBARA Baytess Lacy. 1989. Nanise’ A Navajo Herbal. One hundred plants from the Navajo Reserva- tion. Navajo Community College Press, Tsaile, Arizona. An eas text on the common plants found on the Navajo Indian Nees cas in the “Four Corners Area” of Arizona, Utah, Colorado and New Mexico, a 25,000 s mule area. Each slate 1s presented with an illustration, scientific name and pronouncation guide, the derivation and the Navajo name. Each plant is ge! described and its associa- tion with other plants is briefly mentioned. The Navajo uses of the plant is also described followed by references from the ethnobotany literature. An interesting book from an un- usual region of the United States. John E. Ubelaker. KINDscHER, Ketty. 1987. Edible Wild Plants of the Prairie. An Ethnobotanical Guide. The University Press of Kansas, Lawrence, Kansas 66045. Paper. An introductory text that introduces wildflower enthusiasts to the value of numerous native and introduced plants of the Great Plains. The text resulted from an 80 across Kansas and eastern Colorado by the author. Presented are lin c illustration of common ane Indians names for the cones followed bya btiefdesceipdion and habi substantial section involves the food uses of the plants as well as food mythology and belief by the American Indians. The book is well written and enjoyable to read. John E. Ubelaker. STUBBENDIECK, JAMES AND ELverRNE C. Conarp. 1989. Common Legumes of the Great Plains, An Illustrated Guide. Illustrated by Bellamy Parks Jansen. University of Nebraska Press, 901 N. 17th St., Lincoln, NE 68588-0520. Hardbound $35.00. 330 pp. illustrated manual treating 107 is in detail common to the Great Plains. This includes full page illustrations accompan the text of each taxon. The text contains a ee description, synonymy, agronomic ma wildlife data, food, medicinal, and other es by American Indians and pioneers. Dichotomous keys are provided from the family level down to the species treated within each genus when there are more than one per genus. The excellent diagrammatic illustrations greatly enhance this publication. wfm Stipa 14(3):320. 1991. A NATURAL INTERGENERIC HYBRID IN THE x = 6 GROUP OF THE ASTEREAE (ASTERACEAE) RONALD L. HARTMAN Rocky Mountain Herbarium, University of Wyoming Laramie, WY 82071-3165, U.S.A and MEREDITH A. LANE R. L. McGregor Herbarium, University of Kansas 2045 — Ave., Lawrence, KS 66047-3279, U.S.A. ABSTRACT A single plane found west of nee Puebla, Mexico, is recognized as a hybrid between members of two x = 6 genera of Astere termediacy ree and reduced fertility. The cl is cs) as ea Aumile (Kunth) Benth. X I[socoma (Haplopappus) veneta (Kunth) E. Greene. The existence of this and two other ae hybrids of similar origin supports the hypothesis that the genera of Astereae having x = more closely related to one another than any is to genera having other base chromosome numbers, but should not be interpreted as evidence for uniting the genera. KEY WORDS: Asteraceae, Astereae, Haplopappus, Isocoma, Xanthocephalum, hybrid Mexico. RESUMEN Una planta encontrada al oeste de Zacatepec, Puebla, México ha sido reconocida como un ae entre dos x = 6 géneros de Astereae basado en fertilidad reducida y caracteres . La planta es considerada Xanthocephalum bumile (Kunth) Benth ‘pti veneta (Kunth) E. Gree natural sostiene x [socoma ne. La existencia de éste y otros dos hibridos de one ¢ la hipotesis de que los géneros de Astereae que tie mas cercanamente relacionados unos a Otros gue cualqui efa de éstos con otros generos con if- erentes nuimeros cromosomicos base, no obstante ésto no debe ser interpretado como evidencia para unir los generos. Found west of Zacatepec, Puebla, Mexico was a single plant, Hartman G Funk 4127, 19 Aug 1976 (RM and TEX), which has the following combination of features unlike that of any plant previously known to science: Sprawling perennial herb; stems longitudinally ridged, 15—25 cm long, reddish- brown to purple, densely villous, less so with age, the internodes 2 m long. Leaves alternate, often with fascicles of secondary leaves in axils, lanceolate to heh oe or pecasionully linear, 15 — 30 mm long, 1 — 5 mm wide, the apex mucronate, the base gradu- Sipa 14(3):321 — 329. 1991. 322 ally tapered, the margins with | — 3 (-4) salient, mucronate teeth per side, often not paired, the adaxial surface sparsely to een wa! villous, pitted on drying, the veins obscure, less so on the slightly paler abaxial surface. Capitulescence a terminal, corymbose cluster of {—8 heads; peduncles 5-30 mm long, bracteate, the bracts linear to scale-like, densely villous, sparsely so with age. Heads radiate, 7-8 mm high and 14— 18 mm wide in 7—9.5 mm high in fruic (pressed material); involucres hemispheric, 5 —6 mm high, 7—9.5 mm wide, the ee in, 34 — flower, series, imbricate, oblong to narrowly oblanceolate, appressed, 1-5 1 n long, the lower portion thickened, cua the upper 1/2 — 1/4 herbaceous, ia apex mucronate; receptacles flat to slightly convex, alveo- lace, - alveolae rimmed by scales 0.2 — 0.5 mm long. Ray florets heap (Fig. LA), 12 — 15; corolla yellow, the ae Lit be mm long, 0.3 —-0.41 in diameter, mod- ene to densely villous (Fig. 1A), the lamina broadly obl ie to valigic 5—6.5 mm long, 1.8 — 2.2 mm wide, with 4 nerves, the lobes irregular, 0.1 — 0.5 mm long; anthers 4—4, not well developed; ae branches 1.3 — 1. : mm ne either linear a ay Haus stigmatic throughout or with deltate appendages; blong to obovoid, long, tan, antrorsely pubescent; pappus bristles somewhat une aia qui a mm ae tan. oes florets hermaphroditic, 32—40; corolla yellow, goblet- ee the tube I ye & I 2.5— 2.7 mm long, 0.3 —0.4 mm in diameter, sparsely villous, the throat 1.2 — 1.5 mm long, 0.8 —0.9 mm in diameter, glabrous, the lobes narrowly triangular, 0.5 — 0.7 mm long, glabrous; anthers 5, functional; style branches 1 — 1.2 mm long with deltate append- aves; oe obovoid, 2.8 —3 mm long, tan, antrorsely pubescent; Me bristles un- equa il, 4 mm long, tan. Mexico: Puebla: sale flat (elev. ca. 2300 m), ca. 4.8 km WNW of oe on hwy 136. With ie humile, scattered individuals of Locoma venela, and species of Erigeron, Suaeda, Atriplex, Boutelona, and Distichlis in the immediate VIcInity. There are two possible explanations for the differences between this plant and known species: either it is a new species, or it is a hybrid between related but distinct taxa. Based on the data given below, we believe the latter to be the case. The plant was found with Nanthocephalum humile (Kunth) Benth. and lsocoma veneta (Kunth) E. Greene [= Haplopappus venetus (Kunth) S. E Blake}, two species of a group of genera of the Astereae that several authors have considered to be related. The bases for this assessment are the common base chromosome number of x = 6 and shared morphological characters including goblet-shaped disk corollas (first noted by Jackson 1966), deltate style-branch appendages on the disk florets, and rectangular epidermal cells on the adaxial surface of the ray corolla, as shown in Figure IC (Hartman 1990; Hartman et al. 1987; Lane 1980, 1982; Lane & Hart- man 1984, 1985; Lane et al. 1987) among other features (De Jong & Bea- man 1963; Keil & Stuessy 1977). In addition, many members of the group grow in saline or alkaline habitats (Robinson 1893; Steyermark 1937; Turner 1972; Mayes 1976; Watson 1977; Lane 1983). Further, Jackson (1966), Jackson & Dimas (1981), and Venugopalan (1966) have reported experimental hybrids between some of the species belonging to this group o QJ 1. Scanning electron micrographs of ray florets (scale for A = 1 mm, scale for Band C = n). A. Tube and lower portion of lamina of a ray floret from Hartman & Punk 4127 (RM) showing the poorly developed anthers (light colored) in proximity to two rudimentary adaxial lobes (dark colored) and the villose tube. B. Adaxial epidermis of the lamina of a floret shown in Fig. LA. C. Adaxial epidermis of the lamina of a ray Aoret of Nanthocephalum humile (Lane 2395, TEX), showing an epidermal pattern typical of all members of the x = 6 genera of Astereae. Specimens were prepared and photographed as described by Lane (1982). FIG. S i ] i eral I f adaxial surfaces of lol f di Ilas (scale = 5O 2m for all). A. Kantbcealen humile (Lane 2395, TEX). B. Hartman & Funk 4127 (RM). C. Lsocoma veneta (Hartman 3830, TEX). Specimens were prepared and Siti hed as described by — (1982). The epidermal pattern Son is typical of all members of the 7 = 6 genera of Astereae. 324 of genera, Lane (1980, 1983) found two natural interspecific hybrids in Nanthocephalum. Hartman & Funk 4127 shares a number of features with both of the parental species proposed here. The epidermal cells of the adaxial surface of the disk corolla lobes are identical to those of both X. humile and I. veneta (Figs. 2A-C), the disk corollas are goblet-shaped, the disk style-branch appendages are deltate, and the plant was found in a saline habitat. How- ever, while the pollen stainability of both species is 98% or greater (Jackson & Dimas 1981; Lane 1980; see Table 1), that of the putative hybrid is only 31.4% (205 of 399 observed grains were stained in lactophenol cotton- blue). Further, only 11 of 19 (57.9%) expanded achenes contained em- bryos. These indications of reduction in fertility are similar to those found in the natural and artificial hybrids mentioned above (Jackson 1966; Jack- son & Dimas 1981; Lane 1980, 1983) As indicated in Table 1, Hartman & Funk 4127 has young stems, pedun- cles, capitula, involucres, and phyllaries like those of X. Aumile, but capitulescence, receptacles, achenes, and pappus similar to 1. veneta. In all the other features listed in Table | and shown in Figure 3, the specimen is intermediate between the two taxa (e. g., habit, leaves, floret numbers, fea- tures of the style branch appendages, and achene lengths). The most striking features of this plant are found in the “ray” florets (Fig. 1A), which have densely villous tubes, unusually deep sinuses, and adaxial epidermal cells (Fig. 1B) quite unlike those of X. Aumule (1. veneta is eradiate) shown in Figure 1C. These florets, unlike the ray florets of “good” taxa of the Astereae, contain stamens (although rudimentary) and the style- branch appendages in some of the florets resemble more closely those of disk florets. The same phenomena occur in another natural radiate-eradiate hybrid between Machaeranthera restiformis B. Turner and M. gypsophila B. Turner (Turner & Sanderson 1971; Turner 1973). These anomalous “ray florets” appear to be highly modified disk florets (pers. obs. of the authors). Jackson & Dimas (1981), who experimentally hybridized 1, veneta with Haplopappus aureus A. Gray (a radiate species with » = 6), found that the presence or absence of ray florets is a single-gene character (see also Gottleib 1984), alchough length of the lamina when ray florets are present is appar- ently quantitatively inherited. Our observations suggest that length as well as other features of the lamina are polygenically controlled and that some genes involved in disk floret structure can be “turned on” by the allele for presence of ray florets. These hypotheses await testing. Recently, Guy Nesom brought to our attention two additional putative hybrids. The first (G. Castilla C. & M. Vazquez 3003, TEX) was collected in the same area (Mexico: Puebla: road from Zacatepec to El Carmen at $25 F e drawings of disk florets, showing goblet- wer corollas (scale bar as indicate Xena humile (Lane 2395, TEX). B. Hartman & Funk 4127 ( ). C. lsocoma veneta ( nse . TEX). Achene pubescence of Hartman & Punk 4127 Het lsocoma veneta is not shown, and only a fn - the pappus bristles are depicred. 220 border with Tlaxcala, 12 Jan 1984). It agrees in general with the descrip- tion of Hartman & Funk 4127 in most morphological features except the leaves are smaller, 10 — 20 mm long. Interestingly, the ray corollas exhibit one of the following conditions with respect to the adaxial petals: both are suppressed as in a normal floret, one or both is present but reduced in size, both are united into a narrow lamina 1/2 — 4/5 as long as the abaxial one, or one or both is present as a lateral lobe of the abaxial lamina. The second (HH. Iltis, A. Jose, & A. Lassergne 802, TEX) was collected approximately 140 km WNW of the site for Hartman & Funk 4127 (Mexico: Mexico: on Tapie 1. C omparison of Nanthocephalum humile (data from Lane 1980, 1983), Hartman and Funk 4127 (data from the specimen), and /socoma veneta (data compiled from Hall 1928 pp. 223 — 224, Jackson & Dimas 1981, and personal observation of Hartman 3830). Nanthocephalum humile Hartman & Funk 4127 Isocoma veneta ABIT prostrate, perennial herb sprawling, perennial herb shrub YOUNG STEMS reddish to purple, villous — reddish to purple, villous green, glabrous to puberulent Leaves linear to narrowl lanceolate to oblanceolate to obovate- oo linear-lanceolate spatulate-oblong MARGIN usually er 1—3 salient teeth/side 2—5 salient teeth/side CaprruLa solitary 4—8, pedunculate, in 4—8, + sessile, in corymbose clusters corymbose clusters PrEDUNCLES villous, bracteate villous, bracreare pubescent but not villous, ebracteate INVOLUCRES hemispheric hemispheric broadly turbinate Height 4.43—6.4 mm 5-6 mm 5 —8 mm Wipr 5.7 -6.9 mm 7—9.5 mm 4-6 mm PHYLLARIES not resinous not resinous resinous Recerrac es reticulate, scales none alveolate, alveolae alveolate, alveolae rimmed by scales rimmed by scales Ray Fiorers 14— 32, pistillate 12— 15, hermaphroditic ) (anthers rudimentary) Coroiia Tunes trichomes is densely villous ——— and uniseriate Disk Frorers 29 —46 e “" 32—40 15 — 25 (30) COROLLA Lenarn 2.9—4.6 mm 4—5.7 mm 5S —7 mm Coroiia Tunes trichomes glandular trichomes glandular and glabrous or trichomes uniseriate sparse, uniseriate Styie-Brancu APPENDAGES papillae elongate, papillae elongate, papillae short, attenuate rounded rounde ACHENES glabrous, sparsely silky-villous, densely silky-villous, . brown tan light tan LENGTH 1—2.4 mm 19-3 mm 1.8—4 mm Pappus none or low scaly crown bristles bristles LENGTH 0 15-4 mm 3-6 mm POLLEN STAINABILITY 99.2% 51.4% 98% rar former bed of Lago Texcoco at kilometer post 7, WSW of Texcoco, 10 Jan 1978). It fits closely the description of Hartman & Funk 4127 except the plant is older and most of the pubescence has been lost and the ray corolla ts shorter (tube 2.5 — 3 mm long; lamina 2.7 — 3 mm long) and either has a narrow, adaxial lamina nearly equalling the abaxial one or is normal in this respect. Both of these putative hybrids were found to have developed em- bryos in only 10% of the expanded achenes (1 of 10; limited number avail- able). Pollen stainability was also much lower in these specimens than in Hartman & Funk 3027; Castillo and Vasquez 3003 had 28.1% stainable pollen (307 of 1090 grains observed), and I/tis et al. 802 only 13.9% (145 of 1042 grains observed). Jackson's (1966) hypothesis that taxa of Astereae having goblet-shaped disk corollas are related has certainly been supported by his and others’ more recent work, including the present paper. However, if Haplopappus phyllocephalus DC., H. annuus (Rydb.) Cory, and H. aureus (the “phyllocephalus group” of Hartman 1976, 1990) were to be placed in Isocoma (Haplopappus sect. lsocoma sensu Hall 1928) based on the hybrids reported by Jackson & Dimas (1981), then Xanthocephalum sensu Lane (1983) should also be accrued, based on the Xanthocephalum humile X Lsocoma (Haplopappus) veneta hybrid reported here. There are strong Pe logical and chloroplast DNA restriction site data (D. Morgan, Y. Suh, Simpson, M. Lane, unpubl. data) that suggest that the monotypic a anodoria (Robinson 1892) is very closely allied with I[socoma and Xanthocephalum. Further, Xanthocephalum is clearly related to Grindelia (in- oe Prionopsis, Morgan, Suh, Simpson, Lane, unpubl. data) to which Olivaea (De Jong & Beaman 1963) is also allied (Lane 1982). Although the experimental and natural hybrids and other data discussed here certainly do indicate relationship of these taxa, we believe that the “lumping” of the several genera, before the detailed morphological and DNA-systematic investigations currently underway (Nesom et al. 1990; Lane, unpubl. data; Y. Suh and B. Simpson, pers. comm.) are completed, would result in a very large genus that would be not only systematically uninformative but also a nomenclatural nightmare. If all were joined to Hall’s (1928) Haplopappus (in which case the generic name would be Xanthocephalum based on priority) as has been suggested by some workers, then it would follow that yet other genera of Astereae should also be united with it. Ultimately, the collapse of most if not all of the tribe into one ora very few genera would result. Such a situation would obscure rather than clarify the phylogeny of the Astereae; for this reason we continue to recognize distinct genera, even though their members may occasionally hybridize. 328 ACKNOWLEDGMENTS This study was supported by NSF grant number BSR-850863 1 to MAL and an Ohio State University Graduate School postdoctoral fellowship to RLH during 1976—77, when the plant was collected. We thank B. L. Turner for discussion of the plant, R. C. Jackson, G. Nesom, J. L. Strother and an anonymous reviewer for their comments on the manuscript, J. Panero for translating the abstract, G. Nesom for bringing to our atten- tion the two additional hybrids, and V. A. Funk for assistance in the field. REFERENCES DE JONG, D. C. D. and J. H. BEAMAN. 1963. The genus Olivaea (Compositae- Astereae). Brittonia 15:86 —92 GOTTLIEB, L. D. 1984 list 123; eee lee HALL, H. M. . The genus Haplopappus, a phylogenetic study in the Compositae. ubl. Carnegie ee Wash. 389: vii + 386 HARTMAN, R. L. 19 . Genetics and morphological evolution in plants. Amer. Natura- 9 pp. 976. A conspectus of Machaeranthera (Compositae: Astereae) and a biosystematic study of section Blepharodon. Ph.D. dissertation, Univ. Texas, Austin. 1990. A conspectus of Machaeranthera (Asteraceae: Astereae). Phytologia 68:439 — + 465. M. A. LANE and G. K. BROWN. 1987. Haplopappus 1: Hall’s illusion. mer. J. Bot. 74:735. rae R. C. 1966. Some intersectional hybrids and relationships in Haplopappus. Univ. Kansas Sci. Bull. 46:475 — 485. and C. T. DIMAS. 1981. Experimental evidence for systematic placement of the Happs pylopbaas complex (Compositae). Syst. Bot. 6:8 —14 ag Ve 2 LOT. Soil on the research area was a Delnorte very gravelly loam (loamy- skeletal, mixed, thermic family of shallow Typic Paleorthids) (Rives 1980). These are calcareous, light colored, very gravelly soils with indura- ted caliche within 50 cm of the surface. The research area was surveyed in July 1986 and guayule shrubs were categorized into five distinct morphological groups based on growth habit, leaf shape and number of teeth, and branching of the peduncle. Fifty plants of each group were randomly selected and marked. Shrub height and two canopy diameter measurements were recorded. Ten leaves and peduncles were randomly collected from each plant, placed in a plant press, and returned to the laboratory. Terminology used in describing leaf characters follows Radford et al. (1974). The leaf shape and number of teeth on each margin, leaf length, and le w width were recorded. Each peduncle was measured and the number of branches denoted. In March 1987 and 1988, one branch from each shrub was harvested for resin and rubber analyses. The branches were air dried, defoliated and ground in a Fitzmill Comminutor with a 2.36 mm screen. Resin and rubber contents were determined according to the procedure outlined by Black et al. (1983). Average plant height and canopy diameter, leaf length and width, anc peduncle length are reported as the mean + standard error. Resin a — | oma ow rubber values were analyzed by analysis of variance and the means were separated by Tukey’s Studentized Range (HSD) Test (@ = 0.05) RESULTS Guayule plants in the five morphological groups varied considerably in growth habit (Fig. 1). The dense, intricately branched canopies of shrubs in Group I contained fine, tapered, smal Mer diameter stems similar to Group IV. Stems merged gradually into a peduncle, which branched two to three times. The branches were about the same length as the peduncle (Fig. 2). Leaves in Group I were smaller than other groups (Table 1). Leaf shape was usually oblanceolate to obovate, and the margins were coarsely toothed with two to four teeth (Fig. 2). Group I shrubs consistently produced lower rubber contents than Groups II, HI, and [VY (Table 2). Group II shrubs were the tallest, reaching a mean height of 48 cm (Table 1). The canopies were open with minimal branching, and stem diameter was greater than in other groups. Unlike plants in Groups I and IV, there was an abrupt termination of the stem at the base of the peduncle. The naked peduncle generally branched once (rarely two times) with the branches extended beyond the peduncle (Fig. 2). Rubber content was significantly greater in Group II shrubs, and ranged from 7.2 to 13.1% in 1987 and from 6.2 to 12.0% in 1988. 334 FIG. 1. Growth habit of guayule plants in (A) Group 1, (B) Group I, (C) Group HI, (D) Group IV, (E) GroupV, and (F) mariola. Group HI shrubs followed a low growth habit and resembled Group I in height and canopy diameter (Table 1). However, in Group II, the diverg- ing system of larger branches resulted in a symmetrical, closely branched canopy, not the interwoven system as in Groups I and IV. Like Group II, the peduncle usually branched once (Fig. 2), the branch extended beyond the peduncle, and the distinction between stem and peduncle was abrupt. Leaves were intermediate in size compared to the other groups, and leaf shape and margin characteristics matched those in Groups II and V. 335 Taste 1. Morphological characteristics of guayule shrubs within five morphological groups. Shrub Peduncle Leaf Group Height Diameter Length Branches Length Width a CCI) (cm) woe (GM) I 39 + 0.9! 54 + 1.3 15.4 + 0.1 2-3 3524 0.03 1.1 + 0.01 I] 48 + 0.9 57 + 1.2 14.0 + 0.1 0-2 5.1 + 0.04 1.2 + 0.02 Il 39 + 0.8 5221.2 13.44 0.1 0-1 4.5 + 0.03 0.9 + 0.01 IV 43 + 0.7 64 + 1.6 19.6 + 0.1 2-3 45+ 0.04 0.9 + 0.01 Vv 46 + 1.1 42 + 1.2 14.8 + 0.1 0-1 5.6 + 0.04 1.0 + 0.01 ' Mean * standard error. FIG. 2. Leaf and peduncle morphology in (A) Group I, (B) Group I, (C) Group HI, (D) Group TV, and (E) Group V. Canopy characteristics of Group IV shrubs were similar to Group I and included: (1) close, interwoven network of stems, (2) fine, tapered, small diameter stems, and (3) gradual transition of stem to peduncle. Leaf size was comparable to Group III; however, leaf shape was narrowly elliptic to elliptic (Table 1). The growth habit and branching characteristics in Group V were similar to Group II (Fig. 1): plants were erect with an average height of 46 cm (Table 1); canopies were open with minimal branching; and stems termina- 336 ted abruptly at the base of the peduncle. Corresponding to Groups II and Ill, the peduncle generally branched only once with the branch extending above the peduncle. Mean rubber content of Groups IH, II], and V was significantly greater than Groups I and IV (Table 2). DISCUSSION Lloyd (1911) stated that the monopodial growth of the guayule seedling was terminated by the development of the first inflorescence and followed by the rapid growth of several of the uppermost branches. The growth of these branches was also ended by the formation of an inflorescence. Thus, a constantly divaricating system of stems was produced, which resulted in a symmetrical, closely branched shrub. Through the failure of some branches to develop, irregular forms were often observed and attained a greater height than the symmetrical plants. Groups I] and V were readily discernable in the field, and included upright, erect shrubs with less rebranching than other groups. Plants in Group II] were low-growing with the symmetrical, closely branched growth habit. As guayule leaves mature, they are characterized by a single tooth loca- ted near the middle of one margin (Lloyd 1911). Subsequently, a tooth appears on each margin, and a second pair can develop about halfway between the original two and the apex. The guayule stem, unlike mariola, terminates abruptly at the base of the peduncle, and the peduncles general- ly branch only once near the tip. The morphology of Groups H, III, and V was similar to these typical P. argentatum characters: (1) one to two teeth on either leaf margin, (2) peduncle branching one or two times with the branch extending beyond the peduncle, and (3) stems terminating abrupt- ly at the base of the peduncle. Tair 2. Average resin and rubber content of guayule shrubs within five morphological groups harves- ted in March 1987 and 1988. Resin Content Rubber Content 1987 1988 1987 1988 Group Mean Range Mean Range Mean Range — Mean Range (% (%) I 8.0b! S.9-10.4 7.9b 5.4-12.2 56 3.6-8.5 4.5e 1.9-8.0 Il 8. 2ab S.5-11.1 8. 1b 5.3-10.8 10.5a 7.2-13 da 6.2-12.0 II 1.6-9.4 6.3¢ 4.9-9.3 8.8b 5.0-11.3 8.7b 4.6-12.0 IV 8.4a 7.2-9.8 8.8a 6.8-10.6 6. lc 4.1-8.] 6.0d 3.7-7.9 Vv 7.9b 4.6-9.8 8. 1b S.1-11.0 8.8b 4.2-12.9 7.6¢ 3.1-11.8 ' Means within columns followed by the same letter are not significantly different (a = 0.05). oie, Groups I and IV, with dense, profusely-branched canopies, exhibited the mariola manner of growth, and apparently resulted from the introgres- sion between guayule and mariola. Mariola stems, like guayule, terminate in an inflorescence, but are more slender and support short branches or spurs which are more numerous (Lloyd 1911). This manner of growth results in a close interweaving of stems, in striking contrast to guayule. Leaf morphology in Groups I and IV was intermediate between that of guayule and mariola. Leaves were oblanceolate/obovate and narrowly el- liptic/elliptic in shape, and not the lanceolate/ovate shape typified by Groups II, HI, and V. Morphology of the peduncle in the two groups resembled that of mariola. The peduncles usually branched two to three times and the branches were about the same length as the peduncle Results of this study confirm that guayule plants with morphology similar to P. argentatum (Groups IH, HI, and V) produced the highest rubber content. Selection should be concentrated in these groups with rubber con- tents of over 10%. Shrubs with the erect growth habit consistently yielded the greatest rubber content among the five groups. Shrubs with leaf and inflorescence morphology and growth habit similar to mariola should be avoided when screening plants for high rubber-bearing potential. ACKNOWLEDGEMENTS Mr. Darrell W. Ranne, Mr. James M. Harbour, Jr. and Ms. Brenda J. Brown assisted with data collection. Research support was provided by the USDA/CSRS Native Latex Grant and Bridgestone-Firestone, Inc. REFERENCES BLACK, L.T., G.E. HAMERSTRAND, ES. NAKAYAMA, and B.A. RASNIK. 1983. Gravimetric analysis for a aun the resin and rubber content of guayule. Rubber Chemistry Technology 56:367 — —— E Enh, DSeg and: Me C. JOHNSTON 1979. Manual of vascular plants of Texas. Texas . Found., Renner, Texas. 1881 a Re M. A. and JAROY MOORE. 1987. Guayule: A rangeland source of natural rubber. Rangelands 9:99 — 102 AMER, HERBERT H. 1946. “the evaluation of individual plant selections from a nacue) aaa of guayule (Parthenium argentatum Gray). J. Amer. Soc. Agron. LLOYD, E E. 191 1. Guayule ea areentatum Gray) - A rubber plant of the Chihu- ahuan Desert. Publ. Carne nst. Wash. 139. 213 pp. MEHTA, I.J. 1982. Stem es of — argentatum and P. incanum and their nae hybrids. Amer. J. Bot. 69:502 2 MEHTA, I.J., S.P DHILLON, and G.P. ee ON. 1979. Trichome morphology as an ee of high rubber bearing ee ies argentatum Gray) plants in native populations. Amer. J. Bot. 66:796— 338 RADFORD, A.E., W.C. DICKINSON, J.R. MASSEY, and C.R. BELL. 1974. Vascular plant systematics. Harper & Row, New York, 891 pp. RIVES, J.L. 1980. Soil survey of Pecos County, Texas. USDA Soil Conserv. Serv., Temple, yp. Texa ROLLINS, i C. 1944. Evidence for natural hybridity between ee (Partheninm argen- tatu) ha ae (Partheninm incanum). Amer. J. Bot. 31:93 — ae ee -— . 1950. The guayule rubber plant and its relatives. Contr. Gray Herb., No. 2 pp. ak Jibs ane. FAC, Is REGG: ir Variations in rubber concentration of native Texas guayule. Hort. Sci. 17:742 — CLASSIFICATION AND SYSTEMATICS OF EASTERN NORTH AMERICAN VITIS L. (VITACEAE) NORTH OF MEXICO MICHAEL O. MOORE Botany Department University of Georgia Athens, GA 30602, U.S.A. ABSTRACT Eastern North American Vitis, north of Mexico, as circumscribed here consists of two subgenera: Vitis and Muscadinia (Planchon) Rehder. Subgenus Mascadinia consists of a single species with two varicties. Subgenus Vets Is further divided into five . Series Aestivales and Cinerescentes both consist of a single species, the former with Ce varieties and the latter with four varieties. Series Cord:foliae, Labruscae and Ripariae each contain three species. Three taxa previously recognized as species are regarded as hybrids: V. champinit, V. X doaniana, V. X novae-angliae. Keys, Heese ae synonomies and typifica- tions are included. INTRODUCTION The genus Vitis in North America has long been considered difficult from a systematic standpoint and has been largely ignored by North American systematists since the 1930's. The major classifications of North American Vitis (Planchon 1887, Munson 1909, Bailey 1934) are discor- dant in defining species and subgeneric groupings (Barrett et al. 1969), with the latter two treatments being the most widely accepted (Comeaux 1984). Galet (1967) proposed a monograph of the worldwide genus, but his treatment of the North American species is a compilation of the taxa listed by Munson and Bailey with a few minor revisions. The subgeneric groupings proposed by Galet sharply contrast with those proposed by Munson (1909) and Bailey (1934). Indeed, several authors have stated a need for a thorough taxonomic and nomenclatural revision of North American Vitis (Brizicky 1965, Radford et al. 1968, McGregor 1986). Comeaux (1984) represents the most recent classification of North American Vitis and is based on Munson’s (1909) treatment. However, Comeaux (1984) studied in detail only those taxa native to North Carolina, with the remainder of his classification being derived from a general review of previous literature. Comeaux’s (1984) classification was also never pub- lished, but rather a different classification was followed by Comeaux et al. in 1987. Several other recent studies have also provided information of im- SipA 14(3):339 — 367. 1991. 340 port co the systematics of North American Vstis, either through the use of experimental studies or as taxonomic treatments of portions of the genus (Barrett et al. 1969, Comeaux 1987a, 1987b, Duncan 1975, Matthews 1960, Moore 1985, 1987, 1988, 1989, Moore and Giannasi 1987). Thus, renewed interest in Vitis systematics has resulted in a foundation upon which a modern classification of North American Vitzs can be structured. The classification presented here is the result of a revisionary study that employed phenetic analyses of both morphological and foliar flavonoid data as well as extensive field work (Moore 1990). This study, however, excludes the members of series Occsdentales Munson sensu Munson (1909) and Comeaux (1984) (distributed largely west of the Rocky Mountains) as well as four Mexican and South American members of series Cinerescentes and one Mexican member of subgenus Muascadinia (sensu Comeaux 1984), The proposed classification is in general agreement with Comeaux’s (1984) treatment, but several nomenclatural changes are proposed. MATERIALS AND METHODS Herbarium specimens were borrowed from ALU, AUA, BH, BM, C, CM, CU, FE FLAS, FSU, GH, IND, KY, MICH, MINN, MO, MOR, NA, NCU, NHA, NLU, NO, NY, OKL, OKLA, OS, PH, SA, SMU, SRSC, TENN, TEX, TTC, UARK, UNA, UNM, US, USE VDB, VT, WIS, WVA (acronyms according to Holmgren et al. 1981). All taxa were observed and collected in the field during various trips from 1984 — 1989. These specimens, along with those housed at GA, were studied during the course of research. The method by which nodal diaphragm measurements were taken in this study 1s necessary of brief discussion. In several previous treatments of the genus (e.g., Steyermark 1963; Duncan 1975), the width of nodal diaphragms was used to discriminate between taxa. However, in many such treatments, no indication is given concerning the age of wood from which diaphragm measurements were taken. In wood one year old or older, the nodal diaphragms are frequently wider than in sections made from the current years growth. In this present study, all diaphragm measurements were made from current years growth. Also, measurements were made using a dissecting microscope equipped with an ocular micrometer. In using the following keys to assist in the identification of the native grapes, emphasis must be placed on the use of combinations of characters, as a single character is frequently insufficient. The morphological variation in the native grapes is considerable, but when several characters are consi- dered, correct identifications can be achieved with little difficulty. 341 TAXONOMIC TREATMENT VITIS L. Sp. Pl. 2:230. 1753. — Lecrorypr: V. vinifera L., LINN (as IDC microfiche!). Deciduous woody vines or viny shrubs climbing by tendrils. Bark ex- foliating in strips, lenticels inconspicuous or absent (subgenus Vefzs) or adherent with prominent lenticels (subgenus Muascadinia). Pith brown, in- terrupted by nodal diaphragms (subgenus Vrf/s) or continuous through nodes (subgenus Muascadinia). Tendrils bifid to trifid (subgenus Vefzs) or unbranched (subgenus Mascadinia), present opposite only two consecutive nodes or at three to many consecutive nodes (V. fabrusca, V. X novae- angliae). Branchlets of the season terete to angled, glabrous to densely pubescent. Leaves petiolate, blades simple, lobed or unlobed, palmately veined, cordate to orbicular or reniform, toothed to merely scalloped, often mucronate, bases cordate to less often truncate, glabrous to sparsely or densely pubescent beneath, glabrous to slightly pubescent above. Stipules -aducous, 0.5 —7 mm long, promptly deciduous. Growing tips glabrous to densely pubescent. Inflorescence thyrsoid-paniculate, present opposite only two consecutive nodes or at three to many consecutive nodes (V. labrusca, V. X novae-angliae). Flowers pedicellate, functionally unisexual, lants polygamodioecious. Calyx minute, fused into a collar at the base of —9) apically united petals, — iS) the flower, essentially absent. Corolla of 5 ( 1—3 mm long, separating basally at eet and falling from the plant as a unit. Stamens 5 (43—9), filaments erect in staminate flowers, 2— 7 mm long, reflexed to less commonly absent in pistillate flowers; anthers dor- sifixed, valvate, introrse, ca. 0.5 mm long. Nectariferous intrastaminal disc of five more or less eee glands alternating with the stamens. Pistil 1, 0.5 —2 mm long, ovary 2 (3 — 4)-locular, each locule with two ovules; style very short; stigma capitate. Fruit a pulpy |—4 seeded berry. Seeds obovoid to pyriform, 3—8 mm long, the ventral surface with two long- itudinal grooves on either side of the attached funiculus (raphe), the dorsal surface with a groove running its length, becoming wider toward the cen- ter, forming a circular structure (chalaza) that is either sunken or raised. Considered in this treatment are |2 species and 9 varieties, distributed throughout the United States and Canada, largely cast of the Rocky Mountains. Three hybrid taxa are also found in eastern North America. KEY TO THE SUBGENERA AND SERIES Tendrils simple; bark adherent with prominent lenticels —s ; pith continuous ens OG CS:a fc ding aah Reve nite ws a eb aera ere et Subgenus Mascadinia Tendrils bifid to crifid; bark shedding, the lenticels an CenRD Nene: pit nee by dhiaphtaemS: at DOGS. 4 icccuneyeene sea ee dey ae Vitis) — 342 Nm Nh Leaves glaucous beneath; nodes often glaucous............... Series Aestivales Leaves not glaucous beneath; nodes not glaucous...........0....0..0.00-- 3 Branchlets of the season angled, pubescent with arachnoid or hirtel- lous trichomes, or both, varying to glabrate; mature 3 or 4 seeded berries less than 8 mm in diameter; nodes frequently banded with COA EAC) atence liee Bead eon a Peden ox Series Cinerescentes Branchlets of the season more or less terete, glabrous or pubescent; mature 3 or 4 seeded berries usually greater than 8 mm in diameter; nodes epee! not banded with red pigmentation..................0... 4 4 es heavily arachnoid pubescent beneath, concealing the leaf Oe eee but not always the veins; mature fruits greater than 12 TO CUA CEE sag cy ee eae meee eee aed Series Labruscae A a + ; Bs t. Leaves glabrous to slightly arachnoid pubescent beneath, not concealing the intervein area of the underleaf surface; mature berrtes-less than. 12 qin iaciameter ce w sels ackeweeh anaes eed ed 5 5. Growing tips more or less enveloped by enlarging, unfolding leaves; stipules large, usually greater than 3 mm long; nodal diaphragms usually less than | mm in diameter ...... Series Ripariae 5. Growing ae not enveloped by Swi gs unfolding leaves; stipules small, usually less than 3 mm_ long; nodal Pree usually greater than | mm in ae .. Series Cordifoliae ARTIFICIAL KEY TO SPECIES AND HYBRIDS Tendrils simple; bark adherent with prominent lenticels; pith continuous CONS Odea laeediaee oe kale e hare ede Oabe cece tees aes V. rotundifolia Tendrils bifid to trifid; bark shredding, the lenticels inconspicuous; pit interrupted. by todal divphragin’ 4440452062) ee ebwws Sine Ve uiGa douse eee Ran 2 2. Mature leaves glaucous beneath; nodes often glaucous............. V. aestivalis 2. Mature leaves not glaucous beneath; nodes not glaucous................... 3 3. Tendrils or inflorescences present at three to many consecutive nodes 4 4. Leaves densely pubescent beneath, concealing the leaf under- surface but not always the — nodal diaphragms usually great- er than 1 min Camere 5 3 ow eee eae Se RE eR oe een V. labrusca 4. Leaves ae to anodes a pubescent beneath, the leaf under- surface visible on mature leaves; nodal diaphragms usually less than | mm in diameter ...........0.......00000. V. X = novae-angliae eS) Tendrils or inflorescences present at only two consecutive nodes .......... 5. Leaves densely pubescent beneath, concealing the leaf under- surfaces but not always the veins; fruits greater than 12 mm in GIG fhe ce eena 260) Coa AE LEN ee ewan nee eahs 6 6. Stipules greater chan | mm long; leaves frequently concavely HOIGG date cto lee Sarde ch aea swans eoponaty ede ee V. mustangensis 6. Stipules less than 1 mm long; leaves not concavely folded eral ey estates cts, Womeentae a! ae diet Gude geaaudd GS Eee ea sa V. shuttleworthii 5. Leaves glabrous to moderately pubescent beneath, the intervein area of leaf undersurfaces visible on mature leaves: fruits less or greater than 12 mm in diameter... 260.4040 00e.es00e0i4bvanesads 7 7. Leaves reniform, glabrous beneath at maturity; tendrils absent or present only opposite the uppermost nodes 7. Leaves cordate to cordate ovate, glabrous to pubescent beneath at maturity, tendrils present opposite most nodes 8. Nodal diaphragms less than 1 mm wide, usually less than 0.5 mm wide; growing tips enveloped by enlarging, un- folding leaves 9. Growing tips slightly to densely pubescent; branchlets of the season slightly to densely arachnoid aa inflorescences usually less than 8 cm ong 9. eae tips glabrous to slightly pubescent; branchlets of the season usually lacking arachnoid eb inflorescences usually greater chan 8 cm oo Nodal cee ee greater than | mm wide; oe tips not enveloped by enlarging, unfolding leav 10. Branchlets of the season angled, aon and/or hirtellous pubescent, varying to glabrate; mature 3 or 4 seeded berries less than 8 mm in diameter; nodes 343 frequently banded with red pigmentation ......... V. cinerea 10. Branchlets of the season more or less terete, glabrous or arachnoid pubescent; mature 3 or 4 seeded berries usually greater than 8 mm in diameter; nodes usually not banded with rec jon 11. Mature 3 or 4 seeded berries greater than 12 mm in fon leaves arachnoid — pubescent bene 12. ee moderately to heavily arachnoid pubes- cent beneath, also with hirtellous trichomes PIFMENCACIONS 4 saw dawg eke Ye along the veins; fruits glaucous ....... ' X doantana rs ~~) 2. Leaves only slightly arachnoid pubescent beneath and lacking hirtellous trichomes; fruits not glaucous 11. Mature 3 or 4 seeded berries less than 12 mm in diameter; leaves usually lacking arachnoid pubes- eénée beneath. cage ess eee Le Dae Pewee es 13. Nodal diaphragms greater than 2.5 mm wide, leaf apices usually long acuminate; branchlets f the season with a purplish red cast Nodal diaphragms less than 2.5 mm wide; leaf apices usually acute to short acuminate; branchlets of the season gray to green or brown or with purplish pigmentation only on one side of the branch oO tad 14. Berries usually with — lenticels; — in- fructescences with less than 12. berries; growing tips slightly to densely pubes- cent, leaf blades usually less than 8 cm ied wean S ectgus ie V. Xx champinit 344 long; branchlets of the season usually slightly arachnoid pubescent .......... V. monticola sae without le neicels; infructescences mo ne an 12 berries; growing tips ee to slightly ens leaf blades usually greater chan 9 cm long; branchlets of the season glabrous.............. _V. vulpina Subgenus MUSCADINIA (Planchon) Rehder, Man. Cult. Trees 601. 1927. Section Muscadinia _ non, DC Monogr. Phan. 5:323. 1887. Genus Muscadinia (Planchon) Small, Fl. SE U.S. 756. 1903. — Tyee species: V. rotundt- folia Michaux. Vitis ROTUNDIFOLIA Michaux, Fl. Bor.-Amer. 2:231. 1803. High climbing vine, branchlets of the season terete to slightly angled. Bark of younger woody stems with evident lenticels, that of older stems tight, not exfoliating, that of still older stems exfoliating in plates, pith rown, continuous through nodes, diaphragm absent. Tendrils un- branched, a tendril or inflorescence present at only 2 consecutive nodes, nodes not glaucous, but often banded with red pigmentation. Very young, rapidly growing stems and leaf surfaces usually with thin, loose, grayish arachnoid pubescence or with dense, rusty, arachnoid pubescence at the nodes of the stems and pinkish on leaf surfaces, the pubescence eventually deciduous. Leaves with petioles mostly as long as the blades, glabrous to glabrate; blades cordiform to nearly reniform, very rarely lobed; margins crenate to dentate, apices very short acuminate; upper surface of mature leaves glabrous and lustrous, lower surface not glaucous, but glabrous or pubescent with few to many hirtellous trichomes along the veins and in their axils; stipules | — 2 mm long. Panicles 3 — 8 cm long, rarely longer, usually more or less globose in outline, infructescences with less than 25 berries (or pedicels); 3 or 4 seeded berries 8 — 25 mm in diameter, generally black or purplish, occasionally bronze when ripe, glaucescent, with tan, circular lenticels present on the skin. Seeds brown, oval to ellipsoidal, 5—8 mm long. Inhabiting a very wide variety of sites, both upland and well drained and lowland and poorly drained, including intermittently flooded bottom- lands. (DE to KY, s IN, MO, generally southward to FL, e OK and e TX). Flowering in late April to May; fruit ripening in late July to September. The two varieties of this species can be distinguished morphologically based on the following key: _ Mature fruits greater than 12 mm in diameter; infructescences se with less than 12 berries; leaf blades usually more than 5 cm in len Rady Oe Gt een ee Ge oh ee ek eee eee a eo elt V. ere var. rotundifolia 345 1. Mature tae less than 12 mm in diameter; infructescences with mo 12 berries; leaf blades often less than 5 cm long..... Vv. rotundifolia var, munsoniana ViITIS ROTUNDIFOLIA a var. ROTUNDIFOLIA. V. muscadinia Raf., Amer. Man. Grape Vines 16— 17. 1830. Muscadinia rotundifolia (Michaux) Small, Fl. SE U.S. 757. 1903. — Type: a ie uae ad Floridum (Lecrorype, here designated: microfiche IDC Michaux, no. 122, photo 20! P). — Synrypr: microfiche IDC Michaux, no. 123, photo I! (P). Leaves generally larger than in variety munsoniana, fruits greater than 12 mm in diameter and infructescences with less than 12 berries. Inhabiting a wide variety of sites, both upland and well drained and lowland and poorly drained. (DE to KY, s IN, MO, generally southward to FL, e OK and e TX). Flowering in late April to May, fruit ripening in late July to a aie er specimens examined: ARKANSAS. Hempstead Co.: cree (G ae ORGIA. Brooks Co.: Moore 790 (GA), Clarke Co.: Moore 1011 (GA). FLORIDA. Franklin > Moore 814 (GA); Gilchrist Co.: Moore 407 (GA); Jackson Co.: ce 675 (FSU). NORTH CAROLINA. Davie Co.: Moore 249 (GA). OKLAHOMA. LeFlore Co.: en 20589 (GH). SOUTH CAROL INA. Berkeley Co.: Wiegand & Manning 1955 (GH). XAS. Newton Co.: Lundell 11870 (TX). VITIS ROTUNDIFOLIA Michaux var. MUNSONIANA (Simpson ex Munson) M.O. Moore, comb. nov. — BasionyM: V. munsoniana Simpson ex Munson, Proc. Soc. Promot. Agric. Sci. 8:59. 1887. Muascadinia munsoniana (Simpson ex Munson) Small, Fl. SE U.S. 757. 1903. — Type. FLORIDA. MANATEE Co.: collected along Manatee River, 1883, 1885, 1887, ].H. Simpson, cultivated Munson vineyard, Denison, Texas 1890 (Lecrorypr, here designated: PH!). SYNTYPE: (PH!). Similar to var. rotundifolia, but usually with smaller leaves, fruits less than 12 mm in diameter and infructescences with more than 12 berries. ieee a wide variety of sites, but usually found on drier soils. (FL, s GA, s AL). Flowers and fruits virtually all year in peninsular Florida, but in more northern locations flowering in late April to May; fruit ripening late July to September. Representative specimens examined: FLORIDA. Collier Co.: Moore 764 (GA); Moore 759 (GA). Duval Co.: Curtis 4818 (US). Franklin Co.: Moore 815 (GA). Highlands Co.: Skean 2144 (GA). Lake Co.: Moore 401 (GA); Judd 2453B (FLAS). Monroe Co.: Bazley 314 (BH); Moore 769 (GA). Putnam Co.: Mosre 746 (GA). Subgenus VITIS, Series AESTIVALES Planchon, in DC Monogr. Phan. 23. 1887. — Tver species: V. aestivalis Michaux. Viris AESTIVALIS Michaux, Fl. Bor.-Amer. 2:230. 1803. High climbing vine, branchlets of the season terete, tomentose, arach- 346 noid floccose or glabrous. Bark exfoliating in shreds on mature stems, lenticels absent or inconspicuous, pith brown, interrupted by diaphragms at the nodes, diaphragms | —4 mm thick. Tendrils bifurcate, a tendril or inflorescence present at only 2 consecutive nodes, nodes glaucous or not glaucous, not banded with red pigmentation. Leaves with petioles about as long as the blades, glabrate to pubescent; blades cordiform to orbicular, unlobed to 3-shouldered or 3—5 lobed, often deeply so, when lobed the lobes mostly acute, the sinuses rounded to acute; margins crenate to dentate; upper surface of mature leaves glabrous to puberulent, lower surface glaucous with varying degrees of arachnoid, floccose pubescence, when heavy the glaucescence somewhat obscured, the pubescence whitish to more commonly rusty, hirtellous trichomes also eae ae ed present along the veins and as tufts in the vein axils; stipules 1 — m long. Panicles 7—20 cm long, usually narrowly triangular in saline in- fructescences usually with more than 25 berries (or pedicels); 3 or 4 seeded berries 8 — 20 mm in diameter, black, glaucous, without lenticels. Seeds tan to brown, pyriform, 3-8 mm long. Generally found on well drained sites, woodlands of various mixtures, woodland borders, thickets, fence and hedge rows, scrub, stabilized dunes, less often along stream or river banks, rarely in floodplains or lowland woods. (Throughout eastern North America and southern Canada). Flowering in April to June; fruit ripening in July to September. This species is frequently confused with V. cinerea. However, the glaucous leaf undersurfaces, more heavily glaucous, larger berries, terete less evenly pubescent branchlets, preference for well drained, drier habitats and earlier blooming period distinguishes V. aestivalis from V. cinerea. The three varieties of this species can be distinguished morphologically based on the following key: Branchlets of the season heavily arachnoid pubescent; mature 3 or 4 seeded erries usually greater than 14 mm in diameter; stipules usually less than 2) MM, MONG eh. 3 ewe Bek Bien ae wea ee eS V. aestivalis var. lincecumii 1. Branchlets of the season slightly to moderately arachnoid pubescent, or glabrous; mature 3 or 4 seeded berries usually less than 14 mm in diameter; eee weal greater than 1.5 mm long ........... 00000 2 Mature 3 or 4 seeded berries less than 9 mm in diameter, mature leaves ee to glabrate beneath; nodes usually glaucous; nodal diaphragm sea less than 2 mm in diameter .................. V. a a var. bicolor Mat 2 re 3 or 4 seeded berries greater than 9 mm in diameter; mature oe slightly to heavily arachnoid pubescent beneath; nodes usually not glaucous; nodal diaphragms usually greater than mm in GiaMiCtel #2 Buses ence a ee ee ok ee eee et V. aestivalis var. aestivalis 347 VITIS AESTIVALIS Michaux var. AESTIVALIS. V. /abrusca var. aestivalis (Michaux) Regel, Act. Hort. ae 2:396. 1873. V. vinifera var. aestivalts (Michaux) Kunze, 1 Rev. Gen. PI. 32. 1891. — Type: in sylvestris, a Pensylvania ad Carolinum (Lectotype, here samen microfiche IDC Michaux, no. 122, photo 17! P). — Synrype: microfiche [DC Michaux, no. 122, fe 18! (P). V. aestivalis var. sinuata Pursh, Fl. Amer. Sept. 1:169. 1814. V. stnwata (Pursh) G. Don, Gen. Hist. 1:711. 1831. — SyNrvypes: not seen. V. araneosus LeConte, Proc. Acad. Nat. Sci. Phil secon 6272... 1859. Tire: GE 31A. On the banks of the Oconee at Athens, 14 Sep 1850, John LeConte s.n. ee rotypr, here designated: PH!; tsoLecroryPe: : ')) — Synryees: (PH!). V. lincecumii Buckley var. glauca Munson, U.S.D.A. Div. Pomel: Bull, No, 32-7412. 1890. V. fincecumii var. lactea Small, Fl. SE U.S. 755, 1334. 1903. — Type: TEXAS. North Texas, 26 May 1890, Manson s.n. (HOLOTYPE: PH!). V. simpsonti Munson, U.S.D.A. Div. Pomol. Bull. No. 3:12. 1890, non 1887, nom. illeg. V. ie Bailey, Gentes Herb. 3:207 — 209. 1934. V. aestivalis ssp. small- iana (Bailey) Rogers, Proc. Florida State Hort. Soc. 92:289. 1979, nom. illeg. V. aestivalis var. smalliana (Bailey) Comeaux, Sida 12:286. 1987. — Type: FLORIDA. MANATEE Co.: collected originally from Manatee County, n.d., J.H. Simpson s.n. cultivated Denison, Texas, 25 May 1890 (LEcrorypre, here designated: PH!; ISOLECTOTY PES: MO! Vv. ig Small, FI. ‘SE U.S. 756, 1334. 1903. — Type: FLORIDA. Lake Co.: Lake Eustis, 16-30 Apr 1894, Nash 525 (HoLoTyPE: NY!; IsoTYPEs: ae V. gigas Fennel, J. Wash. Acad. Sci. 30:15 — 19. 1940. — Tyrer: FLORIDA. Sebas- tian River, 20 Jul 1938, J.L. Fennel 713 (HoLorype: US!; 2 sheets, | of fruiting branch, | of growing tip). Leaf undersurfaces with varying degrees of arachnoid pubescence, mod- erately to somewhat heavily glaucous, nodes usually not glaucous, nodal diaphragms usually greater than 2 mm wide, mature 3 or 4 seeded berries 9 — 14 mm in diameter. Found in well drained sites, woodlands, woodland borders, thickets, fence and hedge rows, scrub, stabilized dunes, less often along stream or river banks and floodplains and lowland woods. (Piedmont, Coastal Plain, Mountains, Interior Low Plateau, Central Low- lands, coastal MA to se IA, MO, e OK, e TX to FL). Flowering in April to June, fruit ripening in July to September. Representative specimens examined: FLORIDA. Hamilton Co. : us 379 (GA). Ham- pton Co.: Moore 360 (GA). Wakulla Co.: Moore se ees GEORGIA. Decatur Co.: Moore 805 (GA). Montgomery Co.: Moore 346 (GA). DIANA. Martin Cen. Moore 1056 (GA). MISSOURI. Howell Co.: Moore 1027 (GA). ne Carter Co.: Moore 250 (GA). TEXAS. Rusk Co.: Moore 930 (GA). VIRGINIA. Nelson Co.: Moore 834 (GA). Vitis AESTIVALIS Michaux var. BICOLOR Deam, Shrubs Indiana 207. 1924. V. argentifolia Munson, Proc. Soc. Promot. Agric. Sci. 8:59. 1887. V. aesti- valis var. argentifolia (Munson) Fernald, Rhodora 38:428. 1936. — Neoryre, here designated: WEST VIRGINIA. NicHoras Co.: W side of US 19, 1.7 mi S of the 348 Braxton Co. line, 12.4 mi N of jet. with WV 55, 25 Aug 1987, Michael O. Moore 886 (GA!). — SyNntypes: not found. Similar to var. gestevalis, but with leaf undersurfaces glabrous to glabrate and heavily glaucous beneath, nodes usually glaucous, nodal diaphragms only |— 2 mm wide, mature 3 or 4 seeded berries 8 — 9 mm in diameter. Inhabiting a wide variety of well drained sites, woodlands of various mixtures, woodland borders, thickets, fence and hedge rows and scrub. (Blue Ridge, Ridge and ee Appalachian Plateau, n GA and n AL generally north to Canada). Flowering in late May to June; fruit ripening in July to September. Previously known as V. aestivalis var. argentifolia, Fernald (1936), statec that the name var. dicofor cannot be used because this taxon is not the V. bicolor of LeConte and because var. bicolor was never published. However, Deam (1924) did treat this taxon as V. gestivalis var. bicolor, attributing the combination to Britton and Brown. Britton and Brown never made this combination, but rather listed V. bicolor in synonymy under V. aestivalis. LeConte’s V. bicolor is a dubious name that cannot be associated with any currently recognized taxon. Deam (1924) did give a good description of this taxon including characters that distinguish it from V. gestivalis var. aestivalis and thus did feel that it should be treated as a variety of V. aesti- valts. It can thus be concluded that Deam’s use of the name V. aestivalis var. bicolor is a valid publication which has priority over Fernald’s (1936) name on V. aestivalis var. argentifolia. Representative specimens TT CONNECTICUT. Hartford Co.: Moore 851 (GA). GEORGIA. oS Co.: Owen 201 (AUA). ILLINOIS. Tazwell Co.: Chase 3467 (MIN). KENTUCKY. Bell Co.: McF baphend 3019 (BH). NORTH C AROLINA. Avery Co. Moore 254 (GA). OHIO. seg Co.: Tandy 1741 (OS). PENNSYL — EG Co.: Adams 4128 (GH). VIRGI NIA. Patrick Co.: Moore 232 (GA). WEST VIRGIN Nicholas Co.: Moore 886 . A). WISCONSIN. Columbia Co.: ie SoM, i VITIS AESTIVALIS Vat. LINCECUMH (Buckley) Munson, Proc. Amer. Pomol. SOc. co 97. 1886. V. /incecumi? Buckley, Proc. Acad. Nat. Sci. sees 62 451. 1861. — Type: TEXAS. Eastern Texas, 1861, S.B. Buckley 5.n. (HOLOTYPE: US!). Similar to var. gestivalis, but with branchlets of the season more or less densely tomentose, nodal diaphragms generally less than 2 mm wide, leaves more frequently deeply 3 to 5 lobed, berries that are generally larger than 14 mm in diameter and are heavily glaucous, and larger seeds, 7 —8 mm. Vetrs aestivalis var. lincecumii also has an earlier time of anthesis than var. aestivalis and is more drought resistant. Inhabiting well drained sites, woodlands of various mixtures, woodland borders, thickets, fence and hedge rows and scrub. (TX, east of the Trinity 349 River, ¢ TX east of Austin, w LA). Flowering in April, fruit ripening June to September. In the original publication of this name (Buckley 1861), the specific epithet was spelled “/insecomii”, but the holotype has the name spelled “lincecumi?” in Buckley’s handwriting. Munson (1909) determined that this taxon was named after Dr. Gideon Lincecum, and speculated that the spelling “/znsecomi?” probably came through as an error of the typesetter. Thus, in accordance with article 73.1 of the International Code of Bo- tanical Nomenclature, the spelling of this name should be corrected to “/incecumit. Representative specimens examined: LOUISIANA. Bienville Parish. Moore 664 (GA). TEXAS. Cherokee Co.: Moore 931 (GA). Henderson Co.: Lundell & Lundell 9509 (SMU). Leon Co.: Moore 936 (GA). Milam Co.: a 680 (GA). Morris Co.: Correll & Correll 12445 (SMU); Correll & Correll 25409 (NY). Rusk Co.: 7 56465 (SMU). Smith Co.: Shinners 15094 (SMU). Wood Co.: Holmes ae (NLU) Subgenus VITIS, Series CINERESCENTES Planchon, in DC Monogr. Phan. 5:323. 1887. — Tver species: V. cimerea (Engelm. in Gray) Engelm. ex Millardet Viris CINEREA (Engelm. in Gray) Engelm. ex Millardet, Mem. Soc. Sct. Phys. Nat. Bordeaux 2(3):319 — 330. 1880. High climbing vine in floodplains and lowland woods, along stream banks, pond margins and fence rows. Branchlets slightly to distinctly angled (the angling often difficult to sce with the unaided eye), branchlets of the season covered with dense, short, straight (hirtellous) trichomes and/ or thin to dense arachnoid pubescence, varying to glabrate. Bark exfoliat- ing in shreds on mature stems, lenticels absent or inconspicuous, pith brown, interrupted by diaphragms at nodes, diaphragms 1.5 to 3.5 mm thick. Tendrils bifurcate to trifurcate, a tendril or inflorescence present at only 2 consecutive nodes, nodes of branchlets of the season often banded with red pigmentation, nodes not glaucous. Leaves with petioles about as long as the blades, puberulent to pubescent with hirtellous trichomes, thin arachnoid pubescence commonly present as well; blades cordiform, unlo- bed to 3-shouldered, occasionally 3-lobed, the apex acute to more commonly acuminate; margins crenate to dentate; upper surface of mature leaves glabrous to pubescent, lower surface not glaucous, slightly to mod- erately arachnoid pubescent, varying to glabrous, the pubescence mostly whitish; hirtellous trichomes also commonly present along the veins and as small tufts in the vein axils; stipules | — 3 mm long. Panicles 10 — 25 cm long, usually broadly triangular in ae infructescences usually with more than 25 berries (or pedicels); 3 or 4 seeded berries 4 — 8 mm in diame- 350 ter, black, with little or no glaucescence, lenticels absent. Seeds brown, obovoid, 2—4 mm long. Usually found in moist habitats. (s IN tos PA, south to FL, west to TX, north toOK, KN, MO and IL). Flowering in late May to June; fruit ripen- ing in July to October. This species is frequently confused with V. aestz- valis. See the discussion provided under V. aestivalis In recent treatments of the genus (e.g., Radford ef a/. 1968; Godfrey and Wooten 1981), the author citation for V. cinerea is given as Engelm. ex Millardet. Still other treatments (e.g., Steyermark 1963; McGregor 1986) cite only Engelm. as the author citation. Gandhi and Brown (1989), how- ever, use the following: V. cinerea (Engelm.) Engelm. ex Millardet and dis- cuss the reasoning for their citation of authorship. Since this taxon was first published as a variety of V. aestivalis in Gray’s Manual (1867), with the name being attributed to and the description provided by Engelmann, it is clear that the initial citation should be V. aestivalis var. cinerea Engelm. in Gray. Millardet was the first to elevate this taxon to the species level, also attributing the name to Engelmann but providing a description not given sy Engelmann. Thus, the correct citation ts clearly V. cinerea (Engelm. in Gray) Engelm. ex Millardet. To eliminate Gray’s name from the author citation also eliminates the author of the original publication in which the name appeared from the citation, making it quite difficult to trace the nomenclatural history of this taxon. The four varieties of this species can be distinguished morphologically based on the following key: Berries moderately to heavily glaucous; leaf blades glabrous to cane usually less than 10 cm long; central Texas Tee eae . cinerea var. helleri Berries only slightly to not aoa leaf blades pubescent, varying to glabrate, usually more than 10 cm long e Texas east and northward 2 Branchlets of the season aes to en hirtellous pubescent, often with arachnoid pubescence as well less uniformly hirtellous pubescent on veins ............. V. cinerea vat. cinerea Branchlets of season without pre hircellous trichomes (if present, then om by arachnoid pubescence); leaf undersurfaces usually without hirtellous trichomes, or, wh sparsely so ~) t ; leaf undersurfaces usually more or in) en present, only very oe Branchlets slightly to densely arachnoid pubescent; nodes usually not banded with red pigmentation; leaves slightly to densely arach- noid pubescent beneath; Coastal P Branchlets glabrate to ‘only igh usually banded with re cat arieg steed . cinerea var. floridana y arachnoid pubescent; nodes pigmentation; leaves glabrous to as slightly arachnoid pubescent as Piedmont and Mountain Ww om V. cinerea var. baileyana 351 Vitis CINEREA (Engelm. in Gray) Engelm. ex Millardet var. CUNEREA. V. — var. cinerea Engelm. in Gray, Manual ed. 5:676. 1867. — Type: ILLINO- IS. The Engelmann farm, Sep 1867, G. Engelmann s.n. (LECroTYPE, here designa- ted: a ISOLECTOTYPE: MO!). — Syntype: (MO!). V. cinerea var. canescens (Engelm.) Bailey ex Gray, Syn. Fl. N. Amer. 1(2):425. 1897. V. aestivalis var. canescens Engelm., Amer. Naturalist 2:321. 1869. — Type: Mis- sissippit Valley (HOLOTYPE: GH!). Branchlets of the season covered with short, straight hirtellous trichomes, occasionally with arachnoid trichomes as well. Leaf under- surfaces are moderately arachnoid and/or hirtellous pubescent. Inhabiting floodplains, lowland woods, ponds and stream margins. Native to the rich bottomlands of the Mississippi basin. (s IA, s IL, s IN south toe KN, e OK, e TX east to a few scattered localities in AL and panhandle FL). Flowering in late May to June, fruit ripening in July to October. Representative specimens examined: ALABAMA. Lowndes Co.: Moore 734 (GA). ARKANSAS. Ashley Co.: Moore 349 (GA). Marion Co.: Moore 300 (GA). ILLINOIS. Richland Co.: Moore 1053 (GA); Schuyler Co.: Moore 1047 (GA). KENTUCKY. Hickman Co.: Moore 284 (GA). LOUISIANA. Bossier Parish. Moore 335 (GA). MISSISSIPPI. Low- ndes Co.: Moore 344 (GA). OKLAHOMA. Pottawatomie Co.: Moore 982 (GA). TENNESSEE. Lake Co.: Moore 278 (GA). Viris CINEREA (Engelm. in Gray) Engelm. ex Millardet var. FLORIDANA Munson, U.S.D.A. Div. Pomol. Bull. No. 3:12. 1890. V. simpsoni Munson, Proc. Soc. Promot. Agric. Sci. 8:59. 1887. aries Small, Fl. SE U.S. LORIDA 755. 1903. — Tyre: F . Manatee Co.: originally from Manatee River, J.H. Simpson s.n., cultivated in vineyard of T.V. Munson, 990 (LECTOTYPE, here designated: MO!; isoLecrotyPE: MO!). — Synrypes: (BH H!). V. sola Bailey, Gentes Herb. 3:203. 1934. V. aestivalts ssp. sola eis Rogers, Proc. Florida State Hort. Soc. 92:289. 1979, nom. illeg. — Type: FLORIDA. Swamp near eens 20 Sep 1894, A.H. Curtiss 4791 (Lecrorype, here designated: NY!, H!; tsotecrorype: NY!). — Paratypes: (MO!, as photo BH!). V. eo ssp. divergens Rogers, Proc. Florida State Hort. Soc. 92:289. 1979, nom. ifleg. Similar in general appearance to V. cinerea var. cinerea but differs from var. cinerea by having branchlets that are arachnoid pubescent, often densely so, and generally lacking the dense hirtellous pubescence characteristic of V. cinerea vat. cinerea. The leaf undersurfaces of V. cinerea var. floridana also tend to be more densely arachnoid pubescent than is common in V. cinerea vat. cinerea. Common in floodplains, lowland woods, stream and pond margins. (Coastal Plain of VA, SC, NC, GA, FL, ALand MS). Flowering in late May to June; fruit ripening in July to October. This variety is frequently con- fused with V. aestivalis. See the discussion provided under V. aestivalzs. 352 Comeaux and Fantz (1987) provide a discussion of the somewhat convo- luted nomenclatural history of this taxon. Representative specimens examined: ALABAMA. Lowndes Co.: Moore 732 (GA). FLORIDA. Collier Co.: Moore 763 (GA). Gadsden Co.: Moore 804 (GA). Jefferson Co.: Moore 591, (GA). Taylor Co.: Moore 402 (GA). Walton Co.: Moore 202 (GA). GEORGIA. Early Co.: Moore 261 (GA). Randolph Co.: Moore 268 (GA). Telfair Co.: Moore 382 (GA). Wilkinson Co.: Moore 381 (GA) Vitis CINEREA (Engelm. in Gray) Engelm. ex Millardet var. BAILEYANA (Munson) Comeaux, Castanea 52(3):212— 213. 1987. V. virginiana Munson, U.S.D.A. Div. Pomol. Bull. No. 3:3, 14. - nom. aes V. baileyana Munson, Leaflet, 20 Jun 1893. — Type: VIRGINIA. fountain valleys, southwest Virginia, 1890, J.G. Wertz s.n. y) ee PE, ee ieee PH!; isoLECTOTY PE: Similar in general appearance to V. cznerea var. floridana, but differing in having branchlets of the season glabrous to glabrate, nodes usually banded with red pigmentation and lower leaf surfaces glabrous to glabrate. Inhabiting a variety of habitats but more common in moist soils, flood- plains, lowland woods, stream and pond margins. (Piedmont and Mountains, GA and AL to se IN, s OH and s PA). This taxon intergrades into V. cinerea var. floridana along the fall line between the Piedmont and Coastal Plain in AL, GA, NC, SC and VA. Representative specimens examined: GEORGIA. re Co.: Moore 171 (GA); ce 190 (GA); Moore (GA); Jones Co.: Moore 259 (GA ORTH CAROLINA. 10. Moore 238 (GA). Yadkin Co.: ie 241 (GA). ou H CAROLINA. Sparano Moore 819 (G . TENNESSEE. Lawrence Co.: Sharpe et al. 9700 (T ae INIA. Albemarle Co.: ae (BH). Roanoke Co.: Wertz 5.1. (MO). ~ o) ~ Viris CINEREA (Engelm. in Gray) Engelm. ex Millardet var. HELLERI (Bailey) M.O. Moore, comb. nov. — Basionym: V. elias var. ao saree gh Syn. Fl. N. ee 1:424. 1897. V. hellert (Bailey) Small, Fl. 754 4. 1903. — ‘Wee: TEXAS. Kerr Co.: 1600 — 2000 ft, aes ae oe ear 1750 (Lecrorypr, here designated: BH!; tsotecroryres: BH!, photos BH!). as V. berlandieri Planchon, Compt. Rend. Hebd. Seances Acad. Sci. 91:425. eon V. cinerea var. berlandter: (Planchon) Comeaux, Proc. Texas Grape Growers 1986. 1987, nom. — — Type: NEW MEXICO and TEXAS, 1834, ii 2412 (HOLOTYPE: PH!). Similar in appearance to V. cinerea var. cinerea, but differing by having berries that are moderately to heavily glaucous, branchlets of the season that generally lack hirtellous pubescence and are not as prominently angled, and leaf blades that are usually less than 10 cm long with under- surfaces that are only sparsely hirtellous pubescent (or glabrate). Inhabit- ing a variety of moist habitats, floodplains, lowland woods, stream and 529 pond margins. (TX, most common on the Edwards Plateau, but also found in the Cross Timbers and Prairies and the Blackland Prairies). This variety intergrades with V. cénerea var. cinerea southwest of the Brazos River (Com- eaux, 1987a). Comeaux (1987a) combined this taxon with V. cznerea as V. cinerea vat. berlandieri (Planchon) Comeaux. However, in doing so, no clear indication of the basionym was given as is required under article 33.2 of the Interna- tional Code of Botanical Nomenclature and thus the name was not validly published. Nevertheless, the name “var. e//er7” is the oldest varietal name attributable to this taxon and therefore must be used if this taxon is recognized at the varietal level and if V. cordifolia var. eller? 1s considered as a synonym of it. Gandhi and Brown (1989), however, did not accept Com- caux’s treatment and preferred recognizing this taxon as a distinct species. Present evidence justifies the treatment of this taxon as a variety of V. cmmered. Bee specimens examined: TEXAS. Bandera Co.: Moore 683 (GA). Coryell : Moore 943 (GA); Moore 944 (GA). Kendall Co.: Moore 682 (GA). Kerr Co.: Correll & ae 17231 (TEX). Real Co.: Moore 691 (GA); Cory 19088 (GH). Travis Co.: Ripperton & Barkley 14522C (OKL). Uvalde Co.: Moore 958 (GA), Moore 689 (GA). Subgenus VITIS, Series CORDIFOLIAE Munson, U.S.D.A. Div. Pomol. Bull. No. 3:7. 1890. — Tver species: Vitis cordifolia Michaux (= V. vualpina L.). Vitis VULPINA L., Sp. Pl. 203. 1753. — Tver: VIRGINIA (HoLoryPe: LINN as IDC enna no. 281.7!). _ ~ /, cordifolia Michaux, Fl. Bor. pe 2:231. 1803. — Lecrorype, here ea eae as 23, photo 3! (P). — Syntype: as microfiche IDC V. age LeConte, Proc. Acad. Nat. Sci. Philadelphia 6:273. 1853. — Type: sINTA. Noitbile, n.d., John LeConte s.n. (Lecrorype, here designated: PH!, ISOLECTOTYPE: PH!). V. as vat. fide Engelm., Amer. Naturalist 2:231. 1869. — SYNTYPEs: not fou Vv. cordifolia var. sempervirens Munson, Rev. ae 5:165. 1896. V. lex Bailey, Gent. Herb. 3:217. 1934. — - FLORIDA. Manater Co.: originally from south Florida, cultivated in ney of ua — 10 May 1890 (LecroryPE, here designated: BH!). — Synrvprs: (BH!). High climbing vine, branchlets of the season slightly angled when very young but becoming terete, very young stems and emerging leaves glabrous to sparsely arachnoid pubescent. Bark exfoliating in shreds on mature stems, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms | — 2.5 mm thick. Tendrils bifurcate, a tendril or inflorescence present at 2 consecutive nodes only, nodes not 354 glaucous, not banded with red pigmentation. Leaves with petioles about as long as the blades, sparsely to moderately pubescent with hirtellous trichomes or glabrous; blades cordiform, often 3-shouldered to shallowly 3-lobed, deeply lobed only on ground shoots; margins irregularly dentate- serrate, bases typically cordate, apices acute to short acuminate; upper surface of mature leaves typically glabrous to very sparsely hirtellous pub- escent, often lustrous, lower surface not glaucous, typically green, with short, straight hirtellous pubescence along the veins and in their axils, varying to more or less glabrous, rarely with very sparse arachnoid pubesc- ence; stipules 1.5 —3 mm long. Panicles 9— 19 cm long, usually narrowly triangular in general outline, infructescences typically with more than 25 berries (or pedicels); 3 or 4 seeded berries 8 — 12 mm in diameter, black, very slightly, or more typically, not at all glaucous, lenticels absent; seeds dark brown, ovoid, 3—5 mm long. In upland, well-drained woodlands of various mixtures, woodland borders, fence and hedge rows, thickets, less commonly in floodplains or lowland woods (se NY to MO and e KN, generally southward to penin- sular FL and nc TX), Flowering in May; fruit aaa July to August. Representative specimens examined: ARK Marton Co.: nae 301 (GA). FLORIDA. Dixie Co.: Moore 317 (GA); Gadsden an ape 798 (GA). NOIS. Shelby Co.: Moore 1050 (GA). MISSOURI. Howard Co.: Moore 1033 ce ). NORTH CAROLINA. Brunswick Co.: Moore 374 (GA). OKLAHOMA. McCurtain Co.: Moore 716 (GA). TENNESSEE. Carter Co.: Moore 251 (GA); Rutherford Co.: Moore 273 (GA). VIRGINIA. ve 835 Nelson Co.: (GA). — oe) _ Viris PALMATA Vahl, Symb. Bot. 3:42 — 43. 1794. — a Te in Virginiana, n.d., no collector (Lecrorype, here designated: — SYNTYP! Ch). V. rubra Michaux ex Planchon, in DC Monogr. Phan. 5:344. 1887. — Lecroryer, e designated: as microfiche IDC Michaux, no. 123, photo 2!(P). — SyNryPE: as er microfiche IDC Michaux, no. 123, photo 5! (P) Relatively slender, high climbing vine, the branchlets of the season subterete and usually entirely dark crimson or purplish-red until mature, upon maturity the branches then of a reddish-brown to chestnut color, glabrous to very thinly arachnoid. Bark exfoliating in shreds on mature stems, pith brown, interrupted by nodal diaphragms, diaphragms 2.5 — 4 mm thick. Tendrils bifurcate, red-pigmented when young, a tendril or inflorescence present at only 2 consecutive nodes, nodes not glaucous. Leaves with slender petioles that are somewhat shorter than the blades, glabrous to puberulent; blades generally cordiform, commonly deeply 3 (5) lobed, the lobes attenuate acuminate, sinuses acute to rounded; margins dentate-serrate; upper surface of mature leaves glabrous, lower 355 surface not glaucous, glabrous or pubescent with only hirtellous trichomes along the veins and in their axils; stipules 1.5 —3 mm long. Panicles 6—18 cm long, usually narrowly triangular in outline, infructescences usually with more than 25 berries; 3 or 4 seeded berries 8— 10 mm in diameter, bluish-black to black, with very little or no glaucescence, lenticels absent. Seeds dark brown, globose, 4— 7 mm long, nearly filling the berry. River banks and alluvial floodplain woodlands (IL and IN south to MO, TX, we AL, c panhandle of FL). Flowers the latest of all native species, mid to late June; fruit ripening late July to October. Representative aan: examined: ALABAMA. Hale Co.: Sens 49 (UNA). FLORIDA. Gadsden Co.: Moore 797 (GA); Moore 802 G A). INDIA x Co.: Deam 24145 (US). Posey Co.: "Tyen 4256 (US); Deam 39930 (GH). See on . A Oe Par- ish: Thieret 20841 (FSU); Smith 458 (TENN). MISSISSIPPI. LeFlore Co.: Moore 347 (GA). Neshoba Co.: Smith 883 (FSU Viris MONTICOLA Buckley, Proc. Acad. Nat. Sci. Philadelphia 62:450. 1861. V. aestivalis var. monticola (Buckley) Engelm., Amer. Naturalist 2:321. 1869. — Type: TEXAS. Hays Co.: Crescit in Texas, n.d., B. Backley 5.n. (LECTO- typr, here designated: US!). — Synrype: (PH!). High climbing vine, branchlets of the season angled when young but becoming terete at maturity, young stems and leaves slightly to moderately arachnoid pubescent. Bark exfoliating in shreds on mature stems, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms | — 2.5 mm thick. Tendrils bifurcate, a cendril or infloresc- ence present at only two consecutive nodes, nodes not glaucous, usually not banded with red pigmentation (but occasionally the red-banding present). Leaves with petioles about half as long as the blade, sparsely to moderately pubescent with arachnoid trichomes, glabrate at maturity; blades cordi- form, often 3-shouldered to shallowly 3-lobed; margins irregularly dentate-serrate, bases typically cordate, apices acute to short acuminate (occasionally long acuminate); upper surface of mature leaves typically glabrous, usually lustrous, lower surface not glaucous, typically green, glabrous to sparsely hirtellous pubescent; stipules 1.5 — 3 (-4) mm long. Panicles 3 —7 cm long, usually globose in general outline, infructescences typically with less than 25 berries (or pedicels); 3 or 4 seeded berries 8 — 10 mm in diameter, black, very slightly, or more typically, not at all glaucous, lenticels usually present. Seeds dark brown, ovoid, 5—7 mm long. In upland, well-drained habitats of various mixtures. Endemic to the Edwards Plateau in sc TX. Flowering in May, fruit ripening July to August. 356 Representative specimens examined: TEXAS. Bandera Co.: Moore 935 (GA). Bexar Co.: Clare 641 (BH). Blanco Co.: Whitehouse 546 (NY). Comal Co.: Palmer 12181 (GH-A). Kendall Co.: Palmer 13651 (GH). Kerr Co.: Moore 962 (GA); Cory 24043 (BH). Llano Co.: (NY). Real Co.: Cory 42700 (TEX); Cory 42701 (GH) Ranisey s.n, Subgenus VITIS, Series LABRUSCAE Planchon, in DC Monogr. Phan. 5:323. 1887. — Tver species: Vitis Lebrusca L. VITIS LABRUSCA L. oor Pl. 202. 1753. — Tyee: America Septentrionalt (LECTO- tyPr, here ee) LINN, as IDC microfiche no. 81.5!). — Synrype: LINN, as IDC microfiche no. 281.6!, \ labrusca var. labruscoides Eaton, Man. Bot. 496. 1818. — SyNrypEs: not seen . labrusca var. alba Pri Treatise on the Vine 181. 1830. Vo debrasca forma alba ?rince) Fernald, Rhodora 4 11:431. 1941. — Syntyprs: noc seen. ’ labrusca var. rosea Prince, Treatise on the Vine 182. 1830. — SyNryPEs: not seen. : labrusca vat. ae Fernald, Rhodora 42:462—463. 1940. — Type: VIRGINIA. ae s Crry Co.: swampy thickets southeast of Charles City, 22 Aug 1939, M. Fernald and B. Long 11074 (HOLOTYPE: GH!; isorypes: GH! NY! PH! US!). High climbing vine, branchlets of the season obscurely angled when young, becoming terete at maturity, young stems and leaves densely tomentose, occasionally with spinose pubescence with glandular tips. Bark exfoliating in shreds on mature stems, lenticels inconspicuous or absent, pith brown, interrupted by nodal diaphragms, diaphragms 0.5 — 2.5 mm wide. Tendrils bifurcate to occasionally crifurcate, continuous, a tendril or inflorescence present opposite virtually every node, occasionally absent Opposite lowermost nodes, nodes not glaucous, not banded with red pigmentation. Leaves ac petioles about as long as the blades, thinly arachnoid pubescent to glabrous; blades cordiform, often 3-shouldered: margins crenate to crenate-dentate, bases typically cordate, apices usually acute, upper surfaces of mature leaves glabrous to slightly pubescent, dull, lower surface not glaucous, typically whitish to yellowish due to dense arachnoid tomentum which obscures the leaf undersurface but not the veins; stipules 2—4 mm long. Panicles 6— 14 cm long, usually globose to cylindrical in general outline; infructescences usually with less than 25 berries, occasionally with less than 12. Berries greater than 12 mm in di- ameter, black, very slightly, or more typically, not at all glaucous, lenticels absent. Seeds brown, obcordate, 5-8 mm long. Inhabiting a very wide variety of sites, both upland and well drained and lowland and poorly drained, including intermittently flooded bottomlands (ME, NH and VT south ton GA, n AL, n MS, north to ne AR, se MO, € IL and s MI). Flowering in May to June, fruit ripening in September to October. a7 Representative specimens examined: GEORGIA. Rabun Co.: Jones 23062 (GA). . DIANA. Porter Co.: Deam 29810 (IND). MAINE. Oxford Co.: Moore 855 (GA PENNSYLVANIA. Fayette Co.: Moore S881 (GA). Pike Co.: He 846 os TENNESSEE. Cooke Co.: Hannewell 14254 (GH). VIRGINIA. Carroll Co.: Moore 231 (GA); Carroll Co.: Moore 245 (GA). Nelson Co.: Moore 836 (GA). ane Co.: Moore 378 (GA) VITIS SHUTTLEWORTHI! House, Amer. Midl. Naturalist 7:129. 1921. \ coriacea Shuttlew. ex Planchon, in DC Monogr. Phan. 5:345. 1887, nom. aes , non Miq. 1863. _ es coriacea (Shuttlew. ex Planchon) Bailey ex Gray, S . N. Amer. 1:429. 1897. — Typr: FLORIDA. borders of the Manatee ee a8 1845, ae 1/1 Goreme BM!). Moderately high climbing vigorous vine, branchlets of the season oval to terete, densely tomentose when young, becoming more thinly tomentose with age. Bark exfoliating in shreds on 2 year old stems, lenticels absent or inconspicuous, pith brown, interrupted by diaphragms at nodes, diaphragms typically 2.5 —6 mm thick but frequently continuing halfway into the internode. Tendrils bifurcate to trifurcate, a tendril or infloresc- ence present opposite only 2 consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half to three quarters the length of the blade, densely tomentose; blades broadly cordate to nearly reniform, typically unlobed but varying to 3-shouldered or, less often, deeply 3—5 lobed, when lobed the lobes acute and the sinuses rounded; margins with shallow, broad scalloped, obtuse teeth, typically nearly entire, leaf bases cordate to truncate; upper surface of mature leaves floccose to glabrous, lower surface not glaucous but densely and evenly covered with white to rusty tomentum, typically concealing the leaf under- surface but not always the veins; stipules minute, less than 1 mm long, promptly deciduous. Panicles 4— 10 cm long, the rachis arachnoid floccose, usually broadly short triangular in outline, infructescences with less than 25 berries, occasionally with less than 12. Berries large, greater than 12 mm in diameter, dark red to purple-black, with little or no glaucescence, lenticels absent. Seeds dark brown, ovoid to rounded, 5 — 6 mm long. Generally found in woodlands of various mixtures, woodland borders, thickets and lowland woods in peninsular FL (endemic to peninsular FL). Flowering in early April to early May, fruit ripening in June to August. Representative specimens examined: FLORIDA. Citrus Co.: Moore 776 (GA). Charlotte Co.: Moore 753 (GA). Collier Co.: Moore 760 (GA). DeSoto Co.: Moore 752 (GA). Glades Co.: Moore 749 (GA). Hardee Co.: Moore 322 a Hillsborough Co.: Pardue s.n. (USEF). Manatee Co.: Moore 786 (GA). Sarasota Co.: Moore 787 (GA); Moore 788 (GA). 358 Vitis MUSTANGENSIS Buckley, Proc. Acad. Nat. Sci. Philadelphia 62:451. 1861. — Tver: TEXAS. Near Austin, Apr 1860, S.B. Buckley s.n. (LECTOTYPE, here designated: PH!). — Synrype: (US!). High climbing vigorous vine, branchlets of the season oval to terete, densely tomentose when young, becoming more thinly tomentose with age. Bark exfoliating in shreds on 2 year old stems, lenticels absent or in- conspicuous, pith brown, interrupted by diaphragms at nodes, diaphragms 1.5 — 3 mm thick. Tendrils bifurcate to trifurcate, a tendril or inflorescence present opposite only 2 consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half to three quarters the length of the blade, densely tomentose; blades broadly cordate to nearly reniform, usually concavely folded, typically un- lobed but varying to 3-shouldered or deeply 3 —5 lobed, when lobed the lobes acute and the sinuses rounded; margins with shallow, broad scalloped, obtuse teeth, typically nearly entire, leaf bases cordate to truncate; upper surface of mature leaves floccose to glabrous, lower surface not glaucous but densely and evenly covered with white to rusty tomentum, typically concealing the leaf undersurface but not always the veins; stipules 1.5 —4 mm long, promptly deciduous. Panicles 4 — 10 cm long, the rachis arachnoid floccose, usually broadly short triangular in out- line, infructescences with less than 25 berries, occasionally with less than 12. Berries large, greater than 12 mm in diameter, black to less commonly dark red, with little or no glaucescence, lenticels absent. Seeds dark brown, ovoid to rounded, 6-7 mm long. Generally found in woodlands of various mixtures, woodland borders, thickets and lowland woods (e TX and extreme w LA north to s OK, with one disjunct population in Wilcox County, AL). Flowering in late May to early June, fruit ripening in August to September. In several early publications (e.g., Munson 1909; Bailey 1934), this species was known as V. candicans Engelm. ex Gray. Engelmann and Gray published this name in 1850, but the description of this taxon was quite vague, stating only that “Under the name of V. candicans (n.sp.) Engelm. ined., I have from Lindheimer, as also from Mr. Wright, Texan specimens of what appears to be a variety of V. californica Benth., with the leaves somewhat less dentate and more densely tomentose underneath”. Addi- tionally, it is not at all clear how Gray is treating the above description, as he did not list it as a new species, but rather included it under what appears to be the description of a new variety of V. aestivalis. However, Gray did not give this apparent new variety a name but only states “var. tomento albo, nec fulvo. Shady banks of streams, New Braunfels. Climbing high trees. Berries the size of peas, in large bunches, black; the taste vinous and 359 pleasant. Flowers very odorous.” Thus, the name V. candicans must be con- sidered ambiguous, making the name V. mwstangensis the valid and legit- imate one for this species. Representative specimens examined: ALABAMA. Wilcox Co.: Moore 728 (GA). OKLAHOMA. Marshall Co.: Goodman 5858 (GH). TEXAS. Anderson Co.: Moore 932 (GA). Comal Co.: Moore 687 (GA). Gonzales Co.: Webster & Wilbur 2977 (SMU). Grayson o.: Moore 713 (GA). Hays Co.: Moore 686 (GA). Leon Co.: Moore 935 (GA). Llano Co.: Moore 964 (GA). Mason Co.: Moore 693 (GA) ‘®) Subgenus VITIS, Series RIPARIAE Munson, U.S.D.A. Div. Pomol. ll. No. 3:7. 1890. — Tver species: V. riparia Michaux. Viris ACERIFOLIA Raf., Amer. Man. Grape Vines 14. 1830. — Dea. here designated: TEXAS. WitparGer Co.: growing along Beaver Creek o 283, Sof Vernon, ina rest area 1.5 mi S of jct. with Farm Road 1763, 13 Jun ne Moore 700 (GA!; IsONEOTYPES: PH!, US!). — Synrypss: not found. V. foneit Prince, Treatise on the Vine 184. 1830. — Synrypes: not s V. solonis Hort. Berol. ex Planchon, Vignes Amer. 119. oa ps cdi var. solonts yt. Berol. ex Planch) Planchon, Vignes Amer. 118. — SYNTYPES: not found. V. nuevo-mexitcana Lemmon ex Munson, Trans. Amer. Hort. Soc. 3:132. 1885. — Synrypes: not found. V. solonis var. microsperma Munson, Rev. Vitic. 3:158. 1895. V. longii var. microsperma (Munson) Bailey ex Gray, Syn. Fl. N. Amer. 1:423. 1897. — SyNTyPEs: not found. Typically a stocky, erect, shrubby, much branched low to moderately high climbing vine, branchlets of the season slightly angled when young but becoming terete, very young stems and leaves whitish arachnoid pub- escent, mature stems glabrous to arachnoid pubescent. Bark closely persistent for several years, then shredding in thin plates, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms usually less than 1.0 mm wide. Tendrils bifurcate, a tendril or inflorescence present at only two consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half as long as the blades, often partially conduplicately folded, typically moderately to thinly arachnoid pubescent to glabrate; blades broadly cordate, often 3- shouldered to shallowly 3-lobed; margins sharply dentate-serrate, bases typically broadly cordate, apices typically short acuminate; upper surface of mature leaves slightly arachnoid pubescent to glabrate, lower surface not glaucous, slightly arachnoid pubescent but also with sparse hirtellous pub- escence along the veins, varying to glabrate; stipules 3—6 mm long. Panicles 5—9 cm long, compact, globose in general outline, 1n- fructescences typically short pedunculate, making the clusters appear al- most sessile, typically with greater than 25 berries, but occasionally with 360 only 12 to 25; 3 or 4 seeded berries 8 — 12 mm in diameter, black, heavily glaucous, lenticels absent. Seeds reddish-brown, pyriform, 5—6 mm long. Inhabiting moist to slightly drier sites, river banks and alluvial flood- plain woodlands, but also along hedge rows and fence rows (nc TX, ne NM, se CO, sw KN, w OK). Flowering in April to May, fruit ripening in July to August. In earlier treatments of the genus, (e.g., Munson 1909; Bailey 1934), this species was known as V. /ongii Prince. Since both V. acerifolia and V. longit were published in 1830, Bailey (1934) stated that he could not choose between the two names and that both descriptions were equally good. Since Prince had the species in fruit, while Rafinesque apparently did not, and since the name “/ongir” had been used for many years, Bailey decided to continue using that name. However, Rehder (1946) later deter- mined that Rafinesque’s publication was dated May, 1830 in the preface while the copyright date of Prince's publication was September, 1830. Thus, the name “acer?folia” has priority over the name “/ongii.” Representative specimens examined: OKLAHOMA. Custer Co.: Moore 708 (GA). Dewey Co.: Ne/son 6095 (OKL). Grady Co.: Moore 978 (GA). Harmon Co.: Moore 701 (GA). Washita Co.: Moore 707 (GA). Woodward Co.: Moore 702 (GA). TEXAS. Collings- worth Co.: Moore 697 (GA). Donley Co.: Moore 698 (GA). Hemphill Co.: Cory 16224 (BH). Wilbarger Co.: Moore 700 (GA). Viris RIPARIA Michaux, Fl. Bor.-Amer. 2:231. 1803. V. cordifolia var. vulpina, (L.) Eaton, Man. Bot. 497. 1818. V. cordifolia var. riparia (Michaux) Gray, Manual ed. 5:113. 1867. V. ite ssp. riparia (Michaux) Clausen, Cornell Univ. Agric. Exp. Sta. Mem. 298:8 49, ype: ad ripas et in insulis fuviorum Ohio, Mis- SISSIPpl, tC. (HOLOTYPE: as sie de IDC Michaux no. 122, phoro 19, ee specimen! P) V. riparia var. praecox Engelm. ex Bailey, Amer. Garden 14:353. 1893. — SYNTYPES: not found — N . vilpina var. syrticola Fernald and Weigand, Rhodora 25:212. 1923. V. riparia var. syrticola (Fernald and Weigand) Fernald, Rhodora 41:431. 1931. — Type: NEW YORK. Osweco Co.: Selkirk, sand dunes overlying Silurian shales and schists by Lake Ontario, 23 Aug 1922, Fernald, Weigand and Eames 14388 (HoLoryPe: GH!; ISOTYPE: GH!). Moderate to high climbing vine, branchlets of the season terete, young stems and leaves glabrous to slightly hirtellous pubescent (varying to slightly arachnoid pubescent in some Louisiana specimens). Bark exfoliat- ing in shreds on mature stems, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms usually less than 0.5 mm wide. Tendrils bifurcate, a tendril or inflorescence present at only two consecutive nodes, nodes not glaucous, not banded with red pigmen- 361 tation. Leaves with petioles about half as long as the blades, slightly to moderately hirtellous pubescent, blades cordiform, 3-shouldered to shallowly 3-lobed, margins sharply dentate-serrate, bases typically cor- date, apices typically short acuminate; upper surface of mature leaves glabrous, often light yellowish-green, lower surface not glaucous, typi- cally green and with hirtellous trichomes along the veins and in their axils, varying to glabrate; stipules 3—5 mm long. Panicles 7 — 12 cm long, usually narrowly triangular in general outline, infructescences typically with more than 25 berries; 3 or 4 seeded berries 8 — 12 mm in diameter, black, heavily glaucous, lenticels absent. Seeds dark brown, pyriform, 5—6 mm long. Inhabiting a wide variety of habitats but preferring moist soils, stream banks, pond margins, alluvial woodlands but also on roadsides, hedge rows and fence rows (s New Brunswick west to se Saskatchewan, south to n VA, w TN, n MS, LA, e TX, north toe KN, e NB, e SD and e ND. Also reported from the Pacific Northwest). Flowering in April to June, fruit ripening in August to September. Representative specimens examined: ARKANSAS. Miller Co.: Moore 724 (G IOWA. Davis Co.: Moore 1041 (GA). Van Buren Co.: Moore 1042 (GA). MISSOURL. a. Co.: es 1035 (GA). NEW ee. gee : Moore 870 (GA). Schuyler Co.: Moore 575 (GA). Sullivan Co.: Moore 847 (GA). Warren Co.: ge 8609 (GA). VERMONT. Addison ae Moore 868 (GA). pan Cos Moore 865 Cc. ViITIS RUPESTRIS Scheele, pee 21:591. 1848. — Neoryee, here desig- nated: MISSOURI. Drenr Co.: all around gravel bed deposits covering large area around Meramec River ee T33N, R4W, sect. 14, 2 mi SE of Max, 10 Aug 1936, J. A. Steyermark 12842 (MO!). — Synrypes: not found. V. rupestris var. dissecta Eggert ex Bailey in Gray, Syn. Fl. N. Amer. 1:422. 1897. V. bile forma dissecta (Eggert ex Bailey in Gray) Fernald, Rhodora 41:431. — Type: MISSOURI. JEFFERSON Co.: brooks on hillsides, 22 May 1892, ae sm. (LECTOTYPE, here designated: NY!; sotecroryprs: F!, US!). Sprawling to low climbing, much branched vine, branchlets of the sea- son slightly angled but becoming terete at maturity, very young stems and leaves glabrous or slightly hirtellous pubescent. Bark persistent for the first several years, then shredding in plates, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms less than 1 mm wide. Tendrils bifurcate, commonly present only opposite the uppermost nodes and then only at two consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half as long as the blades, glabrous to slightly hirtellous pubescent; blades typically reni- form, conduplicately folded, particularly when young, often 3- shouldered, rarely shallowly 3-lobed; margins dentate-serrate, bases typi- 362 cally truncate to broadly cordate, apices acute to short acuminate; upper surface of mature leaves typically glabrous, often lustrous, lower surface not glaucous, typically green and glabrous, occasionally with sparse hirtel- lous pubescence along the veins and in their axils; stipules 3—6.5 mm long. Panicles 4-7 cm long, usually globose in general outline, in- Seaegog i typically with less than 25 berries, occasionally with less than 12; 3 or 4 seeded berries 8 — 12 mm in diameter, black, slightly glaucous, lenticels absent. Seeds light brown, ovoid, 5—6 mm lon Herbarium records indicate that this species was once distributed from south central Texas, through northern Arkansas, Missouri, northern Tennessee, Kentucky and northern West Virginia and northwestern Mary- land to southwestern Pennsylvania. It has apparently been extirpated from many of these regions and is now only found along calcareous, gravelly banks, river bottoms, stream beds and washes in south central Missouri and extreme northern Arkansas. Flowering in April to May, fruit ripening in August to September. It 1s a critical species as it is important in viticul- ture as a rootstock. Representative specimens examined: MARYLAND. paneer Co.: : Steele s.n. (NY). MISSOURI. Oregon Co.: Palmer & Steyermark 41716 (MO). Phelps Co.: Eggert 5.7. (BH). Iron Co.: Palmer 18103 (GH). PENNSYLVANIA. Lane Co.: tore SM, OKLAHOMA. Comanche Co.: Demaree 13141 (GH). TENNESSEE vidson Co. Gattinger 400A (GH). TEXAS. Johnson Co.: Reverchon 5.n. (SMU). ae Co. Ruth 368 (US). Tom Green Co.: Tweedy 134 (US). HYBRIDS Viris X champinu Planchon (pro. sp.), Vigne Amer. 6:22. 1882. — Neoryer, here designated: TEXAS. Grayson Co.: originally from Llano County, cultivated Denison, Texas, Munson vineyard, 25 Apr 1890, EM. Ramsey s.n. (NY!; ISONEOTYPE: MO!) — SyNnryprs: not found. High climbing vine, branchlets of the season somewhat angled when young, becoming terete when mature, young stems and leaves arachnoid pubescent, becoming glabrate with age. Bark tardily exfoliating in shreds, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms 1.5 — 2.5 mm thick. Tendrils bifurcate, rarely trifurcate, a tendril or inflorescence present at only two consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half as long as the blade, thinly arachnoid pubescent to less commonly glabrous; blades cordate to occasionally nearly reniform, often 3-shouldered to very shallowly 3-lobed, occasionally partially condupli- cate; margins crenate to slightly serrate, bases typically cordate, apices acute to short acuminate; upper surface of mature leaves typically glabrous, often lustrous, lower surface not glaucous, typically green, thinly arach- 363 noid pubescent to glabrate; stipules 2.0 — 5.5 mm long. Panicles 3 —7 cm a usually globose in general outline, infructescences usually with less an 25 berries (or pedicels); 3 or 4 seeded berries greater than 12 mm in ie black, very slightly, or more typically, not at all glaucous, lenticels absent. Seeds brown, ovoid, 5— 6 mm long. Inhabiting well drained a lcaicows soils in sc Texas, on and adjacent to the Edwards Plateau. Flowering in April to May, fruit ripening in July to August. Interpreted here as a hybrid between V. mustangensis and V. rupes- tris, Comeaux (pers. comm.), however, presently feels that the origin of this taxon may be more complicated. It is now rare in nature (Comeaux, 1987b). eee specimens examined: NORTH CAROLINA. oe - — Co. (BH). TEXAS. Burnet Co.: Biltmore Herbarium 14842 (US). Bell Co.: ik s.n. (BH). Travis Co.: Manson s.n. (MO). § arse Tos te SM, Gas Originally from Coryell County, Moore ee GA), vated, on, Monin Sm, (BH); cultivated, Denison, Manso , (MOR). ae from Ue County, Manson s.n., (MO). No collector, no ees no location (29 May 1896) (B Vitis X doaniana Munson ex Viala (pro. sp.), Mission Vitic. Amér. 101. 1889. — Typr: TEXAS. WitparGer Co.: Wilbarger County, 1886, cultivated in the Munson vineyard, Denison, 24 Apr 1890, T.V. Munson s.n. (LECTOTYPE, here designated: NY!). — Synrypes: (CU! BH! FLAS! US!). High climbing vine, but shrubby and sprawling without support, branchlets of the season terete when mature, young stems and leaves densely tomentose. Bark tardily exfoliating in shreds, lenticels absent or inconspicuous, pith brown, interrupted by nodal diaphragms, diaphragms 1—2 mm thick. Tendrils bifurcate to occasionally trifurcate, a tendril or inflorescence present at only two consecutive nodes, nodes not glaucous, not banded with red pigmentation. Leaves with petioles about half as long as the blades, thinly arachnoid pubescent; blades cordate, often 3- shouldered to shallowly 3-lobed; margins crenate to crenate-serrate, bases typically cordate, apices acute to short acuminate; upper surface of mature leaves slightly to moderately arachnoid pubescent, lower surface not glaucous, typically dull green, slightly to moderately arachnoid pubes- cent; stipules 3 —6 mm long. Panicles 4— 10 cm long, usually globose to short triangular in outline, infructescences with less than 25 berries (or pedicels); 3 or 4 seeded berries greater than 12 mm in diameter, black, heavily glaucous, lenticels absent. Seeds dark brown, ovoid, 6-7 mm g. Inhabiting well-drained, drier soils in sand hills, plains and timber regions (n TX to s OK). Flowering in April to May, fruit ripening in July to August. A hybrid between V. mustangensis and V. acerifolia, once more 364 common in nature than it is at present, that was named for Judge J. Doan of Wilbarger County, Texas, who manufactured wine from the berries of this species. The town of Doans in Wilbarger County is named after Judge Doan where populations of this hybrid can still be found. Representative Lane examined: MASSACHUSETTS. cultivated, Arnold Arboretum, Pa/mer s.n., (MOR). NEW YORK. Besa . o.: cultivated, Bean 668 (BH). OKLAHOMA. Filles Co. jes 975 (GA). TEXAS ilbarger Co.: Moore 974 (GA), Manson son. (NY). Cultiv ee Denison, 6 May on me s.”. (BH); cultivated, Dent- son, Aug site Munson s.”, (BH); cultivated Denison, Munson s.n. (BH): cultivated Dent- son, originally m Wilbarger Co.: 24 Apr 1890 Munson s.n. (BH); cultivated, Denison, 7 Aug 1895 a son, (BH). — —~ cL Viris X novae-angliae Fernald (pro. sp.), Rhodora 19:146. 1917, — Tyee: MAINE. Penosscor Co.: thicket by river, Orono, 27 Jun 1906, M.L. Fernald yu. (LECTOTYPE, here designated: GH!; isotecroryprs: GH! NY! PH!). Syntype: (GH!). — Paratyrrs: (GH!, PH!) High climbing vine, branchlets of the season terete at maturity, young stems and leaves densely tomentose. Bark exfoliating in shreds on mature stems, lenticels inconspicuous or absent, pith brown, interrupted by nodal diaphragms, diaphragms 0.3— 1.1 mm thick. Tendrils bifurcate, con- tinuous, a tendril or inflorescence present at three to several consecutive nodes, but frequently not present opposite all nodes as in V. dabrusca, nodes not glaucous, not banded with red pigmentation. Leaves with petioles one half to nearly as long as the blades, sparsely arachnoid pubescent to glabrate; blades cordiform, often 3-shouldered; margins crenate to irregu- arly dentate-serrate, bases typically cordate, apices acute to short acumi- nate; upper surface of mature leaves typically glabrous, lower surface not glaucous, typically green when mature, more or less densely arachnoid pubescent on young but expanded leaves, only slightly arachnoid pubes- cent on fully mature leaves; stipules 2.5 — 6.0 mm long. Panicles 7 — 13 cm long, usually triangular in general outline, infructescences typically with more than 25 berries, but occasionally with only 12 to 25; 3 or 4 seeded berries greater than 12 mm in diameter, black, slightly glaucous, lenticels absent. Seeds brown, 6—8 mm long. Thickets, largely alluvial, as well as roadsides, pond and stream margins, and fence and hedge rows (s ME south to n PA and n NJ). Flower- ing in June, fruit ripening in August to September. A hybrid between V. labrusca and V. riparia that is common in the New England region. _ pe ean specimens examined: MAINE. Franklin Co.: ea 852 ee Knowlton yn. (USF). Waldo Co.: Seymour 30055 (WT). MASSACHUSETTS. Middlesex Co.: Smith sn. (MO). He Co.: Williams s.n. (GH). NEW eae ine Co. Moore 559 (GA). Cheshire Co.: Batchelder s.n. (PH). Merrimack Co.: Moore 856 (GA); Moore 858 (GA). PENNSYLVANIA. Lackawanna Co.: Glowenke 8343 (PH 365 Nomina nuda et dubiosa Names without diagnosis or of such uncertainty as not to be cited confid- ently in regular synonomy. V. amara Raf., Amer. Man. na Vines 1830. Vi americana Baca ed. Rep. hexade 2, vol. a 1804 V. angulata a Amer. Man. Lee Vines V. bicolor LeConte, Proc. Acad. Scare C2272. 1893. V. bifida Raf., nee Man. Grape ee 12. 1830. V. blanda Raf., Amer. Man. Grape Vines 12. 1830. V. boulderensis Daniels, Univ. Missouri Stud., Sct. Ser. 2(2):159. 1911. V. bracteata Raf., Amer. Man. a Vines 9. 1830. V. bracteata LeConte, Proc. Acad. me ciae jaa ad Gi27 1s; LO: V. caerulea Munson ex Viala, ce Vitic Amer. 113. 1889. V. callosa Raf., Amer. Man. Grape Vines 9. 1830. V. campestris Bartram, Travels Carolina 400. 1791. V. candicans Englem. ex Gray, Boston J. Nat. Hist. 6:166. 1850. V. canina Raf., Amer. Man. Grape Vines 11. 1830. V. ciliata Raf., Amer. Man. Grape Vines 13. 1830. V. ee Raf., Amer. Man. Grape Vines 15. 1830. V. concolor Raf., ere Man. Grape Vines 14. 1830 V. digitata Raf., Amer. Man. Grape Vines 9. | V. dimidiata Raf. Amer. Man. Grape Vines 13. 1830, V. diversifolia Prince, Treatise on the Vine 183. 1830. V. ferruginea Raf., . Man. Grape Vines 12. 1830. V. floridana Raf., ees Man. Grape Vines 17. 1830. V. falva Raf., Amer. Man. Grape Vines 8. 1830. V. glariosa Raf., ae mer. Man. Grape Vines 10. 1830. V. hbyemalis Raf., Amer. Man. Grape Vines 9. 1830. V. linoensis Prince, Treatise on the Vine 185. 1830 V. incisa Jacq., Hort. Schoenbr. 4:14. 1804 V. inteerifolia Raf., Amer. Man. Grape Winies 18. 1830. V. latifolia Raf., Amer. Man. Grape Vines 10. ne 0. V. lecontiana House, Amer. Midl. Naturalist 7: V. longifolia Raf., Amer. Man. Grape Vines 13. 1830 V. luteola Raf., Amer. Man. Grape Vines II. 1830. Vo mrissourtensts cae Treatise on the Vine 184. 1830. V. norton: Prince, Treatise on the Vine 186. 1830. V. peltata Raf., Amer. Man. Grape Vines 17. 1830. V. poiretia Raf., Amer. Man. Grape Vines 18. 1830. V. populifolia Raf., Amer. Man. Grape Vines 15. 1830. V. prolifera Raf., Amer. Man. Grape Vines 18. 1830. V. rugosa Rat., Anien Man. Grape Vines 11. 1830. V. saxatilis Rat., Amer. Man. ee Nee 8. 1830. V. serotina Bartram, Med. Re exade 2, vol. 1:22. 1804. at es © Nh V. sylvestris Batein, Med. Rep: on de 2, . Te C, caryopsis, x 10 1/2. BP 1 ari x 2/3. B, spikelec, 1. Sclerochloa dura. A, plant, FIG. at 2 Lolinm perenne, Matricaria discoidea, Poa angustifolia, P. annua, Polygonum arenastrum, Stellaria media, and Veronica hederifolia. The specimens of S. dura we have seen were collected from late winter (“February,” Texas) to early summer (20 July, Washington), but mostly from mid April to early June. Green during the first part of the growing season, the plants eventually become light yellow-brown. At this time they are easiest to locate, as areas infested by the species change color. Yellow-brown ballfields are a common sight where the species is dominant. The plants often persist, unshattered and dead, until mid July (at least in Oklahoma and Utah, where most of our observations of growing plants were made). Seed dispersal is often accomplished, we believe, through dis- turbance of these dried plants. The dispersal units may consist of one or more florets, one or more spikelets, irregular pieces of inflorescence, more or less intact inflorescences, or even entire culms or plants. Like many grasses, S. dura has had a tortured nomenclatural history. It was originally described by Linnaeus (Linnaeus 1753) as Cynosarus durns. Thereafter, Scopoli (1772) included it in Poa; Villars (1787), in Festuca: and Lamarck (1791), in Elewsine. Beauvois (1812) established the genus Sclerochloa to accommodate the species. There, except for a transfer to Sesleria by Kunth (1829), it has rested ever since. The chromosome number of §. dura was reported by Stace (1980) as 2n = 14 and by Tsvelev ieee ox = A foie species of Sclerochloa, 8. woronowii (Hack.) Tsvelev, originally described as a species of ie in 1912, ts known from Syria, Iraq, the Caucasus, and Afghanistan (Bor 1968). The description of §. dura below is based mainly on our study of many herbarium specimens of this species; figures in brackets are literature data we could not confirm. We offer it as a supplement to descriptions of the species we have seen (Bor 1968, 1970; Clayton & Renvoise 1986; Cope 1982; Cronquise et al. 1977; Gould 1975; Gould & Shaw 1983: Hegi 1906; Hitchcock 1935, 1950; Ladd 1983; Maire 1955: Roshevits 1980; Rugolo de Agrasar 1980; Stace 1980; Swallen 1931; Tsvelev 1983; Watson & Dallwitz 1980, 1988). Rosengurtt et al. include data on lipids in the central endosperm of S. dura ; Watson & Dallwitz (1988), data on anatomy (see also Dallwitz {1980} and Watson et al. {1986}). om SCLEROCHLOA DURA (L.) Beauv., Ess. Agrost. 98. 1812. Cynosurus durus L., Sp. Pl. ed. 1:72. 1753. Poa dura (L.) Scop., Fl. Carn. 1:70. 1772. Festica dura (1..) Vill., Hist. Pl. Dauph. 2: 94. 1787. Eleusine dura (L.) Lam., Tab. Encycl. eee ee 1:203. 1791. | 29 Crassipes annuns Swallen, Amer. J. Bot. 18: oy L931. ™ a lone {\\ i ss tq ~ Rene’ Rp PEAY ion wa, yh can ae ee ena TAL _ Sclerochloa dura. Documented distribution by county in the United States. ) FIG. 374 Annual. Plants often matted, occasionally solitary, green but becoming stramineous in age. CULMS generally prostrate or procumbent to ascend- ing but sometimes erect, branched, 2 — 18(30) cm tall though mostly less than 15 cm, many from tillering at basal nodes, the nodes glabrous, the internodes glabrous, solid or hollow with narrow lumen, more or less flattened. LEAVES basal and cauline, strongly overlapping toward base, usually overtopping inflorescences, the junction of sheath and blade not well defined; sheaths closed and tubular in lower 1/4 to 1/2, open above, more or less rounded on lower leaves, rounded to keeled on upper leaves, longer than internodes, glabrous, the margins conspicuously and broadly hyaline, the upper sheaths often inflated; auricles absent; /igules membra- nous, broadly triangular, (0.3) 0.75 — 2 (3.3) [3.5} mm long, glabrous, the margin entire to more or less lacerate, the apex acute; collars pale white to yellowish white, glabrous; b/ades flat or folded, (0.15) 0.5—5 (7) cm long, 1-4 mm wide, glabrous above and below or scaberulous on midrib, the apex boat-shaped, the margins entire or scaberulous. INFLORES- CENCE oblong to broadly elliptic, 1—4 [5} cm long, 0.5—2 [4] cm wide, often partially enclosed in the upper leaf sheath(s), the spikelets overlapping on short, thick pedicels (or nearly sessile) arranged along one side of a more or less zig-zag rachis; middle (and sometimes lower) nodes bearing short branches with 2 — 5 spikelets, spikelets solitary at upper (and usually lower) nodes, rarely with all nodes bearing only single spikelets; no general mode of disarticulation (see text above). SPIKELETS narrowly ob- long, laterally compressed, (3.4)5 — 12 mm long; florets (2)3 — 4(7), upper one or two sterile, the first floret more or less sessile, remaining florets on rachilla joints 1 — 3.5 mm long and ca. 0.5 mm wide; g/wmes weakly dor- sally compressed, both shorter than first lemma, awnless, chartaceous, glabrous, the apices blunt or emarginate, the margins broadly hyaline; first glume lanceolate to narrowly oblong, 1.4 —3(3.7) mm long, nerves (1) 3 (5); second glume oblong to elliptic, longer than first glume, 2.6—5. (6.2) mm long, nerves (3 or 5) 7 (9); lemmas awnless, oblong to narrowly lanceolate, laterally compressed, (3.4) 4.5 — 5.8 (7) mm long in first floret, (0.4) 1—4.5 (5.9) mm long in remaining florets, chartaceous-indurate, glabrous or scaberulous on midnerve toward apex, incompletely and irregularly (5) 7—9 nerved, the nerves parallel, the apex obtuse, the margins broadly hyaline; pa/eas dorsally compressed, ca. 0.5 —1.5 mm shorter than to equalling the lemma, 2-nerved, glabrous or, in upper half of keels, scaberulous, the keels slightly winged in upper 1/2; apex blunt to variously lobed or notched; margins broadly hyaline. STAMENS 3, anthers 0.8 — 1.3 {1.5} mm long. LODICULES 2, broadly oblong to oval, 0.75 —2 mm long, clawed at base, the apex entire to somewhat oye) lacerate, the margins entire. CARYOPSES yellowish brown, narrowly lanceolate in outline, 2. 1— 3.5 mm long, 0.8— 1.4 mm wide, rugulose, weakly trigonous, beaked by remnants of persistent styles/stigmas. ACKNOWLEDGEMENTS We acknowledge aid from John R. Baird, Jeremy Bruhl, Douglas Ladd, Clive Stace, and Kathleen Stewart. The illustration of S. dura is the work of Paul W. Nelson. Herbaria from which we borrowed specimens of S. dura are ARIZ, CAS, CHSC, CINC, COLO, DAO, DS, E FLAS, GH, HSC, , ILL, ISC, KANU, KNK, KSC, LL, MICH, MIN, MO, MONT, MU, USE UTC, VDB, WARM, WILLU, WIN, WIS, WS, WTU, and WVA. A list of specimens we examined is available upon request to JWT. The following herbaria had no specimens of S. dura: ASC, BAYLU, BHSC, CSCN, FHKSC, ILLS, KNFY, KSTC, NMC, NSMC, PUSC, ROPA, SAT, SDU, SOC, SRSC, TTC, UNLV, UNM, WCW, and WWB REFERENCES ALLEN, C. M. 1980. Grasses of Louisiana. University of Southwestern Louisiana, Lafay- Ctl. BEAUVOIS, A.-M.-E-J. P de. 1812. Essai d'une nouvelle agrostographie,; ou nouveaux genres des graminées. Imprimerie de Fain, Paris. BLACK, J. M. 1978. Flora of South el Sid.-dis; le 1. Lycopodiaceae- Orchidaceae. D. J. Woolman, Government Printer, Adelai BOR, N. L. 1968. Gramineae. Volume 9 mm C. C. Townsend ik Guest, eds., Flora of Iraq. Ministry of Agriculture, Republic of ng {Baghdad}. «1970, Gramineae. Lfg. 70/30 in K. H. Rechinger, ed., Flora Iranica. Akade- mische Druck- u. Verlagsanstalt, Graz, janes CLAYTON, W. D. and S. A. RENVOISE. 1986. Genera graminum. Her Majesty's Stationery Office, London. COPE, T. A. 1982. Poaceae. No. 143 in E. Nasir and S. I. Ali, eds., Flora of Pakistan. No place indicated, no publisher cas ave A., A. H. HOLMGREN, N. H. HOLMGREN, J. L. REVEAL and P. K. OLMGREN. 1977. a, flora. Vol. 6. The A canentyledons. Columbia Ga Press, New York ere , M. J. 1980. A general system for coding taxonomic descriptions. Taxon 29:4 DE Ser - R. J. 1980. Gilkey’s weeds of the Pacific Northwest. Oregon State University Press, Corvallis. FRECKMANN, R. W. 1988. Sclerochloa dura (L.) Beauv. (Poaceae) [in Tennessee}. Castanea 53:311. 1988. GAINES, X. M. and D. G. SWAN. 1972. Weeds of eastern ee and adjacent areas. Camp-Na-Bor-Lee Associates, Davenport, Washingt GOODMAN, G. J. 1974. Notes on Oklahoma plants. es Oklahoma Acad. Sci. 54:98 — 99. 376 GOULD, FE W. 1975. The grasses of Texas. Texas A&M University Press, College Station and R. B. SHAW. 1983. Grass systematics. 2nd ed. Texas A&M University Press, College Station. ILL, S 1988. New plant records for Maryland with an additional note on Nymphaea tetragona (Nymphaeaceae) pollination. Castanea 53: HITCHCOCK, A. S. Publ. 200. oO oo 1935. Manual of the grasses of the United States. U.S.D.A. Misc. ——___.. 1950. Manual of the grasses of the United States. Revised by Agnes Chase. U.S.D.A. Misc. Publ. 200. KUNTH, K. S. 1829. Revision des graminées publiées dans les Nova genera et species plantarum de Humboldt et Bonpland. Paris. LADD, D. 1983. Sclerochloa dura (..) Beauv. in Missouri. Missouriensis 4(3):73 — 75. LAMARCK {“de la Mark”}. 1791. Tableau sk pe et méthodique des trois regnes de la nature. Botanique, vol. 1. Chez Panckoucke, Paris. LINNAEUS 753. Species fe intarum. Impensis . Salvit, Holmiae. ao S, D. T. 1977. Additions to the Louisiana flora. Sida 7:220 — MAIRE, R. 1955. Flore de Afrique du Nord. Vol. 3 Paris. a E. and D. KORNECK. 1982. Sclerochloa dura (L.) P.B. im unteren Nahegebiet. dess. Florist. Briefe 31(4):63 — 64. ee NGURTT, B., A. LAGUARDIA, and B. R. ARRILLAGA DE MAFFEI. 1971. El endosperma “central lipido en la 11(2):383 — ROSHEVITS, ‘ Y. 1980. Grasses. An introduction to the study of fodder and cereal grasses. Translated from Russian. Indian National Scientific Documentation Centre, New Delhi. RUGOLO DE AGRASAR, Z. E. 1980. Sclerochloa dura (L.) P de Beauv. (Gramineae), adventicia en la Argentina. Darwiniana 22:557 — SCOPOLI, J. A. 1772. Flora Carniolica. Impensis Joannis Paulikrauss, Vienna STACE, C. A. 1980. Peis Beauv. Fl. Europaea 5:27. SWALLEN, J. 18:684 — 685. TSVELEV, N. N. 1983. Grasses of the Soviet Union. reer poe es C ompany, New Delhi — . Monocotyledonae. Paul Lechevalier sistematica de gramineas. Adansonia II. 560. Crassipes, a new grass genus from Utah. Amer. J. Bot. Part 2. Translated from Russian. VILLARS Libratres, oe WATSON, L. ane : 7 Dallwitz. 1980. Australian — pene The Australian National University, Research School of Biological Sciences, Canberra and «1:98 oire des plantes de Dauphiné. Vol. 2. Chez l'Auceur & Chez les 8. Grass genera i the world. INT KEY set for tnteract- ive identification and information retrieval. The Australian National University, Research School of Biological Se cie Canberra. R. JOHNSTON. Grass genera of the world: 728 detailed dese riptions from an nuitomaced data base. Australian J. Bot. 34:223 — 230. SEEDLING MORPHOLOGY IN CLEMATIS (RANUNCULACEAE) AND ITS TAXONOMIC IMPLICATIONS FREDERICK B. ESSIG Department of Biology University of South Florida, Tampa, FL 33620, U.S.A. ABSTRACT Seeds of 58 species of Clematis and Clematopsis were obtained from a variety of sources, germinated, and their seedling and juvenile morphology observed. Two very distinctive patterns emerged, each consisting of a cluster of characters. In Type I eae the eophylls are alternate and toothed. Hypocotyls are elongate (except in C saa elevat- ing the cotyledons and apical bud above ground, and buds are lacking in the axils of the cotyledons. Such seedlings are similar to those found in related genera such as ae and are found in Clematis in the infrageneric taxa Clematis, Lasiantha, pies Tubulosae, Atragene, Meclatis, Chetropsis, Bebaeanthera, Naraveliopsis, Papuastae, and in the genus Clematopsis. In Type I seedlings, leaves are paired from the beginning, but the firse 1 — pairs are usually reduced to phe Leaves are generally entire, often becoming lobec lor divided, but not toothed. Hypocotyls are short, keeping the cotyledon bases and the epicotyl at first subterranean. Buds are typically present in the axils of the cotyledons. Such seedlings are found in the infrageneric taxa Crispae, Viticella, Patentes, Rectae, and Angustifo- liae. These differences in seedling morphology and some correlated characters suggest a fundamental split in the genus and a basis for a revised infrageneric classification. RESUMEN Semillas de 58 especie de Clematis y Clematopsis fueren obtenido de orfgenes diversos, se germino, y sus eae juvenil se observid. Dos modelos distintos se manifestaron, cada uno consite en un grupo de varios caracteres. En las plantas de semilleros del Tipo Uno los eofilos son alternos y de nes Los hipocotilos son alargados (con exclusion de Clematopsis), y elevan los cotiledones y la yema cimera sobre la tierra. Yemas son ausente de las axilas do los cotiledones. Plantas de seimlleros de este tipo son semejante a aquellas hallado en géneros relatados como Anemone. Se encuentran en las taxa infragenéricas Clematis, Lastantha, Connatae, Tubulosae, Atragene, Meclatis, C ale ini Naraveltopsts, Papuasicae, de Clematis y en el Be nero Clematopsis. En pla | Tipo II, todas sey son re per las parea 1 — 3 son usualmente reduciendo a carafilos. Las hojas son sualmente enteras, frecuentemente lobados pero no dentados. Hipocotilos son corto, y las ee de i eoulledones y la es cimera se _ subterraneo. Yemas se encuentran en las . Pla Je semilleros de este tipo se hallan en las taxa infrageréricas axilas de | as Crispae, Viticella, Patentes, Rectae, y een Estas differencias morfologicas de las plantas de semilleros y algunes caracteres correlativos sugeron una division fundamental in el género y un fundamento para revisar la snakes pases Sipa 14(3):377 — 390. 1991. INTRODUCTION The genus Clematis is a large and diverse genus of the presumedly archaic family Ranunculaceae. Found on every continent except Antarctica, the approximately 300 species of Clematis occur in nearly every climatic zone from the taiga to the equatorial tropics, and display a wide variety of both vegetative and floral forms. There has been no comprehensive revision of the genus since that of Kuntze (1885), but there have been recent efforts to develop a modern infrageneric classification. Authors have subdivided the genus in various ways (see Keener & Dennis, 1982, for a review), some dividing it into subgenera, others dividing it into sections. Tamura (1967) divided Clematis into 12 sections (Table 1) in the most comprehensive of recent Classifications. His great familiarity with Asiatic Ranunculaceae allowed him to define fairly precisely various infrageneric taxa occurring in that region, but species from other regions, particularly Africa and New Zealand, are sometimes difficult to place in his system. Tamura did not attempt to group his sections into subgenera, perhaps feeling that there was insufficient basis for recognizing major divisions within the genus. Keener and Dennis (1982), on the other hand, divided the native and naturalized North American species into four subgenera, drawing upon earlier subgeneric concepts. They did not attempt to incorporate the old world taxa into their system, or to further divide their subgenera into sections, etc., stating that a new world-wide monographic treatment would be needed in order to accomplish this. Thus, the reconciliation of Tamura’s sectional classification with the subgeneric system of Keener and Dennis remains to be done. Tamura’s system emphasizes floral and inflorescence characters, as can be seen in Table 1. The best and most natural classification systems generally result, however, when a full range of characters from flower, fruit, seed, seedling, and vegetative shoots are employed. Study of additional characters, particularly vegetative characters, and perhaps also cytological and chemical characters, is therefore needed in this genus. A comprehens- ive new classification should reflect major lines of evolution at the subgene- ric level, and link the sections, subsections, etc. in a hierarchical fashion. In building up a collection of tropical and subtropical species of Clematis at the University of South Florida Botanical Garden, striking differences in seedling morphology and related vegetative features were noticed, suggesting that a survey of the genus would be worthwhile. Little has been recorded concerning seedling morphology in Clematis, despite the fact that numerous species have been cultivated over the past two centuries. Lubbock (1892) described and illustrated the seedlings of three species (C. recta L., C. orientalis L. (as C. graveolens Lindl.), and C. (Atragene) alpina 319 Taste |. Classification of the Clematis alliance according to Tamura 1956, 1967), ith di cingt ishit g& haere as reported by Tamura; asterisk indicates taxa known to have Type II seedlings. Genus CLematis [leaves opposite, sepals valvate] Section Viorna [sepals erect, often colored, stamen filaments hairy} Subsection Connatae [woody tees leaves toothed] Subsection Tvbulosae erect semi-shrubs, leaves toothed] *Subsection Cripae [perennial herbs or weak-stemmed vines, leaves entire] Section Bebaeanthera [as in Viorna, but flowers fasciculate with new growth] Section Afragene [flowers with stamen-derived “petals,” leaves toothed] Section Meclatis {sepals spreading to erect, mostly yellow to orange, stamen filaments hairy] Subsection Orrentales [flowers few to many in axillary or terminal clusters Subsection Tanguticae [single Howers terminating new shoots} Section ees {sepals spreading, whitish, stamen filaments ais leaves mostly toothed} Subsection a ee {flowers 3 — 3.5 cm diam., stamen filaments dilated downward] Subsection Vitalhae [flowers « 3cm diam., stamen filaments filiform Subsection Dyorcae [as above, but flowers mostly dioecious] Subsection Aritatae {as above, but stamen connective projected] Subsection Papuasicae {similar to above, difference in structure of panicle} Subsection Crass/foliae [stamen filaments rugulose, leaves Cortacious, entire] *Subsection Rectae [anthers clongate; leaves entire} *Subsection Angustifoliae [as above but Howers large, with 6 sepals] Section Cherropsis {flowers fasciculate with new growth, large, bisexual, sepals spreading] Section Lasiantha fas above but flowers dioecious} Section Viticella [flowers large, sepals spreading, colored, stamens glabrous, leaves entire} Subsection F/ordae [flowers solitary, axillary, subtended by two bracteoles, styles plumose] *Subsection Viticella {as above, but styles short, not plumose] fall} Section Prerocarpa {achenes strongly compressed and winged, leaves entire] Section Fraticella [woody shrubs with very small leaves] *Section Patentes [as above but flowers from bud produced in f Section Naravelropsis [anther connectives much prolonged, similar to Papuasicae} Genus Archiclematis [leaves alternate throughout, otherwise similar to Connatae] Genus Clematopsis [flowers large, sepals imbricate, otherwise similar to Connatae] Genus Naravelia [Howers with elongate, petal-like staminodes] (L.) Miller). The seedling morphology of C. recta described by Lubbock agrees with the “Type II” morphology described in this paper, while that for C. graveolens and C. alpina agree with the “Type I” morphology descri- bed here. Erickson (1945) illustrated C. fremontii S. Watson var riehli Erickson, and it agrees with “Type II.” This study was undertaken then in anticipation that unrecognized and overlooked vegetative features, such as those of the seedling, might provide clues to the major lines of evolution in the genus that are ambiguous when only floral features are used. MATERIALS AND METHODS Efforts were made to obtain seed of species representing all recognized infrageneric taxa in Clematis and of several closely related genera, following 380 the classification of Tamura (1967, see Table 1). According to Tamura, three genera, Archiclematis, Clematopsis, and Naravelia, are distinct from Clematis, but closely related. Each has been included in it by various previous authors. Therefore, they have been considered in this study, although I thus far have only obtained seed of Clematopsis. Altogether, Tamura’s smallest units (subsections and undivided sections), plus the three related genera, make 26 initial units for systematic study. Seedlings of 58 species, representing 20 of these 26 units (Table 2) were observed. Seeds were obtained from a variety of sources, including botani- cal gardens, commercial seed companies, and private collectors. Seeds of native Florida species and some others were collected by the author. Seed from cultivated sources frequently prove to be misidentified or of dubious or mixed ancestry. Therefore, great care has been taken to assure that the material reported upon has been accurately identified. Identity of all specimens is being verified as the plants become mature, and specimens whose identity or infrageneric placement is still uncertain are not inclu- dec Seeds were germinated in a greenhouse at the USF Botanical Garden some only after stratification and/or a long period of dormancy. Many plants were later transferred to an outdoor experimental plot. All access- 1ons were photographed after the first leaf appeared and often at later st- ages. Seedlings of many species were preserved and examined under a dis- secting microscope. Voucher specimens and photographs are being made as each specimen blooms for the first time. Taste 2, Species examined (all cultivated at USF Botanical Garden). Name Laxon Type USF Acc. # C. addisonit Britton Crispac I 87-40 C. alpir ill Atragene I 87-41 C. apiutolia DC Vitalbac I 82-22 C. ariscata R. Br. aes I 87-57 C. baldwinit Torrey & A. Gray crispae Il 86-25 C. barbellaca Edgew. ae anthera I 87-139 Cc. ae Ker. -Gawl Vitalbae I 87-2 C. buchaniana DC Connatae I ae C. cam sailors. BiG Viticella I C. catesbyana Pursh Dioicae I 85- , 86-35 C. chinensis Osbeck Rectae Il 88- C. chrysocoma Franchet Cheiropsis | 87-48 Taser 2 (Concinued) AAD id ana Te C) Dees Sie Ge prs a) Cams Gat i ire Tan Gea 1 ANAAAD one NRA R C) ¢) ¢ | tee. UT te, 9 L Loa NRRL GE Fae a an Ge Pan a) EE CL an C C. CHOs CY COICY OG >. cirrhosa L. crispa L. denticulata Vell. drummondii Torrey & A. Gray filamentosa Dunn flammula L. fusca Turcz. ». gentianoides DC. >. glaucophylla Small grata Wallich . heracletfolia DC. . hexapetala Pall. hirsutissima Pursh integrifolia L intricata Bunge kirilowit Maxim. lasiantha Nutt leschenaultiana DC. liguscicifolia Nutc. macropetala Ledeb. mandshurica Rupr. microphylla DC. napaulensis DC. ». papuasica Merr. & Perry . patens Morr. & Decne peterae Hand.-Mazz. picroti Miquel >. pitcher: Sargent >. ranunculoides Franchet recta L. _ vitalba L. >. vicicella L. EMATOPSIS ». villosa DC >. scabrosifolia Viguier & Perrier . leas var. kirkii ancthitolia Hook. Lasiantha Connatae Dioicae Atragene Rectae ? Bebaeanthera Meclatis Meclatis Papuasicace Picrotianae Crispac Connatae Viticella Oliver 8 87-38, 87-78 88-32, 87-39 86- la, 86-42 88-3 39 86-45 86-¢ 88 47 -42 FIG. | ee eae A. Type I seedling of Clematis heracleifolia DC. B. Type IU seedling of Clematis crispa L. ©. Type I seedling of Clematis catesbyana Pursh. D. ioe I] ee of Clematis terniflora DC. RESULTS The specimens studied fall into two major categories with respect to four distinct sets of characters involving not only seedling morphology, but also aspects of the adult foliage and the achenes. The features associated with and discussed in detail e] each type of seedling are summarized in Table 3, below. A summary of Tamura’s taxa falling into the two categories, along with the taxa that have not yet been studied 1s presented in Table Seedling phyllotaxy In Type I seedlings (Figure 1A,C), the first several eophylls (seedling leaves) are alternate, and closely spaced, forming a small rosette at the apex of the hypocotyl. This alternate phyllotaxy later gives way to the opposite 383 phyllotaxy typical of the adult plants, usually at the time chat internodal elongation begins. This may happen as early as the third and fourth leaves in some taxa, but in Connatae and Clematopsis, leaves may remain alternate throughout the first juvenile shoot. The genus Archiclematis has been segre- gated from Clematis on the basis of its permanently alternate phyllotaxy. In section Meclatis, the first eophyll ts typically followed quickly by a second eophyll without internodal elongation, giving the appearance of opposite or subopposite leaves. Internodal elongation in Meclatis begins after 2 or 3 eophylls, while the leaves are still alternate. Type II seedlings (Figure 1B,D, 2L) contrast strongly in that leaves are opposite from the beginning, and several sets of paired cataphylls are produced before any leaflike eophylls are produced. Internodal elongation is present from the beginning also, even during the catophyll stage, except that the first pair of catophylls may be produced immediately above the cotyledons (e.g. in Clematis crispa L.). Tasie 3. Characters distinguishing Type I from Type Il Clematzs. Character Type | Type I Phyllotaxy of seedling alternate opposite Hypocoty] mostly elongate suppressed (except in Clematopsis) Initial shoot condensed rosette elongate Cataphylls Eophyll margin Adult foliage Regenerative buds Growth form Achenes absent toothed most often coothed and membranous, or entire and glossy-coriaceous n aerial leaf axils or (in Clematopsis) in subterranean axils of rosetce cophylls woody vines or shrubs; many rooting at aerial nodes laterally compressed, but narrow, turgic several pairs present entire often lobed or dissected but | not toothed, mostly membranous in subterranean axils of cotyledons and some cataphylls perennial erect herbs, weak- stemmed vines or sometimes woody vines; these regenerating from subterranean buds broad, very flat 384 2. Eophyll and leaf morphology. In Type I seedlings, leaf shape varies considerably (Figure 2C — K), but eophylls are typically broad, sometimes 3-lobed, with small veins diverg- ing from the peripheral region and terminating in marginal teeth. This pattern usually persists in the adult foliage, and most of the taxa with Type I seedlings have conspicuously dentate foliage throughout the plant. In Section Mec/atis the first eophylls are narrow and little-toothed (typically entire to irregularly 1-toothed — Figure 21,J), but subsequent leaves are dentate. Section Aftragene, considered a distinct genus by some authors, differs from the common form only in that the first eophylls are deeply divided (Figure 2C). In Type [I seedlings, The first eophylls (after the cataphylls) are mostly elliptic-ovate and entire, although in C. terniflora the first eophylls are sometimes 3-lobed at the tip (Figure 1D). Adult foliage may be variously divided and lobed but never toothed as in Type I species. 3. Cotyledon, hypocotyl orientation, and habit. In the terminology of Duke and Polhill (1981) most Type I seedlings are phaneroepigeal, 1.e. the cotyledons and epicoty! are elevated above ground by an elongate hypocotyl. As adults, Type I plants are mostly woody vines, or in subsection Tvbulosae, suffrutescent shrubs. Branching can occur only from aerial nodes above the hypocotyl. Many species, however, readily form adventitious roots when aerial shoots touch the ground, and can spread quite rampantly in this way. In the three species of C/ematopsis examined, which otherwise have all the characteristics of Type I species, the cotyledons emerge from the ground, but the hypocory! does not elongate and the cotyledon bases, and initially the epicotyl, remain below ground (phanerohypogeal). Several eophylls are produced without internodal elongation, forming a small rosette, and these subterranean nodes form a rootcrown with buds that can repeatedly regenerate the plant if the top dies off due to drought, fire or normal seasonal cycles. Type II seedlings are all hypogeal, as the hypocotyl does not elongate, although the blades of the cotyledons may emerge (phanerohypogeal). In this one respect they are similar to the species of C/ematopsis mentioned above. However, in Type II plants, an elongate shoot ts produced directly, without formation of a rosette. Regenerative buds are produced in the axils of the cotyledons and some of the lowest cataphylls. he predominant growth form in Type II C/ematzs, at least in subsections Crispae and Angustifoliae, and in Clematis recta, 1s a perennial herb or weak- stemmed vine, in which stems die back to the ground each winter. The 385 underground rootcrown established by the seedlings allows for repeated renewal of the plant in successive growing seasons. Other taxa with Type II seedlings, such as Clematis terniflora and several of its Eurasian relatives in section Rectae, along with the sections Veticel/a and Patentes have persistent FIG. 2. Variation in first seedling leaf, including outgroup comparisons with Anemone (A,B), A-K represent Type | seedlings, L is Type Il. A. Anemone pulsatilla L. (Pulsatilla vulgaris Miller). B. Anemone berlandiert Pritzel. C. Clematis eae —e Miller. D. Clematis chrysocoma Franchet. E. Clematopsis & Perry. G. Clematis gentianoides DC. H. Clematis filamentosa janes — villosa as E Clematis papuasica Merr Dunn. [. Clematis ortentalis L. J. laa pie Korsh. K. Clematis microphylla DC. (one of two ee leaves). L. Clematis fusca Turcz. (first and second pair of leaves are numbered; "b” indicates position of buds in axils of cotyledons). 386 Taste 4. Summary of Tamura’s (1967) infrageneric taxa displaying Type | and Type IH morphologies. Type l Type Ll Undetermined Vitalbae Crispae Crassifoliae (prob. 1) Pierotinae Rectae (Eurasian group) Rectae (tropical Asian) (prob. 1) Dioicae Vicicella Fruticella (7) Bebaeanthera Patentes Pterocarpa (prob. II) Lastantha Angustifoliae Floridae (prob. ID Cheiropsis Naravelia (prob. 1) Aristatae Archiclematis (prob. I) Atragene Clematopsis woody stems, but even in well-established plants, new shoots can arise from the subterranean buds at the base of the original shoot. There is a strong trend toward cryptohypogeal germination in this group, in which the cotyledons remain within the seedcoat below ground. The specimens studied of Clematis viorna, reticulata, fusca, texensis, pitchert, glaucophylla, and patens were cryptohypogeal, while C. integrifolia, crispa, baldwinit, terniflora, hexapetala and kirilowii were phanerohypogeal. 4. Achene shape. Achenes 1n Type [ taxa, although laterally compressed, tend to be small and turgid, while those in Type II taxa tend to be very broad and flat, and often have a conspicuously thickened rim. Of all the specimens examined, just one appears to be intermediate between Type Land Type II seedlings. Specimens of Clematis microphylla DC from Australia have seedlings with an elongate hypocotyl, with the eophylls strongly 3-lobed and toothed. Eophylls are paired from the begin- ning, however, and there are buds tn the axils of the cotyledons, the in- ternodes are elongate after the first pair of leaves, and the achenes are broad and flat. Whether this species is phylogenetically intermediate between the two types or represents convergence or reversal in some characters remains to be elucidated through further study. DISCUSSION This survey of seedling morphology and correlated characters of the foli- age and achenes reveals two well-defined patterns in Clematis (Table 3), suggesting a fundamental and natural division in the genus that could provide the basis for clearly defined subgenera. Taxonomic division of the genus along Type I and Type II lines (Table 4) would, however, require a radical departure from the traditional system of Tamura ( 1967), which was based primarily on floral characters (Table 1). It would cut across Tamura’s two largest sections, Clematis and Viorna, and require a regrouping of the ler sections. Traditionally, section C/ematis is defined as having numerous, small, up- right flowers, usually produced in complex dichasial panicles, and with thin, spreading, whitish sepals (true petals are lacking in the genus) and glabrous stamens. Section or subgenus Vzorna, on the other hand, is characterized by relatively large, generally nodding, urn-shaped flowers with rather thick, colored, erect sepals and hairy stamen filaments, and which are either solitary or in few-flowered inflorescence units. Both sections, however, contain subgroups with Type I and Type I morpholo- gies (Table 1). The smaller sections are mostly distinguished on the basis of minor variation from one of these two patterns, and most likely will not be _ sma found to contain more than one seedling type. Adherence to the traditional system of classification (Table 1) would require the interpretation that the rather extended set of specialized Type II vegetative characters, including fundamental differences in embryonic development, evolved independently several times, presumably in response to similar ecological conditions. The alternate system, based ona division between Type I and Type II seedling morphology, requires the in- terpretation that similar floral types, particularly small white flowers produced in masses, have evolved at least twice in the genus, in response to a common pollination strategy. Which of these two alternatives most likely reflects the actual phylogeny of the genus, and should therefore serve as the basis for an infrageneric classification? The most parsimonious alternative is the latter one, 1.e. a primary division along the lines of seedling morphology, with later radta- tion and convergence of pollination types. The changes involved in seed- ling morphology are complex, involving many changes 1n the shape, vena- tion and phyllotaxy of the embryonic leaves, and in the growth pattern of the seedling axis. The changes required to shift pollination strategies are by contrast rather simple: increased branching of the inflorescence, reduc- tion in size and pigmentation of the flowers, and loss of hairs on the stamen filaments. Similar shifts have occurred in many plant families. 388 Some additional information can be interjected at this point. Data on historical hybridization within the genus, although somewhat scant, supports the natural division of Clematis along Type I and Type II lines. This genus has been popular in horticulture for several centuries and many hybrids have been made. As far as can be gathered from the horticultural literature, however, no hybrids have ever been made between Type I and ype IL taxa, even those having similar looking flowers and placed tradi- tionally in the same section. On the other hand hybrids have been made between species with small white flowers and species with large colored flowers placed traditionally in different sections, but sharing the same see- dling and vegetative morphology. Notable are the crosses between Type | taxa Vitalbae (section Clematis) and Tubulosae (sectionViorna), between Type Il taxa Rectae (section Clematis) and Crispae (section Viorna), and between Rectae and section Viticella (also Type II) (Table 5). Many of the most popular garden hybrids arose from crosses among the various large- flowered Type II taxa that are placed in separate sections in Tamura’s system. Multiple attempts by the author at hybridization betweenC /ematis verniflora DC (Type UL) and the superficially similar C. catesbyana Pursh (Type I), which are traditionally placed in the same section, caused initia- tion of achenes, but these all aborted after a few weeks. Based on the data presented here, the following phylogenetic scenario tor Clematis is suggested: Type I seedling characters and related morphol- ogy represent the ancestral or plesiomorphic condition, as they occur in related genera such as Anemone (Figure 2A,B). The ancestral population of Clematis therefore had Type I seedlings, coarsely toothed foliage, and flowers with large, colored, erect to spreading sepals and hairy stamens. These early Clematis were essentially like many members of the modern subsection Covnatae. An early lineage developed Type II characters, appar- ently in response to strongly seasonal climates. These characters included the suppression of hypocotyl elongation, the resulting hypogeal germina- tion, and the regenerating rootcrown. Within both Type | and Type II line- ages, one or more groups shifted, in parallel, to small, more numerous, white flowers with glabrous stamens, adapting to a rather common and successful pollination syndrome. Other Type | and Type II groups retained the ancestral type of flower. TAXONOMIC CONCLUSIONS Although considerable study is still needed before a complete new in- frageneric classification can be developed for Clematis, the system of sub- genera employed by Keener and Dennis (1982) and earlier workers can be supported and extended to the old world taxa defined by Tamura (1956, 389 Tanie 5. Historic hybrids among infrageneric taxa. Type | x Type I Vitalbae X Tubulosae (C. X jouintana C. K. Schneider, fide Hortus Third, 1976) Vitalbac X Tubulosae (C. X takedana Makino, fide Ohwi, 1965) Type I] xX Type Il Viticella X Crispae (C. X ertostemon Decne., fide Hortus Third, 1976) I ; “(C, X cylindrica Sims, fide Kuntze, 1885) Viticella X Florida (C. x jackmanit T. Moore, fide Hortus Third, ) Florida X Crispae (C. X durandii Durand, fide Kuntze, 1885 and Hortus Third, 1976) Florida X Patentes (C. X /awsontana T. Moore & Jackmann, fide Hortus Third, 1976)) Crispae X Rectae (C. X aromatica Lenne & Koch, fide Kuntze, 1885 and Hortus Third 1976) crae X Viricella (C. x violacea A.P. DeCandolle, fide Kuntze, 1885) Rectae X Viticella(C. X rvbromarginata, fide Lloyd 1965) 1967), with the following specific modifications suggested by the current data: 1. Type I taxa include the type species (Clematis vitalba L.) of subgenus Clematis sensu Keener and Dennis, and therefore Type I characters can be considered definitive for subgenus Clematis. 2. Type Il taxa include the type species (Clematis viorna L.) of subgenus Viorna sensu Keener & Dennis (and genus Coriflora Weber 1982), and therefore Type I characters should be considered definitive for subgenus Viorna. 3. Part of subsection Rectae (the two series, Rectae and Chinenses, defined by Tamura in 1956) should be transferred from subgenus Clematis to subgenus Viorna. The remaining series in subsection Rectae (series Crassifoliae, Meyenianae, and Uncinatae) need further study, but based on the shape of their achenes and rather different foliage, most likely will be excluded from Rectae. 4. Subsection Angustifoliae should be included under subgenus Veorna. It differs very little from subsection Rectae. 5. Tamura’s subsections Connatae and Tubulosae of his section Viorna should be transferred to subgenus Clematis. 6. Subgenus Veticella (Moench) Keener & Dennis should be reconsidered. It shows much affinity with other Type II taxa, and probably should be included as a section under Veorna. 7. Subgenus Asragene should be reconsidered. It has Type I seedling morphology and differs from subgenus Clematis only in the usual presence 390 of petal-like staminodes. It possibly should be included as a section under subgenus Clematis. 8. Clematopsis has been excluded from Clematis in the past primarily because of its broad, imbricate sepals, which contrast with the valvate sepals of Clematis. Otherwise, it has the characteristics of the genus Clematis and fits in with the old world complex of Type I taxa. Thorough study of the African Clematis is needed in order to determine the appropriate status for this taxon. Placement of other sections, and formal infrageneric reorganization of Clematis, is deferred pending more complete studies. Recognition of the two major phyletic lines in the genus, should, however, make it easier to proceed with revisionary and phylogenetic studies. It is recommended that future uses of the subgeneric taxa Clematis and Viorna reflect the changes outlined here. ACKNOWLEDGEMENTS I thank all of the individuals and institutions who kindly provided seed for this study. REFERENCES DUKE, J.A. & R.M. POLHILL. 1981. Seedlings of Leguminosae, pp. 941-949 in a _ ae Systematics, ed. R.M. Polhill & PH. Raven (eds). Royal Botanic ardens ER - KSON, . 0. 1945. The Sl ia var. rehliz population in the Ozarks. Ann. Missouri Bot. Gard. 32:413 — GRAY, A. 1895. ° el pp. | — 57, in B.L. Robinson (ed.), Synoptical flora of North America, Vol. pt ree pea Co. BAILEY HORTORIUM es Hortus ee 1976. Mac cmillan KEENER, C. S. & W. M. DENNIS. 1982. The ao. “asahention Clematis eT anes in temperate North Ape north of Mexico. Taxon 31:37 —44. KUNTZE, O. 1885. ae der gattung Clematis. Verh. Bot. ae Prov. Brandenburg 26:83 — LLOYD, C. see pon ae Life. London. LUBBOCK, J. 1892. A contribution to our knowledge of seedlings, vol. 1, pp. 78-99. Rance) D. Appleton & Co., N-Y. OHW 1, J. 1965. Flora of Japan (English ed. by E Meyer & E. Walker), p.443. Smithso- nian. SPACH, E. 1839. Histoire naturelle des vegetaux. Phanerogames 7:268 seri, - 1956. Notes on Clematis of Eastern Asia. II]. Acta Pyioea Geobot. Lor? TAMURA, va 1967. Morphology, ecology and phylogeny of the Ranunculaceae. VII. Sci. Rep. Osaka Bee 3. WEBER, W.A. 1982. New names and combinations, principally in the Rocky Mountain flora. Phytologia 51: 372 — 374 A NEW SPECIES OF CONRADINA (LAMIACEAE) FROM NORTHEASTERN PENINSULAR FLORIDA ROBERT KRAL Vanderbilt University Department of General Biology ox 1812, Station B Nashville, TN 37235, U.S.A. ROBERT B. MCCARTNEY Woodlanders, Inc. 1128 Colleton Ave. Aiken, SC 29801, U.S.A. ABSTRACT Conradina etonia, a new species ey mint endemic to northern peninsular Florida is diagnosed, described, and figured. Differences between it and its closest morphological grandiflora Small, are detailed. Attention is given to the fact that C. efonia is a relative, C. | that it narrow endemic, that it is on land being developed presently for residential use, anc ity. therefore should receive a high conservation priori Conradina (Lamiaceae) as currently treated (Shinners 1962; T. C. Gray, unpublished Ph.D. thesis 1965) is a genus of five allopatric species bicen- trically confined to the southeastern United States. One center is interior, where the arenaceous rocky bars and bluffs of Cumberland Kentucky and Tennessee streams support C. verticillata Jennison, a threatened species. The other center is lower Coastal Plain. Conradina canescens, the most abun- dant and polymorphic species, occupies coastal dunes, white sand scrub and contiguous longleaf pineland from southeastern Mississippi eastward across southern Alabama and the Florida Panhandle. Inland locally 1s the rare C. glabra Shinners, a narrow endemic growing chiefly around the rims of steepheads on the east side of Florida’s Apalachicola River. The two remaining species (C. brevifolia Shinners and C. grandiflora Small) are found in sand scrub habitat in Central Florida (Polk and Highlands counties) and in eastern peninsular Florida, respectively. As mentioned, of the four species in the southern center, only C. canescens is at once abundant within a fairly large range and exhibits considerable variation. The other three have more restricted ranges and show less variation within or between popula- Sipa 14(3):391 — 398. 1991. o92 tions. Most circumstantial evidence, both morphological and geographic, points to a relatively recent origin for the species in peninsular Florida. ll four Coastal Plain Conradina are confined to sandy soils. Major portions of Florida with deep, well-drained sand once supported extensive forests of Longleaf Pine (Pinus palustris) with a ground layer dominated by Wiregrass (Aristida stricta). This system was maintained by and dependent upon relatively frequent, low-intensity ground fires. The more xeric Long- leaf Pine-Wiregrass ecosystems are here referred to as “sandhill.” These contrast with those finer textured (often aeolean) deposits of white sand which support the scrub ecosystem characterized by Sand Pine (P. clausa) and shrubby evergreen oaks such as Quercus geminata and Q. myrtifolia and are referred to here as “sandscrub.” Historically this latter system burne infrequently but catastrophically. All who have studied the flora of the two basic systems know that both are very rich in endemics and that many species occupy very limited and circumscribed ranges within them. Scrub ecosystems, once exclusive to Florida or best developed there, have been so heavily impacted by agricultural and other development along with fire suppression that only fragmentary, often degraded, remnants exist today. Conservation agencies are actively working to preserve key areas, particu- larly in the Lake Wales Ridge area of Central Florida where an unusual concentration of rare endemics occurs. The typical scrub habitat and several of the plant and animal species associated with it reach their northern geographic limits in the center of the Florida peninsula west of Saint Augustine. One such northern exten- sion of deep sand scrub was investigated by the junior author on September 20, 1990. This area along Etonta Creek northeast of Florahome in Putnam County, was noted to have several species not only characteristic of south Florida scrub but also at or near their very northern limits (i.e. Persea humilis, Llex cumulicola, Garberia fruticosa). An endangered bird, the Scrub Jay, was also seen, but curiously the Scrub Palmetto (Sabal etonia Swingle ex Nash), named for this locality, was not observe Most interesting of all, however, was the discovery within a small area of the Etonia Scrub of a distinctive Conradina, seemingly a new species. A limited amount of material for exsiccatae and for propagation was collected and living plants from cuttings are now part of the comprehensive collec- tion of southeastern woody Lamiaceae in cultivation at Woodlanders, Inc., in Aiken, South Carolina. Dr. R. K. Godfrey was given directions to the site and, accompanied by Mr. Angus Gholson, visited the location on 12 October 1990, so as to collect an abundant sample for definitive study. This indispensable aid is hereby gratefully acknowledged. The affinities of this new plant are plainly with its nearest geographic a2 neighbor in the genus, C. grandiflora Small, which ranges intermittently in the coastal scrub of eastern peninsular Florida from Dade County north- ward to an area in Volusia County roughly 70 air miles southeast of the Etonia Creek site. In general habit and in the large size and general con- figuration and pigmentation of flower the two look much the same. There are, however, striking differences as will be explained below. We name the new discovery C. efonia, after the remarkable locality where it was found. CONRADINA ETONIA Kral & McCartney, sp. nov. Figs. 1, 2. Conradina etonia, sp. nov.; differt aC, grandiflora toliis latioribus, hebetiviridibus, nervis lateralibus distinguibilibus, utroque latere 2—4; paginissurculorum juvenorum, lami- orum foliorum et tuborum calicorum informer patenti-puberulis. Virgate shrubs to 1.5 m high, with numerous, frequently arching primary branches, the new shoots slender, ca. 1 mm thick, quadrate, downy-spreading-puberulent, scattered-glandular, reddish-brown, on older growth thickening, terete, the outer bark exfoliating in long, narrow, gray strips, exposing red-brown or orange-brown smooth inner bark. Leaves deciduous in 2 — 3 years, spreading to ascending, each node with axillary buds typically developing short, leafy shoots, thus foliage appearing fascicled-verticillate; principal leaf blades spreading to spread- ing-ascending, broadly to narrowly oblanceolate or spathulate, 15 — 30 mm long, (2) 3—9 mm wide, tips rounded to broadly acute, margins nar- rowly and tightly revolute, base narrowly cuneate to attenuate on a short (less than | mm) petiole, the adaxial blade surface uniformly downy- spreading-puberulent, copiously and punctately gland-dotted, dull green, the midrib evident at base of a strong median groove, the exposed abaxial surface concave, slightly paler, the puberulence very dense, the midrib strongly raised, likewise with dense spreading puberulence and producing 2 — 4 strong branch nerves per side (a unique trait in this genus!). Cymes produced from all or most nodes from midstem up, (1) 3 — 7 (12) flowered, either subsessile or on spreading-ascending peduncles to 2 (3) mm long, these and the ascending branches all densely and uniformly downy- puberulent, their successive nodes with progressively reduced, narrower, decussately-arranged pairs of similarly downy bracteal leaves, pedicels erect or spreading-ascending, 1—3 mm long, ebracteolate, eoieaine: puberulent. Calyx at anthesis bilabiate, 7.5-—8.1 mm long, the tube cylindro-campanulate, 5 mm long, distally densely villous-annulate within, otherwise smooth, externally 13-nerved, with nerves uniformly downy and intervals smooth to downy, strongly gland-dotted, the upper lip upswept (1.8) 2—2.7 (3) mm long, tridentate, the teeth broadly triangular, strongly hirsute-ciliate, connivent; lower lip directed forward 394 . Conradina etonia (Godfrey with Gholson 84008). — a. Flowering branchlet. — b. Sector of old shoot with branchlet base. — c. Adaxial (left hand) ee abaxial (right hand) views of stem leaf. d. Small sector, ideal cross-section, adaxial surface of leaf (left) and that of abaxial surface (right). —e. Side view of flower at anthesis. 37) L5cm FIG. 2.€ fi ia (Godfrey with G holson 84008) a. A range of pancipels shoot leaf types and sizes drawn to scale. — b. Oy 1 corolla (with small portion of tube removed) to show inner surface, stamens. — c. Gynoecium, ne view. — d. Adaxial (left) and abaxial (right) view “of anthers. — e. Sia base, ideal view, — placement of nutlets on gynophore (left) and a median longisection showing placentation (right). . Calyx, opened to show fruit placement, annulus. — g. Opened calyx to show arrangement of ce teeth. 296 and upcurved, 2.5 — 3.1 mm long, cleft to near base, the teeth narrowly triangular-subulate, likewise variably hirsute-ciliate with hairs under | mm. Corolla strongly bilabiate, 20-25 mm long to tip of lower lip, the slender tube gradually broadened toa geniculation at throat base ca. 2 mm above calyx sinus, funnelform to level of limb, from base to limb densely villose within, upper lip galeate, broadly oblong, 10— 11 mm long, the keel continuous with the upper side of throat, projected outward and for- ward as a strong arch, apically emarginate; lower corolla lip bent abruptly down, slightly longer, trilobed, the laterals ascending-oblong, producing an ascending, oblong-rounded extension, the central lobe shortest, obcor- date; corolla tube and throat adaxially lavender-blue to lavender-rose, the upper lip uniformly lavender, the lower lip and throat medially with a broad longitudinal zone of white or cream mottled with spots and streaks of deep purple, the bordering lobes and edges again uniformly lavender, all edges intermittently pilose-ciliate, the external surfaces with a mixture of downy and villous hairs and gland-dotted. Stamens tetradynamous, both pairs roughly paralleling the upper lip and directly under its keel, the shorter pair extending ca. to its tip, the longer slightly beyond, arching oucward and downward, the anthers slightly divergent on broad connec- tive, ca. | mm long, broadest across the base, the thecae dark purple, white-villous-pilose along the dehiscence line. Style sigmoid, its bifid- stigmatose tip exserted slightly beyond level of anthers. Nutlets mostly 4, obovoid, |— 1.2 mm long, brown, very finely reticulate toward apex — Type: U.S.A. FLORIDA. Purnam Co.: Sand Pine Scrub, undevel ae subdivision, SE on Fla. rd 100, 2 mi from Florahome thence to left on Holladay Rd. 1.9 mi to Blossom Rd. on i thence to intersection Blossom and Garden St. , 20 “ 1990, a B. McCartney, . CHOLOTYPE: SMU; isorype: VCB) dditional | eee examined: Toporyprs: 12 Oct 1990, R.K. Godfrey oe Angus ase ee K FLAS, FSU, GA, GH, K, NCU, NY, PH, RSA, TENN, T IC , US, VDB, VSC, WAT, WILLD. The habitat in detail is a deep white-sand scrub dominated by Pinas clausa, Quercus chapmanii, Q. geminata, Q. myrtifolia, with some Q. laevis and a mixed understory of Serenoa repens, Smilax, Persea humilis, Asimina, Ceratiola, Sageretia minutiflora, Uex cumulicola, Licania, Rubus cuneifolius, Garberia, and herbs such as Eapatorium compositoflium, Liatris chapmanii, Balduina angustifolia, Carphephorus corymbosus, Chrysopsis scabrella, Palafoxia integrifolia, Elephantopus, Polygonella. The Conradina are most abundant in sandy natural or artificial clearings, less so in the scrub itself; it would appear, therefore, that this mint responds abundantly to disturbance (his- torically this probably fire) as do many of the other shrubs and herbs of this habitat type. Bot Those who work with Conradina will agree that C. etonia as described here shares more character states with C. grandiflora than it does with any other Conradina, that it shares similar habitat, and that it 1s closest geographically to that species. Thus the burden of proof is in establishing _ most apparent differences in character between the two: . Indumentum of young shoots of C. efonza is puberulent, the hairs thus minute, spreading, downy; that of C. grandiflora, likewise minute and abundant, is mostly upcurved. That of inflorescence branches in both is also abundant and minute, but that of the former is again spreading, more uniform, while in the latter it may be admixed with some pilosity. 2. Leaves of the two differ strongly, those of C. etonia being distinctly broader with lateral veins quite evident on the undersurface, a feature held by no other known species of Conradina; both adaxial and abaxial surfaces are dull green, each witha uniform spreading-and-downy puberulence (the texture of this adaxial side much like that of the undersurface of Chalk Maple leaves!), that of the adaxial surface slightly less than that of the paler, abaxial side. In C. grandiflora the adaxial side of the leaf is dark green, lustrous, with a minute, incurved, harder pubescence, that of the abaxial side is whitened by a dense mat of white, short, appressed tomentulum and shows no branching from the usually sparsely strigillose midrib. 3. Dimensions, shapes, and pollination guide markings in flowers of both species overlap very much, with calices extremely similar 1n size- range, but those of C. etonia (again) with puberulence denser and more uniform (downy), very rarely with the tube having any pilosity and this confined to a few “whiskers” at or toward its base; in C. grandiflora most specimens show a very strongly gradate pubescence of calyx tube, from some long pilosity over 1 mm to an admix of shorter pilosity, some gland- tipped, and puberulence. Calyx teeth in both are hirsute-ciliate, but such airs in C. efonia are shorter, more uniform. Corollas of these species are hardly distinguishable Stamens of the two species are again very similar as to size and shape, but while anthers of C. etonia are bedecked with a distinctive long fringe of white pilose-villous hairs, those of C. grandiflora have fewer and shorter hairs, these tending to be concentrated more at anther sac tips and bases. The features detailed above convince us that C. etonia could well be the best-marked species in a genus whose species differ mostly in very fine characters. We are hopeful that further exploration of the northern lobes of Florida scrub habitat will yield more records for this beautiful new species. Indeed, it may show us some intermediates. A clue to this is a Volusia County collection of C. grandiflora (Sand Pine sandridge by I-95, 0.6 mi S Port 398 Orange exit, 28 Aug 1974, Kral 54022) in which new shoots have an atyp- ical spreading downiness similar to that in our new species. But even this material 1n no other way resembles C. efonia. Because Conradina etonia is rare and perhaps restricted to a small area of highly vulnerable habitat, we have attempted to expedite the publication of its description and initiate appropriate protection strategies. The known range is within a subdivided tract with streets roughed-in and a few residences built. Botanists, the general public, and affected property owners will hopefully refrain from actions which might further jeopardize the survival of this species and will support efforts to conserve it. REFERENCES pe T.C. 1965. A sad es of the genus Conradina A. Gray (Labiatae). Unpublished Ph. ods, Vanderbilt Unive KRAL, ; - 198 report on some rare, threatened, or endangered forest-related vascular ee of - Sh, USDA Forest Service Tech. Pub. R8-TP2. 1305 pp. SHINNERS, L. H. 1962. cae of Conradina se Sida 1(2):84 — 88. SMALL, J. K. 1933. Manual of the southeastern flora. Publ. by the Author. New York. OBSERVATIONS ON FRYXELLIA PYGMAEA (MALVACEAE) PAUL A. FRYXELL U.S. Department of Agriculture in cooperation wit Texas AGM University College Station TX 77843, U.S.A. JESUS VALDES R. Universidad Autonoma Agraria “Antonio Narro” Saltillc, Coahuila, MEXICO ABSTRACT The rediscovery of the rare Fryxellia pygmaea (Correll) Bates in central Coahuila ts repor- ted, and its ecology and taxonomic affinities are discussed. The plant has a chromosome count of 27 = RESUMEN Se reporta el redescubrimiento de una poblacién de la infrecuente Fryxe/lia pygmaea (Correll) Bates de la parte central de Coahuila, y se discute su ecologia y sus afinidades taxonomicas. El numero cromosomico de la planta es 27 16 The monotypic genus Fryxellia has been one of the least known genera of the Malvaceae. The type was collected by Capt. John Pope in 1854 at an unknown locality in Texas, probably west of the Pecos River (Correll 1968 Bates 1974). The species was subsequently recollected by Robert M. Stewart in 1941 near Puerto del Aire near the southern end of the Sierra de la Encantada in Coahuila. Originally described in the genus Anoda (Correll 1968), the plant was recognized by Bates (1974) to be distinct and to con- stitute a monotypic genus, isolated from other genera of the tribe Malveae (Bates 1974; Fryxell 1988) as the Fryxellia alliance. In early September 1990 we had the opportunity to revisit Puerto del Aire (Fig. |) in an attempt to relocate the plant. We succeeded in finding a population of several hundred plants, perhaps the same population found by Stewart. The population was restricted to a relatively small area of “dry open hillside.” The area occupied by the population was perhaps 100 — 150 m in diameter, beyond which no plants were found. Within the area, how- ever, the population was relatively dense, with individual plants occurring within a meter or two of one another. The population was clearly an old Sipa 14(3):399 — 404. 1991. 400 one, with many individual plants having extensive perennial rootstocks. What edaphic or other factors are involved in restricting the population to this small area were not immediately evident and are not known. It may be speculated, however, that the population observed is in fact an ancient clone, spreading laterally by proliferating rootstocks. Even though the plant has an apparently efficient method of seed dispersal, and subsequent germination tests have shown the seeds to be fully viable, seedling es- tablishment at a new locality in the severe desert environment of central wy Bue oe a | : h s : + wig ee me “EL RI _ Br rd ia ‘| sie, S weil We Pa’ & >a tts. = ME = ge od Oy i Pa SG .q ra eA es : si Hacienda de Cabrae = i Bt 28) 27] 26) 25] 24 ae] 23 ABE 19) 18] 17} 164157 146) 13) 12})11) 10) 9 By 7] 6) SE 4} 3) 2) 1] 0 ay VLIYOW VI 8 FIG. 1. Portion of “La mone map, p. 23, «Maps of the Chihuahuan Desert Region” (compiled by José Garcia), to accompany yazetteer a the Chihuahuan Desert Region, A Supplemenc to the Chihuahuan Desert Flora” (by J. Henrickson and R.M. Straw, 1976). ~ 401 Coahuila may bea relatively rare event, dependent upon a favorable pattern of rainfall in a particular year. A perennial, caespitose habit, coupled with vegetative propagation by root proliferation, may be the secret of survival for this species in this habitat. Unfortunately, we did not consider this possibility at the time we were in the field when, as a test of this hypothe- sis, we could have looked for subterranean interconnections of adjacent plants. The Plants Summer rainfall in this area had been above average in 1990, so that the plants were in relatively good condition. All except the youngest plants were fruiting, with flowers and buds still developing. Fruits were relatively abundant, indicating that the plants were vigorously reproductive. In- dividual plants form small rosettes on the order of 15 cm diameter. They are not acaulescent (as described), although they are indeed caespitose. Each plant has one to several short stems | — 2 cm long with very short, crowded internodes. The rootstocks are thick (ca. | cm or more in diame- ter), well branched, and penetrate deep into the gravelly soil; the roots are apparently food-storage organs. Buds and flowers were observed and photographed, on the basis of which corolla color can be described as a rich orange (Fig. 2). After abscission of the corolla, the accrescent calyx ultimately flares to a rotate form and, as the fruit matures, takes on a reddish coloration on the exposed (adaxial) side. Upon maturation of the fruit, abscission is at the base of the calyx, so that the calyx and the contained fruit together are the diaspore. The calyx evid- ently serves as a sail to be blown over the ground as a form of wind dis- persal. Styles and stigmas were observed and were found to have the abruptly capitate form and glabrous condition that are characteristic of the genus Anoda. These features, together with the general aspect of the fruits, ex- plain Correll’s original placement of this species in Anoda. On the other hand, the dorsal spur, which accounts for the resemblance of the mericarps of Fryxellia to those of Anoda, differs in that the spur clearly has a suture of dehiscence in Fryxellia and but not in Anoda. The sabe 3s F. pygmaea (Correll) Bates are well illustrated by Bates (1974, Fig. , Ac). One item of information can be added, however. The ene is in fact a divided structure, consisting of two awl-like internal growths extending forward from the dorsal wall, not a single such growth as was illustrated. Thus, the endoglossum shows a resemblance to that found in the genus Batesimalva, most nearly to that found in B. pulchella Fryx. FIG. 2. Fryxellia pygmaea. Plant grown in the greenhouse from seed. 403 Questions remain concerning the affinities of the genus Fryxe/lia. The stigma morphology suggests an affiliation with Anoda. The endoglossum structure suggests an affiliation with Batesimalva, as does the leaf form and geographical distribution. Furthermore, the accrescent calyx of E pygmaea shows some resemblance to the somewhat accrescent calyx of B. pulchella. However, other characters, such as the caespitose habit, the orange corolla, the strongly accrescent calyx, and the detailed fruit morphology, clearly justify Bates’ segregation of Fryxel/ia as a distinct genus. Conceivably, Fryxellia may be a connecting link (by reduction of the upper cell of the mericarp, with the endoglossum remaining as a vestige of this former, hypothesized condition) between Batesimalva and Anoda and thus provide an indication of the phylogenetic origin of the Anoda-Periptera alliance. A chromosome count of 27 = 16 was obtained for FE pygmaea (Fig. 3). The base chromosome number for Anoda is x = 15 (Bates 1987), for Batesz- malva x = 16 (Bates & Blanchard 1970). Pollen aperture number (Hashmi 1970, Fryxell 1988), is 3 for Fryxeflia, 3 — 4 for Batestmalva, and usually 30 or more for Anoda. These data indicate a placement of Fryxe/lia closer to FIG. 3. Chromosomes of Fryxc//ia pygnaca. Top, metaphase 1( 2160); bottom, diplotene (X 833). 404 Batesimalva than to Anoda. Finally, it may be asked if this species should be considered as “threatened or endangered.” The plants observed were locally abundant and highly fruicful, producing abundant viable seeds, but were very loca- lized in distribution. What ecological factors mediate this localization are unknown. Only one population of the species is certainly known, and the species can arguably be described as the rarest plant of the Chihuahuan Desert. Yet it was also collected in Texas by Pope, probably somewhere west of the Pecos River, a direct distance of 200 — 300 km (or more) to the northwest. An ample amount of relatively undisturbed, apparently suitable habitat lies in the intervening area, in which the species might be expected to occur. Much of this area is unexplored botanically or at least poorly explored. Therefore, it seems more suitable to describe this species as “insufficiently known” rather than “threatened, endangered, or extinct,” as listed by Valdés and Johnston (1988). The recent collection, duplicates of which will be distributed, is cited as follows: oO. EXICO: Coattuita: Mpio. de Ocampo, Sierra de la Encantada (28° 4-1/2’ N, 102° 25’ W), alt. 1250 m, 8 Sep 1990, Fryxell, Valdés, Carranza, Vazquez & Meza 5006 (ANSM, BRII-SMU, pf, and other duplicates to be distributed). ACKNOWLEDGMENTS Appreciation is extended to Miguel A. Carranza P., Ricardo Vazquez Aldape, and Orlando Meza, who helped with the field work, and to Juan Diaz-Colon and David M. Stelly, who provided the chromosome prepara- tions and photographs. REFERENCES BATES, D. . 1974. Fryxellia, a new genus of North American Malvaceae. Brittonia 26:95 — 100. — ES, D. - 1987. Chromosome numbers and evolution in Anoda and Periptera Malvaceae). Aliso 11:523— 531. oat ,D.M. and O.J. BLANCHARD, Jr. 1970. Chromosome numbers in the Malvales. New or otherwise noteworthy counts relevant to classification in the Malvaceae, tribe ilveac. Amer. J. Bot. 57:927 —934 ( ORRE LL, D:S. 1968. Some additions and corrections to the flora of Texas-VI. Wrightia 74-78 oe PA. 1988. Malvaceae of Mexico. Syst. Bot. Monogr. | — HASHMI, S.H. 1970. The palynology of the Malvaceae of Texas. cn Dictonon. Texas A&M Ue (University Microfilms no. 70-16729) VALDES , J. and M.C. JOHNSTON. 1988. Botanical resource and floristic diversity desterion? in the Chihuahuan Desert Region of Mexico. Third Symposium on Resources of the Chihuahuan Desert: U.S. and Mexico (abstract). 10 — 12 November 1988 at Sul Ross State University, Aipiae Texas. — BIOCIDAL SIDA (MALVACEAE) CHARLES L. BURANDT, JR. Department of Biology, Texas A G I University Kingsville, TX 78303, U.S.A. ABSTRACT The secretions of glandular trichomes of two South American species of Sida are highly toxic to ants and cockroaches. In natural habitat, these secretions most likely provide resist- ance to herbivory by insect or other small arthropods. INTRODUCTION During preparation of a monographic treatment of Sida section Oligan- drae (Malvaceae) two species were observed with an interesting vestiture of stalked glandular trichomes (Fig. 1). One species, Sida jatrophoides L Héritier, is a xerophyte occurring in scattered populations from coastal Peru and the Galapagos. The other, Sida palmata Cavanilles, is a mesophyte occuring along trailsides in northern Peru and southern Ecuador (Burandt 1992). Touching the young stems or petioles of either species left the skin covered with a moist but not sticky film, evidently the secretion of these trichomes. Several observations coincided to suggest and support a hypothesis that the trichome secretions in these species might function to repel or possibly kill walking arthropods: |. The droplets of secretions formed at the eleva- ted tips of the trichomes (Fig. 1) would undoubtedly contact an insect walking on the plants. 2. The trichomes were more densely distributed (especially in young S. jatrophoides) in the inflorescence branches (Fig. 1) and in the petiole area just below the blades. A walking insect would be obliged to cross a veritable mine field of droplets to get to the presumably more delectable leaf blades and flowers. 3. Plant populations studied in the field appeared relatively free of insects, and the leaves were free of evidence of insect herbivory. The lethality of morphologically similar glandular trichomes to arthro- pods has been frequently reported Juniper and Southwood 1986). While toxic compounds may also be present in trichome secretions (Carter et al. 1989, Walters et al. 1989, Dimock and Kennedy 1983, Gerhold et al. 1984, and Williams et al. 1980), their role in pest resistance 1s complica- ted by the fact that most trichome secretions are adhesive and the principal mode of action is by trapping - larvae or adults are immobilized until they Sipa 14(3):405 — 410. 1991. 406 die. Conversely, the trichome secretions of Sida sect. Oligandrae are non- adhesive and any antibioses observed for the these secretions would likely be attributable to the effects of toxins alone. This study presents experi- mental evidence which strongly supports the hypothesis that natural toxins in the secretions in Szda section Oligandrae potentially function as deterrents to arthropod herbivory. METHODS Plants of S$. palmata and S. jatrophoides were germinated from scarified seeds placed in 75 to 95 mm plastic pots filled with sand/clay/peat mix. Seedlings were kept in a growth chamber illuminated with fluorescent “gro-lights” and set fora 12 hr photoperiod with 17° night and 27° C day temperatures. They were supplied with deionized water and commercially available fertilizers. To encourage flowering, fertilizers were discontinued, soils were leached by excess provision of deionized water, and plants were allowed to become water-stressed. Inflorescences were well developed after six months of growth, and trials were begun. Toxicity of the secretions of these species was assayed using fire ants (Solenopsis invicta L.) and German cockroaches (Blattella germanica Buren). Ants were collected as needed from field colonies, whereas immature cock- roaches (5 — 10 mm in length excluding antennae) were gathered from a laboratory-maintained colony. Active specimens of these insects were subjected to various topical applications as described below. After treat- ment, test and control specimens were housed in covered 500 ml glass con- tainers for observation. SIDA PALMATA assay: Trial no. 1. Fifteen ants were individually gathered with an artist’s small paintbrush and repeatedly pressed against secretory trichomes of S, palmata. Ants were then placed collectively in a container. Asa control, 15 ants were “jostled” with a clean paintbrush and placed collectively in a separate container. Mortality was recorded at 21 hrs. rial no. 2. Ten ants were similarly treated but placed in 10 individual containers. Asa control, 10 ants were repeatedly pressed against trichome- bearing parts of a dried specimen of S. pa/mata and placed in 10 individual containers. A second control consisted of 10 otherwise unmolested ants placed in 10 individual containers. Mortality was recorded at 21 hrs. Trial no. 3. Ten ants were individually gathered with a paintbrush and gently placed on plant parts bearing numerous secretory trichomes. If necessary, they were coaxed to walk sufficiently to come into contact with trichomes. Ants that fell were retrieved and again placed on trichome- 407 FIG. 1. Above: branches of leafy inflorescence of §. jatrophoides, below: petiole of S. jatrophoides (ca. 1.25 mm diameter). 408 bearing areas. Ants were then placed in 10 individual containers. As a con- trol, 10 ants were placed individually in 10 containers and, using a clean paintbrush, swabbed with deionized water. Mortality was recorded at 7 i. Trial no. 4. Twelve cockroaches were placed individually in glass con- tainers and anesthetized with CO® gas. Secretions were then collected by passing an artist’s small paint brush over appropriate plant surfaces until bristles were saturated. Cockroaches were then “painted” on their ventral sides with the secretions. As a control, 12 roaches were placed individually in glass containers, anesthetized with CO’, and “painted” on their ventral sides with deionized water. Mortality was recorded at 5 hrs. SIDA JATROPHOIDES assay: Trial no. 1. Twenty ants (two combined trials) ants were individually gathered with a paintbrush and gently placed on plant parts bearing numerous secretory trichomes. If necessary, they were coaxed to walk suf- ficiently to come into contact with trichomes. Ants that fell were retrieved and again placed on trichome-bearing areas. Ants were then placed in 20 individual containers. As a control, 20 ants were placed individually in 20 containers and, using a clean paintbrush, swabbed with deionized water. Mortality was recorded at 8 hrs. Trial no. 2. Thirteen cockroaches were individually placed in glass con- tainers and anesthetized with CO’ gas. Secretions were then collected by passing an artist’s small paint brush over appropriate plant surfaces until bristles were saturated. Cockroaches were then “painted” on their ventral sides with the secretions. As a control, 13 roaches were placed individually in glass containers, anesthetized with CO’, and “painted” on their ventral sides with deionized water. Mortality was recorded at 5 hrs. Trial no. 3. Seven cockroaches were treated as in the preceding trial but “painted” on their dorsal sides with the secretions of S. jatrophoides. As a control, 7 roaches were treated as above but “painted” with deionized water on their dorsal sides. Mortality was recorded at 5 hrs. RESULTS INSECT BEHAVIOR Initial responses of both roaches and ants to application of plant secretions was similar. Brief episodic whole-body convulsions and tremors occurred sporadically within the first few minutes. Individual legs became sporadically or continuously rigid and were dragged or remained variously skewed to the side or to the rear. Rarely, an appendage would disarticulate. Effective walking gradually became impossible and specimens collapsed 409 Taste 1. Lethality of secretions of S. palmata and §. jatrophoides to ants and roaches. S = secretions applied, C = control. For each, the number dead precedes the number treated (N). Trial Number: ] 2 3 4 Total S. palmata Ants ae 15) 115) 9(10) 2(10) 0110) 610) 010) — - 24(35) 3(45) Roache - = = — — = 1212) O(12) 1212) 0(12) = S. jatrophordes Ants 19(20) 2(10) - - - — — = a 19(20) 2(10) Roaches 13(13) 0(13) 7(7) 07) — - - - — 20(20) 0(20) Grand Totals Ants 43(55) 5(55) Roaches 32(32) 0(32) =] > and could not right themselves. In ants and occasionally in roaches, these symptoms appeared to subside within 30 minutes of treatment but later returned, however, with increasing dysfunction eventually involving all appendages. A few ants appeared to completely recover. Insect MorvTaLiry At natural concentrations, secretions of the glandular trichomes of both S. palmata and S. jatrophoides were very lethal. Roach and ant mortality pertrial and per control(s) are summarized in table 1. Of 35 ants variously treated with secretions of S. palmata, 24 died whereas only 3 of 45 control ants died. All 12 roaches treated with S. palmata died whereas none of the 12 control roaches died. Of 20 ants treated with secretions of S. jatrophoi- des, 19 died whereas only 2 of 10 control ants died. All 20 roaches treated with secretions of S. jatrophoides died whereas none of the 20 control roaches died. Totaling the toxicity assays using ants, 43 of 55 of those treated with plant secretions died whereas only 5 of 55 control ants died. Totaling the toxicity assays using cockroaches, 32 of 32 treated roaches died whereas none of the 32 control roaches died. DISCUSSIONS All plants exhibit potential resistance to herbivory. Deterrent morpho- logies range from the simple mechanical resistance provided by toughened tissues to the honed injection devices of the trichomes of Urticaceae. An array of molecular defenses is also available to plants. Ordinary sap, exuded as a result of wounding, may be mildly repellent, or lethal toxins such as pyrethrins and nicotine may be produced. 410 Most plant structures and molecules serve several functions, however, and their roles in herbivore resistance is often subsidiary to other functions and difficult to establish. The dramatic lethality of trichome secretions of Sida 1s thus more remarkable since it appears to be due solely to the effect of toxins. That such a specialization might evolve from glandular trichomes with broader methods of antibiosis is not surprising but is, nevertheless, apparently rare or little reported. It should be interesting to survey addi- tional taxa and assay for toxicity those which possess similar non-adhesive trichome secretions. ACKNOWLEDGEMENTS I thank John Mellen and Cindy Galloway for useful suggestions during the course of this study, Hugh Lieck for photography, and Carol Altman for preparation of the manuscript. REFERENCES BURANDT, C.L., JR. A monograph of Sida sect. Olrgandrae (Malvaceae). 1992. Syst. Bot. In press. CARTER, C. D., T. J. GIANFANGA, and J. N. SACALIS. 1989. Sesquiterpenes in glandular trichomes of wild tomato species and toxicity of the Colorado Potato Beetle. J. Agri. Chem. 37:1425 — 1428. DIMOCK, M. nd G. G. KENNEDY. 1983. The role of glandular trichomes in there- sistance of ene hirsutum {. glabratum to Heliothis zea. Ento. Exp. App. 263'— 268 CinnorD: D. ‘. R. CRAIG, R. MUMMA. 1984. — of trichome exudate from mite-resistant geraniums. J. Chem. Ecol. 10(5):713— 722. JUNIPER, B. and R. SOUTHWOOD, Eds. 1986. oe and the plant surface. Edward Arnold. sige WALTERS, D. S., R. CRAIG, and R. O. MUMMA. 1989. Glandular trichome exudate is the i factor in geranium resistance to foxglove aphid. Entomol. Exp. Appl. 53: 105 — 109. WILLIAMS, W. G., G. G. KENNEDY, R. T. YAMAMOTO, J. D. THACKER, and J. BORDNER. 1980. Tridecanone: a naturally occuring insecticide from the wild tomato Lycopersicon hirsutum t. glabratum. Science 207:888 — 889. THE GENUS HOSTA TRATT. (LILIACEAE) IN KOREA MYONG GI CHUNG! Botany Department, University of Georgia Athens, GA 30602, U.S.A and JONG WON KIM Department of Biology, Pusan National University Pusan 009-735, REPUBLIC OF KOREA ABSTRACT Based on the result of a biosystematic study that employed phenetic analyses of morpho- logical and enzyme electrophoretic data, and fieldwork, six Korean species can recognized: Hosta yingeri S. B. Jones (Tae-huk-san, So-huk-san, Hong islands); H. capitata (Koidz.) Nakai (southern Korea); H. clawsa Nakai (central and northern Korea); H. minor (Baker) Nakai aes and middle-eastern Korea, pa Wan and Ko-jae islands); taquetil (Lévl. Fedde) M. Chung & J. Kim comb. . (=H. venusta EF ae (Che-ju lend a — H. jonesti M. © Chung (southern islands). Keys, typications, synono- mies, descriptions, and distributions are includec INTRODUCTION Hosta is a horticulturally important genus of approximately 22 — 25 species of herbaceous perennials restricted to eastern Asia (Chung and Jones 1989; Jones 1989). Many species and cultivars are widely grown in shady gardens in Asia, Europe, North America, and New Zealand (Chung 1990; Jones 1989). Numerous nomenclatural and taxomonic problems exist within the genus (Aden 1988). Bailey (1930), Stearn (1931), Hylan- der (1954), and Lee (1957) all pointed out that Hosta is taxonomically confused genus. The taxonomic difficulty has been attributed to the pres- ence of relatively few diagnostic characters on dried herbarium specimens (Hylander 1954). In addition, many species of Hosta are so variable ecolo- gically and morphologically that a proper species concept requires morpho- logical, ecological, and biosystematic studies (Chung 1990). Over 2,500 cultivars further confound the taxonomic status of several Hosta species. These difficulties have given rise to broad (Fujita 1976) or narrow (Maekawa 1940, 1969) species concepts; 15 or 25 Japanese species, respec- tively. Although Fujita carefully described morphological characters and ‘Current address: ees of Biological Sciences, RO. 1059, Rutgers University, Pis- cataway, NJ 08855, U.S.A. Sipa 14(3):411—420. 1991. 412 ecological and geographical distributions, his studies were confined to Japan and did not include Korean or mainland Asian taxa. MATERIALS AND METHODS Samples of Hosta rootstocks (978) were collected from 45 localities in South Korea and two on Tsushima Island (H. tswshimensis N. Fujita), Japan (Chung et al. 1991). Observations were made of the habitats, and notes were taken on characteristics of the populations. The rootstocks were grown under uniform conditions in the Botany Growth Facilities at the University of Georgia. Voucher specimens of all collections are ee at GA, SNU, and KYO. Herbarium materials from BH, KYO, L, MO, NA, NY, PE, SNU, TI, and US were ex carne in more: to search type specimens of Korean Hosta taxa and determine the total range of geographic variation and the distribution of each taxon, especially those collected from North Korea and Manchuria, China. In addition, the Herbaria at Tokyo University (TI) and Kyoto University (KYO), where the majority of type specimens of Hosta are kept, were visited by MGC TAXONOMY The taxonomic treatment presented here is the result of a biosystematic study that employed phenetic analyses of morphological data (Chung 1990; Chung and Jones 1990), data from enzyme electrophoresis (Chung et al. 1991), and fieldwork (Chung 1990; J. Kim pers. comm.). Four complexes, one with three species (H. minor, H. taquetit{ =H. venusta}, and H. jonestt) and the remaining three each with a single species (H. yinger1; H. capttata, and H. clausa) can be recognized in Korea. This conclusion conflicts with the sectional treatments of Hosta by Maekawa (1940) and Fujita (1976). For example, Maekawa (1940) and Fujita (1976) treated H. capttata, H. minor, and H. taquetit in section Lamellatae F Maekawa because these three species have ridged scapes. Results by the study of Chung (1990) and Chung et al. (1991), however, showed that H. minor and H. taquetit are Closely related to each other, while H. capitata is quite distinct. In addition, H. tswshimensis and H. jonesii, which belong to section Tardan- thae (E Maekawa) E Maekawa (Chung 1989; Fujita 1976) show close relationships with H. minor and H. taquetiz. Thus, sectional treatments of the Korean hostas must wait until the entire genus has been examined. While examining loans from the 12 Herbaria and specimens at TI and KYO, four type specimens: H. clausa (Nakai s.n.), H. clausa var. normalis EF Mackawa (Nakai 5255), H. ensata FE Mackawa (Nakai 5253), and H. /on- gipes var. alba Nakai (Utiyama 5.n.) were not encountered. Probably, these four type specimens were lost (H. Ohba pers. comm.). We therefore 413 designate lectotypes for two names, H. clausa var. normalis and H. ensata, and neotypes for two names H. clawsa and H. Jongipes var. alba. A KEY TO THE KOREAN SPECIES OF HOSTA 1. Bracts, pappillous at apex, ae when fresh, remaining green at flower- ing, and persistent with the frut 2. Scapes ridged or aay st 3. eaf 6.5— 10 cm long; a. os inner and outer perianth lobes equal CO subequal iis Genelia setae eae ee ees A. taquetii 3. Leaf 14—35 cm long; outer perianth lobes longer than inner ODES cctiqets 48, ieee a hag Se ae oa wit re ees wie nae Suen ee ae H. minor 2. en smooth, not ridged 1. Racem ne transparent lines on perianth ca. 5 mm long; stamens 3+3 (the exceptional length of the second set of SCAMMERS) here a etai Shee hae el et eb hee 4 Seedab ee Sake HA. yingert 4. Racemes secund or subsecund; transparent lines on perianth ca. 15 fim lone; lene th Or stamens Same, 94444404544 ch haem dne howd 4 H. ponest 1. Bracts, not pappillous at apex, whitish green when fresh, fading to whitish bre at flowering, not persistent with the fruits. = Scapes eicece or longitudinally striate; leaf blades scabrous on th nerves below; inner perianth lobes ca. 14 mm wide; anthers whitish yellow with Sample OS 3 eee le gee oe he be eee eae awe H. tapitata 5. Scapes smooth, not sles leaf blades smooth on the nerves below; inner perianth lobes ca. 9 mm wide; anthers dark purple ........ H. clausa Hosta ciausa Nakai, Bot. Mag. (Tokyo) 44:27. 1930. Funkia lancifolia auct. non Sprengel; Komaroyv, Fl. Manshur. 2:328. 1901. Fankia ovata auct. non Sprengel; Komaroy, ib. 2:329. 1901. Hosta oe auct. non Engler; Nakai, J. Coll. Sci. Imp. Univ. Tokyo 31:250. 1911. ta . non (Andrew) Tratt. ; Nakai, ib. 31:251. 1911. Fankia lancifolia auct. non ee E Czerniakovska in Komarov, Fl. URSS, 4:55, ¢. 9 f. 2. 1935. — Type: KOREA. Prov. KYEONG-GI- wang-nung, n.d., Nakai s.n. a: v.). — Neoryee, here designated: KOREA. xe KYEONG-GI-Dbo. Kwang-nung, 16 Aug 1929, Lim s.n. (SNU!). Hosta japonica Thunb. var. dancifolia Nakai, Rep. Veg. Diamond Mts. 167. 1918. nom. ile. Hosta clausa Nakai var. normalis F bean i i Bot. 13:899. 1937. — Type: KOREA. Prov. KANG-WON-D¢ Kum-gang, n.d., Nakai 5255 (n.v.). — Lecroryer, here ee or Prov. KANG-WwON-Do. Mt. Snes 20 Aug 1902, ee sm. (TI!) — Paratypes: KOREA. Prov. Ps heh . Old plants of Par 4 functional] ssp. minima (Anderson 13343 ee. compact ae iat or ae female plant is on ‘the lefc, aad sl branched =} 1 male plane is on the right (with one of irs dichotomous branches broke xual 439 Taste 1. Some quantitative differences between subspecies of P. chartacea. Feature SSp. minima ssp. chartacea Caudex (stem base) width, mm (0.4) 0.7 — 1. ° (1.5) (1.0) ‘ : — 3. ; (4.2) Leaf width, mm 1.2—3.( Flower cluster (cymule) width, mm 1.5 Si a 20. ‘ minima are strictly annual and generally smaller, have more delicate, frequ- ently cruciform, branching and less reddish-brown pigmentation, but have colored epidermal inclusions on the stems. Paronychia chartacea from Lake Wales Ridge (i.e., ssp. chartacea) has repeatedly been described as annual (Small 1925, Core 1941, Chaudhri 1968, Ward 1977, Christman and Judd 1990, Johnson and Abrahamson 1990), but it is often a short- lived perennial. Many specimens exhibit dead tops with green, new branches developed toward the base of the plant. They form larger mats (seldom with cruciform pattern) and generally have darker coloration but lack the purple epidermal inclusions. Much of the distinction between the two subspecies is a matter of degree. Plants of ssp. minima seem somewhat less pubescent than those of ssp. chartacea. The sepal mucro is more prominent in most plants of ssp. minima; the mucro is usually shorter and blunter in ssp. chartacea. At maturity the sepals of ssp. minima frequently spread open, whereas sepals of ssp. chartacea remain directed forward. Collections of ssp. mznima are main- ly from July through October (plants collected in December were overly mature and dry; the stems were more or less disarticulated), whereas ssp.chartacea has been collected throughout the year (mainly August through March). With the description of Paronychia chartacea ssp. minima, both subspe- cies qualify as taxa of special concern and should be officially protected because of the limited range for each subspecies. The species continues to e a Florida endemic (Fig. 2), but it now has an interrupted distribution between two centers of high endemism in the state (the karst lake area 1s adjacent to the Apalachicola River system and could be considered part of that region of high endemism). Other examples linking these two areas of high endemism are rare. An example in Conradina comes to mind (see Shinners 1962). Conradina canescens (TI. & G.) Gray tends to occur in sand near the gulf coast from Wakulla County, Florida, westward through the Apalachicola region to coastal sites in Alabama and Mississippi. It has also been found inland bordering a karst lake in Washington County (Anderson 13295, FSU), but P. chartacea was absent at that particular lake. The closely related Conradina 440 brevifolia Shinners is a scrub endemic of the Lake Wales Ridge in Polk and Highland counties. Wunderlin (1982) apparently considers the two con- specific because he lists the plants from Polk and Highlands counties as C. canescens. An additional endemic, C. glabra Shinners, occurs solely in the Apalachicola River region; whereas another endemic, C. grandiflora Small, occurs just east of the Lake Wales Ridge. Two rare, closely related lupines have somewhat similar distributional patterns. Lupinus westianus Small is endemic to the Florida panhandle, and L. aridorum McFarlin ex Beckner is restricted to the Central Ridge in Orange and Polk counties. Another example is the rare lichen Cladonia perforata Evans, which involves one of these centers of endemism. This Cladonia is known from Okaloosa County in the panhandle (Wilhelm and Burkhalter 1990) and from the Lake Wales Ridge in Highlands County (Buckley and Hendrickson 1988). PARONYCHIA CHARTACEA INIMA PARONYCHIA CHARTACEA ssp. CHARTACEA FIG. 2. Range of Paronychia chartacea in Florida; each subspecies is located in an area of high endemism. Extant and historic sites are plotted for ssp. chartacea; Core (1941) listed a few collections from DeSoto County, but county lines were later redrawn, and those sites are now in Highlands County. 44] ACKNOWLEDGMENTS Field work in Bay and Washington counties was sponsored in part by the Florida Natural Areas Inventory; the agency also supplied distributional data for P. chartacea ssp. chartacea. Ann Johnson (Florida Natural Areas In- ventory), Susan Wallace (Bok Tower Gardens), and Eric Menges (Archbold Biological Station) supplied specimens of P. chartacea ssp. chartacea. Kent Perkins (FLAS) supplied information and loaned additional specimens for this study, John Thieret (K NK) graciously shared his notes on Paronychia, and Mark Garland provided the Latin diagnosis. Richard Rabeler and an anonymous reviewer offered constructive comments on the manuscript. REFERENCES BUCKLEY, A., and T. O. HENDRICKSON. 1988. The distribution of Cladonia perforata Evans on ei southern Lake Wales Ridge in Highlands County, Florida. The Bryologist 91:354— CHAU aoe M. N. 1968. A revision of the Paronychiinae. Drukkerij H. Gianotten, N. V., Tilburg, ane CHRISTMAN, S. P, and W. S. JUDD. 1990. Notes on plants endemic to Florida scrub. Florida Sci. I2 = 13. CORE, E. L. “ 41. The North American species of Paronychia. Amer. Midl. Naturalist 206: re 39 FEDERAL an 1987. Rules and regulations 52(13):2227 — 2234 (21 Jan 87). FERNALD, M. L. 1936. Plants from the outer coastal plain. Rhodora 38:414— 4952. JOHNSON, A. E, and W. G. ABRAHAMSON. 1990. A note on the fire responses of species in rosemary scrubs on the southern Lake Wales Ridge. Florida Sct. 53:138— 143. MULLER, J. W., E. D. HARDIN, D. R. JACKSON, S. E. GATEWOOD, and N. ¢c AIRE. 1989. Summary report on the vascular plants, animals and plant communities endemic to Florida. Florida Game and Fresh Water Fish Comm. Nongame Wildl. Program Tech. Rep. no. 7. 113 pp. + viii. MYERS, BL. 1990: Sas and high pine. /” os of Florida, R. L. Myers and J. J. Ewel, eds. Univ. Central Fla. Press, Orlan SHINNERS, L. H. 1962. Synopsis of a (Labiatae). Sida 1:84— 88. SMALL, J. K. 1925. A new whitlow-wort from Florida. Torreya 25:11 — 12. VANDERKLOET, S. P. 1986. Plant list of the Archbold Biological Station. Lake Placid, Fla. WARD, D. By 1977. vee to the flora of Florida — 2. Paronychia (Caryophyllaceae). Phytologia 35:4 14 — WARD, D. B. 1979. ite afd endangered biota in Florida, vol. 5: Plants. University Presses of Florida, Gainesville. WILHELM, G. S., and J. R. BURKHALTER. 1990. Cladonia perforata, the northwest Florida population. The Bryologist 93:66 — 68. WOOD, D. A. 1990. Official lists of endangered and potentially endangered fauna and flora in Florida. Florida Game and Fresh Water Fish Commission (1 August 1990), Tallahassee. WUNDERLIN, R. P 1982. Guide to the vascular plants of central Florida. University Presses of Florida, Tampa. BOOK REVIEWS GREGERSEN, Hans, SYDNEY DraApeER, AND Dierer Exz. Editors. 1989. People and Trees, the Role of Social Forestry in Sustainable Development. Available from: Publication Sales Unit, The World Bank, 1818 H Street, N.W., Washington, DC 20433. Price Un- known, Tele: 202/477-1234. (8.5 x 11) 273 pp. This book is one of a number in the EDI Sle shai Institute of the World Bank) Seminar Series designed for use in EDI courses and seminars “The distinguishing feature of social aa as distinct ae indluscrial and large-scale government forestry, is the involvement of local, vee ie rural, people in growing trees for their own use. ...The book’s main _ e is as a reference for training people who formulate policies ae design or implement programs on recognize the vital importance of eee trees into farming and pe As stems.” € text is organized into Part 1. Social Forestry and Development with 5 chapters, each oe of 3 to 7 topics and Part 2. eon: and Im Lomein Social BOresty Projects with 10 chapters, each composed of 3 to 8 topics. Seventeen autho ributed to the text with the editors integrating the chapters to eliminate cerca aid to make the chapters consistent. wfm McNEELY, JEFFREY A., KENTON R. MILLER, WALTER V. Reto, Russevy A. Mirrermerr, Timotoy B. Werner. 1990. Conserving the World’s Biological Diversity. World Bank Publications, P.O. 7247-8619, Philadelphia, PA 19170-8619. Order Stock £11384. $14.95 paper (8.5 x 11) 174 pp. This book is published by the World Bank, The World Resources Institute, the Interna- tional Union for Conservation of Nature and Natural Resources, Conservation Internation- al, and the World Wildlife Fund. 2 hapter 1. ee Diver rsity: What it is and Why it is Important (3 topics); Chapter The Values of Biological Diversity (5 topics); Chapter 3. How and Why Biological esources are eee - oe Cc hapter 4. Approaches to Conserving Biological Diversity (5 topics); Chapter 5. The Information Required to Conserve Biolowcal Diversity (S topics); C sees 6. Establishing aie for Conserving Biological pelos: (3 topics); Chapter 7. The Role of Strategies and Action Plans in Promoting Conservation of Biologi- cal Dives (4 topics); eine 8. How to Pay for Conserving Biological Diversity (4 topics); Chapter 9. Enlisting New Partners for aries of Biological Diversity (4 topics); Annex 1, C eee of Life on Earth by Phylum; Annex 2, The World Charter for Nature; Annex 3, International Legislation supporting Conservation of Biological Diversity; eae 4, The Bali Action Plan; Annex 5, The World Bank Wildlands Policy; Annex 6, Glossary. wfm Sipa 14(3):442. 1991. ANNOTATED CHECKLIST OF ARIZONA CONVOLVULACEAE DANIEL E AUSTIN' Department of Botany Arizona State University Tempe, AZ 85287, U.S.A. ABSTRACT Specimens examined in 14 herbaria indicate that there are 30 native and naturalized species of Convolvulaceae in the state of Arizona. Types and select specimens are cited Notes are given on county distributions, eee altitudes, and flowering dates. Com- ments are made on taxonomic problems, abundance and natural history of selected taxa. A lectotype for Ipomoea heterophylla Ortega is selected. RESUMEN Una revision de las muestras de 14 herbarios indica que existen 30 especies nativas y naturalizadas para el estado de Arizona. Se citan los tipos y los eyjemplares estudiados. Se nota la distribucion por los condades, el habitato, la distribucion altitudinal, y las fechas de floracion. Se commenta sobre los problemas taxonémicos, la historia natural, y la abun- dancia de los taxa. Se selecciona un lectotipo para Ipomoea heterophylla Ortega. During preparation of the family Convolvulaceae for the Vascular Plants of Arizona, certain taxonomic notes were found necessary (Austin 1990). The following annotated checklist, which includes all correct names of species known for the state, provides notes on several aspects of sys- tematics, abundance and natural history of Arizonan morning glories. In some cases notes are given on the living colors of floral parts since these details are normally not part of floristic surveys. Included are the corresponding names from Kearney and Peebles (1951) and Kearney et al. (1960) and some additional synonyms. Where types have not been seen, the usage is based on other authors who have seen authentic material, ex- cept in a few cases where it is based on the protologue. 1. CALYSTEGIA lL. CaLysreGiaA LONGrPES (S. Watson) Brummitt, Ann. Missourt Bot. Gard. 52:215. 1965. — Type: NEVADA: in 1872, Wheeler s.n. (US!). Con- volvulus longipes S. Watson, Amer. Naturalist 7:302. 1873 ‘Permanent address: Department of Biological Sciences, Florida Atlantic University, Boca Raton, FL 2242 > S.A. 33431, Spa 14(3):443 — 457. 1991. 444 cin linearilobus Eastw., Proc. Calif. Acad. Sci. Ser. 4, 20:470. 1931. — Type: ZONA: Mazatzal Mi. Eastwood 17264 (CAS!; photo of K specimen at NY!). Known from Coconino, Gila, Maricopa, Mohave and Yavapai cos. The Species grows in chaparral although it has been found extending into the Upper Sonoran zone; 609 — 1706 m; flowering March to October. This rare species, having been collected few times since the 1930s ee 1940s, is endemic to the Southwest, from California (San Diego Co. Inyo Co. and from Kern Co. to San Luis Obispo Co.), s Nevada (Clark Co. "i Utah (Washington Co.) and Arizona. The species appears to be rare throughout its range. Corollas are white with a cream-colored throat, and have pink patches on the limb around the nectar guides (interplicae). Styles, androecia and gynoecia are white, but the nectary is yellow. Flowers are pollinated by Osmia sp. (Megachilidae). 2. CALYSTEGIA SEPIUM (L.) R. Br. ssp. ANGULATA Brummitt, Kew Bull. 35(2):328. 1980. — Tyre: IDAHO. Canyon Co.: Machride 318 (NY!'). Calystegia sepium (L.) R. Br. var. angulata (Brummitt) N. Holmgren in A. Cronquist et al., Intermountain Fl. Vasc. Pl. Intermountain West, U.S.A. 4:77. 1984. Known from a single specimen collected in 1882 from a garden in Cochise Co.; ca 1524 m; flowering in June. These olants are easily confused with the Great Plains taxon C. sylvatica ¢Kit.) Griseb. ssp. fraterniflora (Mackenzie and Bush) Brummitt as shown by the interpretations in Tryon (1939), Correll and Correll (1972) and Lehr (1978). The quadrate sinuses in the leaf bases of C. sy/vatica allow separa- tion from the V-shaped or U-shaped sinuses of C. sepium. 4. CALYSTEGIA MACOUNH (Greene) Brummitt, Ann. Missouri Bot. Gard. 352:215. 1965. — Type: CANADA. SaskatTCHEWAN: Assiniboia, Milk River, ae 1905, Macoun 11883 (not seen). Convolvulus macounti Greene, Pittonia 3:326. Convolvulus sepium sensu Kearney and Peebles Con volv a mterior House, Bull. Torrey Ban Club 32:140. 1905. — Typr: LORADO: near Fort tee Crandall 1625 (NY', US!). Known from Apache, Coconino, Navajo and Yavapai cos. Growing in moist sites, near lakes and streams; 1950 — 2042 m; flowering June to July. This rare species is a Great Plains endemic that had not been collected in Arizona since 1971 until it was relocated in 1990 (Austin & Austin 7661, A Corollas, stamens and gynoecia are white, although the nectary is yellow. Bees (not yet determined) visit the flowers. 445 2. CONVOLVULUS 1. CONVOLVULUS ARVENSIS Linnaeus, Sp. Pl. 153. 1753. — Tyee: SWEDEN: specimen 218.1 (LINN, microfiche!). This introduced European weed was first collected in Arizona in 1843 near the Navajo Ordinance Depot in Flagstaff by Lt. P. Shallert, an army surgeon. A short time later, in 1882, Lemmon collected it in the Huachuca Mountains near Ft. Huachuca. The species is now known from all Arizona counties except La Paz. A collection from Greenlee Co. (Austin & Austin 7634, ASU) is a new county record. Common in disturbed sites, roadsides, and cultivated fields; 341 — 2346 m; flowering April to October. Common name: “BINDWEED.” 2. CONVOLVULUS EQUITANS Benth., PI. oe 16. 1839. — Tree: MEXICO: 1837, Hartweg 98 (oresumpably & C. incanus sensu auctt., non Vahl. Known from all Arizona counties except Yuma and La Paz. Frequent in grasslands and plains; 762 — 1981 m; flowering March to November. Local common names are: “Si-veR BINDWEED” (Little 37, ARIZ), and “Desert BINDWEED” (Wilcox 5.n., ARIZ). The filaments are sae the anthers purple. Styles and the bases of the stigma lobes are white, but the tips of stigma lobes are purple. The nectary is yellow and cup-shaped. Visited by honeybees (Apis mellifera) and bumblebees (Bombus sp.) (Pima Co., Austin G Austin 7561, ASU). 4, CRESSA 1. CressA TRUXxILLENSIS H.B.K., Nov. Gen. Sp. Pl. 3:93. 1819. — Tyee: PERU: Trujillo, Humboldt & Bonpland 3727 (B: microfiche!, IsoTyPE: F!). C. depressa Goodd., Bot. Gaz. 37:58. 1904. — Type: NEVADA: Goodding 726 (UC!). C. insularis House, Bull. Torrey ae es 33:315. 1906. — Type: MEXICO: Revilla- gigedo Isls, Barkelew 252 (U C. erecta Rydb., Bull. Torrey is on 40: 466. 1913. — Type: UTAH: Garrett 870 (NY!). C. minima Heller, Muhlenbergia 8:140. 1913. — Type: NEVADA: Heller & pee 8663a (NY!). C. truxillensis H.B.K. var. minima (Heller) Munz, Aliso 4:96. 1958 C. pumila Heller, Muhlenbergia 8:142. tab. 17. 1913. nomen nudum C. vallicola Heller, Muhlenbergia 8:140. tab. 17. 1913. — Tyre: CALIFORNIA: Hel- ler 8936a UC!). C. truxillensis H.B.K. var. vallicola (Heller) Munz, Aliso 4:96. 1958. C. cretica L. var. truxillensis GH.B.K.) Choisy in DeCandolle, Prodr. 9:440. 1845. Known from Coconino, Maricopa, Mohave, Navajo, Pinal and Yuma cos. Occasional in saline desert, 30-1524 m, flowering May to November. 446 Plants were last collected in Mohave Co. in 1912 (Jepson 5026, ARIZ); in Pinal Co. in 1937 (Peebles 13233, ARIZ); in Maricopa Co. in 1964 (Rea s.n., ASU); and in Navajo Co. in 1978 (Pimkava et al. 13839 ASU). Although last collected in Yuma Co. in 1971 (Hamilton s.n., ARIZ, ASU, DES), the plants were relocated in 1989 (Austin & Austin 7586, ASU). Previously the plants had been separated into species or varieties on the basis of growing erect or prostrate. Both growth forms were growing side by side in Yuma. Plants were not found in flower. 4. DICHONDRA 1. DicHONDRA ARGENTEA Willd., Hort. Berol. 297. tt. 81. 1806. — Type: COLOMBIA: Tolima ca. Honda, Bonpland (B?). Known from a single collection from Bisbee in Cochise Co. in 1931 (Harrison 8256, ARIZ). In New Mexico the plants grow in Chihuahuan desert scrub and Madrean oak woodlands; ca. 1615 m; flowering in September, earlier in New Mexico and Mexico. The species is frequent in Texas and New Mexico as far west as the Organ Mts. Perhaps the Arizona collection represented a western limit to the natural range of the species, or perhaps it was introduced into Bisbee dur- ing the mining operations. Plants have not been relocated in Arizona even though several people have searched for them. 2. DICHONDRA BRACHYPODA Wooton & Standley, Contr. U.S. Natl. Herb. 16:160. 1913. — Tver: NEW MEXICO. Dona Ana Co.: Organ Mountains, Filmore Canyon, 23 Sep 1906, Wooton & Standley s.n. (US!). Known from Cochise and Santa Cruz cos. Occasional in Madrean oak woodlands and lower ponderosa pine zones; 1219— 1889 m, flowering July to October. Although the morphological traits given by Tharp & Johnston (1961) are sufficient to distinguish species, they also differ by colors of flower parts. The corolla lobes are cream, green at base. Filaments and stigmas are green. Anthers are white witha purple stripe and the nectary is dark green. The ovary 1s light green and the styles are cream. Mature fruits collected on 9 Sep 1989 (Cochise Co. Rucker Canyon. Austin & Austin 7611, ASU). Fruits were fully mature by October. 3. DICHONDRA SERICEA Swartz, Prodr. Veg. Ind. Occ. 54. 1788. — Tyee: AICA: Swartz (S?). D. repens Forster & Forster var. sericea (Swartz) Choisy in DeCandolle, Prodr. 9:451. 1845. D. repens sensu auctt., non Forster & Forster. In Arizona the species 1s known exclusively from the Pajarito Mts. in 447 Santa Cruz Co. Rare in streamside vegetation; ca. 1112 m; flowering May to December. The single Arizona location is Sycamore Canyon (Santa Cruz Co. from 1936, Goodding 6620 ARIZ. to 1962, Barr 62-863 ARIZ) near the Mexican border. This population was relocated after an hiatus of 19 years (Austin & Austin 7604, ASU), but its size has apparently declined. Based on herbarium specimens flowers and fruits are uncommon in this population. Flowers were found in April. Petals, anthers and gynoecia are pale green (Austin G Austin 7604, ASU). No flower visitors were seen. de EVOLVULUS 1. EvorvuLtus arstnompes L. var. ANGUSTIFOLIA Torrey, Bot. Mex. Bound. 150. 1858. — Tver: TEXAS. Presipio Co.: ca. the “Grand Canyon” of the Rio Grande. August, Parry (not found in CM, GH, ISC, MO, NY, PH, US or YU). E. alsinoides 7 var. goa (Willd. ea Meded. Bot. Mus. Herb. Rijks Univ. 4:34. 19 — Type: CO. GuerRRERO: ca. Acapulco, Willdenow 6128 ne Known from Cochise, Pima, Pinal and Santa Cruz cos., and reportedly in Gila and Maricopa (Kearney and Peebles 1951, 1960). Occasional in pine-oak woodlands, saguaro desert scrub, and desert grasslands; 762 — 1828 m; flowering February to October. The common name: “Dio bE viBoRA” (Lehr 1978) must represent an error in spelling, or an ortho- graphic variant either of “Tio br viBorA” or “Dios bE viBora,’ although I have found neither of these common names to be in use. Flowers open 8:00-9:00 a.m., and close ca. 4:00-5:00 p.m. (Pima Co.., Austin & Austin 7598, ASU). Corollas are blue on the limb, white in the throat. Stamens and gynoecia are white. No flower visitors were seen. 2. EVOLVULUS ARIZONICUS - Cray, Syn: Flo IN« Amen. 2, 1218. 1878. ypE: MEXICO: sandy prairies, a 1857, hater 1023 (GH!). See Austin to) on peiaes %E typificat Evolvnlus laetus A. Gray, pe Amer. Acad. Arts 17:228. 1882. — Type: ARIZONA: 1881, Pringle s.n. (F!, GH!, US!). E. arizonicus A. ronee > ae A. Gray) Ooststr., Meded. Bot. Mus. as pan Univ. Utrecht 14:76. 193: Known from Cochise, Gila, Graham, Greenlee, Mohave, Navajo, Pinal, Pima, Santa Cruz and Yavapai cos. Occasional to frequent in dif- ferent parts of the state; more common in the southern counties. Plants grow in chapparal, Madrean oak woodlands, and mesquite grasslands; 883 — 1828 m; oe April to October. Common name: “FALSE FLAX” (McGinnies 4, ARIZ Flowers, which open 7:00-8:00 a.m. and close 3:00-4:00 p.m., have 448 blue limbs, white throats, and white stamens and gynoecia, and are visited by bees (not caught but possibly Halictidae). Pollen was gone and some of the anthers were stripped from flowers by 11:30 a.m. (Austin & Austin 7588, U) 3. EvoLvuLUS NUTTALLIANUS Roem. & Schult., Syst. Veg. 6:198. 1820. — Type: on the banks of the Missouri, Nwvtta// (B?). E. id Nutt., Gen. N. Amer. Pl. 1:174. 1818, nom. superfl. — Type: on the banks Missouri, Nuttall (B?). E. sreopbilus Greene, Leafl. Bot. Observ. os 1:151. 1905. — Type: NEW MEXICO: Metcalfe 1228 (NMC!, NY!, UC!, US). Known from Apache, Cochise, Coconino, Gila, Maricopa, Mohave, Navajo, Pima, Pinal, Santa Cruz and Yavapai cos. Occasional in chaparral, Madrean oak woodlands, ponderosa pine zone, pinon-juniper zone, and rocky grasslands; 822 — 2438 m; flowering April to September. Some herbarium specimens are difficult to separate from E. sericeus. The species usually may be separated by the spreading-villose trichomes on the sepals of E. nattallianus (Santa Cruz Co., Austin G Austin 7572, ASU) and appressed-pilose trichomes on E. sericews although there are intermediate specimens (Metcalfe 1228, NMC, NY, US). corolla limbs are pale blue changing to white near the base; the short tube is pale yellow within. Androecia and gynoecia are white. No insect visitors were seen. 4, EVOLVULUS SERICEUS — Prodr. Veg. Ind. Occ. 55. 1788. — Tyee: JAMAICA: Swartz (M, E. wilcoxianus House, Bull. _ Bot. Club 33:315. 1906. — Type: ARIZONA: Wilcox 96 (US!). Known from Cochise, Gila, Graham, Greenlee, Pima, Pinal, Navajo, Santa Cruz and Yavapai cos. Frequent in chaparral, Madrean oak wood- lands, and desert grasslands; 975 — 1889 m, flowering May to October. For those wishing to recognize them, two varieties have been named: var. discolor (Benth.) A. Gray, with upper leaf surface glabrous and var. sericeus, with leaves sericeous on both surfaces. Since these plants may be found growing together outside the U.S.A., I do not recognize the distinc- tion. Specimens that were separated as E. oreophilus Greene were treated by Ooststroom (1934) as E. sericeus var. discolor form B. These are better treated as E. nuttallianus because of their habit, corolla shape and color and in- dumentum on both leaf surfaces. The androecium and the gynoecium are white (Cochise Co. Austin G Austin 7571, ASU). No flower visitors have been seen. 449 6. IPOMOEA te sar BARBATISEPALA A. Gray, Syn. Fl. N. Amer. 2, 1:212. 1886. ae : TEXAS: Wright 507 (GH!, US!). Known from Cochise, Gila, Graham, Greenlee, Maricopa, Pima, Santa Cruz, and Yavapai cos. Occasional in mesquite grasslands and Madrean oak woodlands; 853 — 2438 m; flowering July to December. Flowers open at dawn, and are at first blue on the limb with a white throat. The outside of the tube is white on the folds (plicae) and pale green on the unfolded area (interplicae). As senescence beings, the corolla turns pink and then reddish. Anthers are white, but the filaments are pale yellow. The style is green, the stigma white and 2-lobed, the ovary green, and the disc yellow and cup-shaped (Pima Co., Austin & Austin 7594, ASU). 2. IPOMOEA CaPILLACEA (H.B.K.) G. Don, Gen. Syst. 4:267. 1838. — Type: COLOMBIA: Bonpland (microfiche!). 1. muricata Cav., Icones Pl. 5;52. pl. 478. f. 2. 1794, non L. (1763), non Jacq. (1789). Known from Cochise, Coconino, Pima, Santa Cruz and Yavapai cos. Occasional in Madrean oak woodlands, desert grasslands, and ponderosa pine zones; 1524 — 2499 m; flowering July to September. Flowers open at dawn. The corolla limb is lavender and the tube white, within and without. Androecia and gynoecia are also white. Beeflies (Bombyliidae) visit the flowers (Cochise Co. , Austin & Austin 7569, ASU). 3. [POMOEA CARDIOPHYLLA A. Gray, Syn. Fl. N. Amer. 2, 1:213. 1886. — Tree: TEXAS: Wright 511 (GH!). No Arizona specimens were seen by Kearney and Peebles (1951, 1960) although they suggested this species’ presence in Arizona as probable. A population was reported near Tombstone by Mason et al. (1986) and was relocated in 1989 (Cochise Co., Austin & Austin 7608, ASU). The species is represented by scattered plants extending along the road for about 0.8 miles at an altitude of 1127 m in mesquite-creosote bush scrub (Chihu- ahuan desert scrub sensu Brown and Lowe 1980). No fruits were present on 7 Sep 1989; some mature fruits were found on 26 Sep, but were heavily parasitized by insects. Rare and local. Flowers open at 6 a.m.; mostly wilted by 11 a.m. The corolla tube is pale yellow on the outside; the limb ts pale blue (as in J. tricolor). Stamens are white to pale cream; the style is green, the stigma white. No odor was detectable, and no insects visited between 6:00 and 6:30 a.m. McDonald (1982) found no pollinators on the species in Texas and New Mexico and 450 had a 90% seed set on cultivated plants. The species may be considered autogamous. es A COSTELLATA Torr., oe Mex. Bound. 149. 1859. — Tver: XAS: Wright SOS (GH, Us! 1. futilis A. Nelson, Univ. Wyoming Publ. Sci. 1(3):65. 1924. — Type: ARIZONA: Hanson 1016 (RS, not seen; photo FAU!). Known from Apache, Cochise, Coconino, Gila, Greenlee, Mohave, Navajo, Pima, Pinal, Santa Cruz and Yavapai cos. Common in chaparral, Madrean oak woodlands, and ponderosa pine zones; 975 — 2133 m; Hower- ing July to October. Flowers begin to open at 8:15 a.m. and begin to wilt at 10:30 a.m. The corolla limb is lavender, the tube white within and without. Filaments and anthers are white; white trichomes extend along the filaments from base to apex; the ovary is cream, and the style is green. A stigma was 3-lobed on one flower, 2-lobed on others. The flowers are visited by bumblebees (Cochise Co. Austin & Austin 7584, ASU). 9. [pomora crisruLATA H. Hallier, Med. Rijksherb. Leiden 46:20. 1922. Anew name for Qvamoclit gracilis H. Hallier. — Tyee: MEXICO: based on syntypes including Bourgean 1061 (G-DC!). Quamoclit gracilis H. Hallier, Bull. Herb. Boiss. 7:416. 1899 I. coccinea auctt., non L. Known from all Arizona counties except La Paz. Plants grow in chaparral, Madrean oak woodlands, and ponderosa pine zones; 73 1 — 2773 n; flowering May to November. This is probably the most common and widespread species in the state. “Srar Glory” (Blakeley & Marshall 568 Flowers, which are open from 6:00 a.m. into the late afternoon, are visi- ted by hummingbirds. Since the stigma is held below the level of stamens, the plants may be facultatively allogamous (Pima Co., Austin & Austin 7593, ASU). According to an anonymous reviewer the species is probably predominantly selfing and there may be cleistogamous flowers under low light conditions. Even when chasmogamous the anthers dehisce before anthesis. 7. IPOMOEA HEDERACEA Jacq., Collect. Bot. 1:124. 1786. — Tver: based on Dillentus, Hore. Elth. t. 80, fig. 92 (plate selected lectotype! by Verdcourt, 1957) . desertorum House, Ann. New York Acad. Sci. 18:203. 1908. — Type: ARIZONA: Thornber 29 (ARIZ!, NY!). Although not reported for Arizona by Kearney and Peebles (1951, 451 1960), certain specimens identified as “/. Azrsutula” are of this species, the others are 1. purpurea (Austin, 1990). Known from Cochise, Coconino, Gila, Graham, Maricopa, Pima, Pinal, Santa Cruz, Yavapai and Yuma cos. Common in various disturbed sites, especially cotton fields; 883— 1859 m; flowering August to November. The corollas open at dawn, some closing by 8:15 a.m., and all are closed 11 a.m. Honeybees were seen bypassing flowers after pausing. A bumblebee bypassed flowers of 1. bederacea but visited those of 1. costellata and Convolvulus equitans. However, flowers of I. hederacea were visited by short-tailed black swallowtail butterflies (Papilio indra). Corolla limbs are blue early after opening but begin to include reddish pigments as they begin to wilt and turn more purple. The corolla tube is white within and without. The stamens and the styles are white; the ovary 1s cream (Pima Co., Austin & Austin 7596, ASU). 8. [IPOMOEA LEPTOTOMA Torr., Bot. Mex. Bound. 150. 1859. — Tyrer: MEXICO. Sonora: Thurber 977 (GH!). 1. leptotoma var. wootontt E. Kelso, Rhodora 39:151. 1937. — Tyree: ARIZONA: 10 Sep 1914, Wooton 10 (US!). < emis Torr. f. wootonm (E. Kelso) Wiggins, Contr. Dudley Herb. 4:21. 195¢ Known from Cochise, Gila, Graham, Pima, Pinal, Santa Cruz and Yavapai cos. Occasional in plains, Sonoran desert scrub; 609 — 1371 m,; flowering June to October. The corolla opens as the sun rays touch it, between 7:30 and 8:00 a.m., and close between 10:30 and 11:00 a.m. Corolla limbs are lavender, and there is a white zone between the limb and pale yellow base of the tube. The tube is white without. Orange trichomes adorn filaments from base to apex, the androecium, gynoecium, style, ovary, and disc are white. Skippers (Hesperiidae) visit flowers (Gila Co., Austin G Austin 7001, ASU, which ts a new county record; Pima Co. Austin & Austin 7595, ASU). 9. IPOMOEA X LEUCANTHA Jacquin, Icones Rar. 2:t. 318. 1788. — Type: no specimen known; illustration chosen as lectotype by Austin in 1978. . trtloba sensu auctt., non L. (1753). : lacunosa sensu Shines Pee non L. (1753). Known from Maricopa, Pima and Yuma cos. Plants grow in disturbed sites; ca. 701 m; flowering March to Novembe Known from three old collections (Santa Cruz Co., Pringle in 1884, ARIZ; Pima Co., Thornber in 1912, ARIZ; county unknown, LeRoy s.n., NY); one in 1945 (Pima Co. Goodding & Lusher 128-45, NY), and two recent ones (Maricopa Co. 4 Oct 1979, Heathman s.n., ARIZ, ASU; Yuma 452 Co. 7 Nov 1985, Tuttle s.n., ARIZ). The hybrids are probably not as rare as collections seem to indicate since they are weeds in cotton fields. 10. [POMOEA LINDHEIMERI A. Gray, Syn. Fl. N. Amer. 2, 1:210. 1886. — Type: TEXAS: Wright 508 (GH!, US!). Known from Cochise Co. (Gleeson, 25 Aug 1927, Thornber s.n., ARIZ; Bisbee, 30 Sep. 1930, Thornber s.n., ARIZ); reportedly from Pima Co. (Kearney and Peebles 1951; Kearney et al. 1960). Plants grow in Madrean oak woodlands, and Chihuahuan desert scrub zones; 1066— 1371 m; flowering August to September. An extremely rare species; its continued existence in Arizona is proble- matical. Probably related to and easily confused with I. pubescens Lam., the two collections from Arizona are somewhat intermediate between the two on the basis of sepal shape and pubescence. Ll. [pomMora LONGIFOLIA Benth., Pl. Hartweg. 16. 1839. — Tver: MEXICO: Hartweg (K). Known from Cochise and Santa Cruz cos. Locally common in Madrean oak woodlands; 975 — 1828 m; flowering July to September. The nectary is cream-colored, and the androecium and gynoecium are white. Early in the evening the flowers have a slight sweet fragrance. Flowers are pollinated by moths (Austin 1986). Flowers open 3:00 to 4:00 p.m. and close near dawn; a few were still open at 7:30 a.m. (Cochise Co.., Austin & Austin 7582, ASU). 12. IPOMOEA PLUMMERAE A. Gray, Syn. Fl. N. Amer. 2, L:suppl. 434. 1886. — Type: ARIZONA: Lemmon 2839 (GH)). !. cuneifolia A. Gray, Proc. Amer. Acad. Arts 19:90. 1883, non Meisner (1869). — Type: ARIZONA: Lemmon 2839 (F!, GH!, US!). I. egregia House, Torreya 6:124. 1906, nom. nov. for 1. cwnetfolia A. Gray. Known from Apache, Cochise, Coconino, Gila, Graham, Greenlee, Pima, Santa Cruz and Yavapai cos. Occasional in the ponderosa pine zone; 1219 — 2743 m; flowering April to October. The corollas open by ca. 6:30 a.m., are closing at 10:30 a.m. and are completely closed at 11:50 a.m. Corolla limbs are lavender, the tube white within and without. The ovary and style are green, the stigma and stamens white (Cochise Co., Austin G Austin 7581, ASU) 13. IpOMorA PUBESCENS Lam., Encycl. Meth. Bot. 1:265. 1791. — Typr: AMERICA: collector unknown (K!). I. heterophylla Ortega, Nov. Pl. Descr. Dec. 1:9. 1797. — Typr: MEXICO: Horto egio., 1797, Ortega lee TOTYPE: MA 222592, photo FAU!). 453 The curator of the herbarium in Madrid sent me photographs of six specimens of ses heterophylla in their collection. One of these was collected in Peru (MA 2596) and will not serve as a lectotype because the protologue cites only Cuba and ve Two other sheets (MA 222595, 222597) are labeled with the dates 1798 and 1801, respectively. These will not serve as lectotypes because they are dated after the publication by Ortega. Two of the remaining sheets (MA 222593, 222594) were labeled by J.D. Rodrigues, and are excluded because they are incomplete in ae to the last collection which carries Ortega’s name. Therefore, the speci- ith Ortega’s name as collector is chosen as lectotype. I. indir A. Gray var. ee House, Ann. New York Acad. Sci. 18:196. . — Tyree: ARIZONA: Lemmon 2835 (GH!). Known from Cochise and Santa Cruz cos. Growing in canyons, 106 — 1371 m,; flowering August to September. This is now an extremely rare plant in Arizona. Of the seven sites where it was formerly collected, Bee were relocated in only one (Santa Cruz Co., Austin G Austin 7005, ). The plants are not common in adjacent Mee and should be setae endangered in the state of Arizona. Plants in Texas, New Mexico and Arizona have flowers considerably larger than the populations in Meso-America and South America. Perhaps more than one taxon is involved. Flowers begin to close at 9:40 a.m., and all are closed by 10:15 a.m. Corolla limbs are lavender, and the tubes are white within and without. Styles, stigmas and stamens are white. 14. IPOMOEA PURPUREA (L.) Roth, Bot. Abh. 27. 1787. — Tver: U.S.A.: Dillenius, Hort. Elth. t. 84, fig. 97. 1732 (chosen lectotype! by Verdcourt, 1963). a ict reo , Eclog. Pl. Rar. 1:63. t. 44. 1811. — Type: no specimen found, the plate chosen as lectotype by Austin (1990). Known from Apache, Cochise, Gila, Graham, Greenlee, Maricopa, Mohave, Navajo, Pima, Santa Cruz and Yavapai cos. Found in cultivated fields and other disturbed sites; 304—2286 m; flowering July to November. Occasional to common in counties bordering Mexico. Flowers in cultivated plants have variable corolla colors, but the wild populations are consistently purple on the limb, with pink nectar guides (plicae), and tubes which are white within and without. In wild plants, the ovary is green, and the androecium, style and stigma are white; in culti- vated plants, the ovary ts cream, the style white. Flowers (wild plants) are visited by sulphur butterflies (Cochise Co., Austin & Austin 7014, ASU). tee pees TENUILOBA Torr., Bot. Mex. Bound. 148. 1859. — Tyee: XAS: Bigelow (US!). I, lemmoni A. oy, Proc. Amer. Acad. Arts 19:20. 1883. — Type: ARIZONA: Lemmon 2 , US!). 1. tenutloba Torr. var. jee (A. Gray) Yatskievych & Mason, Weise i: 102. 1984. 454 Known from Cochise, Pima and Santa Cruz cos. Plants grow in chaparral, Madrean oak woodlands, and ponderosa pine zones; 1280 — 1920 m; flowering August to September. A rare species that should be considered threatened in Arizona. Two varieties exist (Yatskievych and Mason 1984): J. tenwiloba var. tenuiloba has white flowers; /. tenuiloba var. lenmoni (A. Gray) Yatskievych and Mason has purple flowers. Flowers of var. /emmoni open before daylight between 1:00 and 5:00 a.m. and close between 7:00 and 8:00 a.m. Corolla limbs are pale lavender and the tube is white within and without. Stamens and stigmas are white. Although the morphology of the flowers indicates adaptation for moth pollination, no moth scales on the stigmas or other evidence of visitation was found. The flowers had no fragrance. Perhaps the plants in Arizona, being on the northern fringe of the range, are autogamous. Plants had been in flower for 10 days by 29 Aug 1989, and fruits were about half grown on a few plants. About 30 flowers were found in the population on 29 Aug; 24 flowers on 30 Aug. The population was rechecked in Bear Canyon 14 Sep and was still alive - no longer in flower (Pima Co., Bear Canyon, Austin & Austin 7592, ASU). Plants grow in the Pinus-Juniperus-Quercus zone in oe Catalina Mts. and Huachuca Mts. Found on quartzite in the Huachuca Mts. (Santa Cruz Co., Austin & Austin 7618, ASU). This substrate is the only one where the species occurs in the Huachuca Mts. (EF Reichenbacher, pers. comm., 1989) 16. Ipomoea THURBERI A. Gray, Syn. Fl. N. Amer. 2, 1:212. 1886. — Tyrer: ARIZONA: Thurber 966 (HOLOTYPE: GH!). I. gentry: Standley, Field Mus. Nat. Hise. 22:46. 1940. — Typr: MEXICO. Cuinu- iA: Rio Mayo, Sierra Canelo. 30 Aug 1936, Gentry 2497 (HOLOTYPE: F! ~— Known from Cochise, Pima and Santa Cruz cos. Grows in Madrean oak woodlands, near lakes; 1158— 1524 m; flowering July to September. For some time the species was thought to be endemic to the United States. Although not included under J. thurberi for Mexico by Matuda (1963-1965), he did include it from Chihuahua, Durango and Sonora under J, awe ME SONORA: vic. El Llano, ca. 9.5 mi W of San Felipe, Sierra Los Locos, 11-12 Aug a a & Martin s.n. (AR Kearney and Peebles (1951) wrote that the plants had “purple flowers opening in the evening.” In fact, the flowers have a pink limb and green throat; they wilt and dry with a green tube and purple limb. Flowers, opening near 6:30 p.m., are visited by sphinx moths (probably Hydes 455 lineata). All flowers examined had moth scales on the stigmas, further indicating moth pollination. Gynoecia and androecia are white. Only 30—50 plants comprise the population (Santa Cruz Co., Austin G Austin 7603, ASU). The species is rare in Arizona and in Mexico (J. A. McDonald, personal communication, Nov. 1989), and should be placed on Arizona's endangered list. Tentatively placed in Ipomoea section Tyrianthinae by McDonald (1987), the species does not belong to that section because it has three carpels. The species belongs to Ipomoea section Pharbitis where it was originally placed by A. Gray. 7. JACQUEMONTIA 1. JACQUEMONTIA AGRESTIS (Choisy) Meisn. in Mart., Fl. Bras. 7:306. 1869. — Tyee: BRAZIL: Martins (M!, photo MO!). Convolvulus agrestis Choisy in DC., Prodr. 9:405. 1845. J. palmert S. Watson, Proc. coe Acad. Arts 24:63. 1889. — Tyrpr: MEXICO: suaymas, Brandegee s.n. (GH , US!). In Arizona, known only from Pima Co. Plants probably grow in semidesert grassland; ca. 1219 m; flowering August to October. This ian was collected in the Baboquivari Mts. several times between the 1920s and 1940s, but it has not been collected since. Although the species is associated with cultivated land in many places in Mexico, its current status in Arizona is uncertain. It may have been brought into the state from Mexico as a weed with plants cultivated by the Tohono O’odham. In Mexico and elsewhere the species is commonly a weed in maize fields and other cultivated crops. 2. JACQUEMONTIA PRINGLE! A. Gray, Proc. Amer. Acad. Arts 17: 228. 2. — Tyrer: ARIZONA: Pringle 295 (GH!). J. pringler var. glabrescens A. Gray, Proc. Amer. Acad. Arts 21:402. 1886. — Tyrer: XICO: Palmer 107 (GH! chosen lectotype by Robertson (1971), but not published; his choice here upheld). Known from Pima, Pinal, Yuma and doubtfully recorded Cochise Co. Frequent in saguaro desert scrub; 914— 1371 m,; flowering April to October There is a specimen supposedly collected in the Chiricahua Mts. (Cochise Co. Chiricahua Mountains, 20 Jul 1895, Toumey s.n. NY!, US!). Since no one else has located the species in this mountain range, nor in the nearby ranges, the specimen probably was incorrectly labeled. The plants are Sonoran Desert endemics ranging from Pima Co., Arizona, south to Los Mochis, Sinaloa, Mexico (Robertson 1971). 456 Flowers, which are white throughout, open at dawn, as the sun strikes them, and close between 3:00-4:00 p.m. Pollination is by the bee Duforea sp. (Halictidae) which drinks nectar but does not actively collect pollen. Numerous bees visit the flowers regularly. Later in the season, fruit set is high, with most flowers producing some seed (Pima Co., Austin & Austin Foo), The following species are now or have been in cultivation in Arizona: Convolvulus cneorum L. (Morning Glory, Bush Morning Glory), Convolvulus tricolor L. (Morning Glory), Convolvulus sabatius Viviani var. mauritanicus (Boiss.) Sa’ad (Morocco Glorybind, Ground Morning Glory), Dichondra micrantha Urban (Pennywort), Ipomoea batatas (L.) Lam. (Sweet Potato, Batata, Camote), Ipomoea carnea Jacq. ssp. fistulosa (Choisy) D. Austin (Tree Morning Glory, Bush Morning Glory), Ipomoea tricolor Cav. (Morning Glory, Heavenly Blue), and Merremia dissecta (Jacq.) H. Hallier (Alamo vine, Mile-a-minute vine). ACKNOWLEDGMENTS Thanks are extended to curators of herbaria(A, ASC, ARIZ, ASU, CAS, DES, GH, MNA, NMC, NY, TEX, UC, UNM, US) for the opportunity to study specimens. Dr. M. Cazier identified the bees. Drs. C. T. Mason, Jr. and D. Pinkava offered suggestions on the original manuscript. My wife, Sandra, helped with the field study and offered suggestions on the manuscript. This study was conducted while the author was on sabbatical leave at Arizona State University. REFERENCES AUSTIN, D. E 1986. Moth pollinated Ipomoea longifolia. Desert Plants 8(1):15 — 16. —_________.. 1990. Comments on southwestern United States Evolvulus L. and Ipomoea L. casei). Madrono 37(2):124 — 132. BROWN, D. E. and C. H. LOWE. 1980. Biotic gone of the Southwest, General Technical Report RM-78, U.S.D.A., Washington, D. CORRELL, D. S. and H. B. CORRELL. 1972. sani acd stra ae of south- western United States. Environmental Protection Agency, Washin ,D. KEARNEY, T. H. and R. H. PEEBLES. 1951. Arizona Flora. Cai. Calif aoe Berkeley. KEARNEY, T. H., R. H. PEEBLES, and collaborators. 1960. Arizona Flora. Ed. 2, with Supplement bi T. Howell, E. McClintock and collaborators, Univ. Calif. na Berkele LEHR, J. H. 1978. ete of the flora of Arizona, Desert Botanical ion: eras MASON, C. T., Jr, R. K. VAN aera ues D. STARR. 1986. M of Arizona VII. ee Plants 8(1):38 — MATUDA, E. 1963— 1965. El genero sie en Mexico I — II. Anales Inst. Biol. 34:85 — 145. 1963; 35:45 —76. 1964; 36:83 — 106. 1965. MCDONALD, J. A. 1982. Biosystematics of the Ipomoea tricolor complex (Convolvu- laceae). Ph.D. dissertation, Univ. Texas, Austin. 457 1987. Revision of Ipomoea section Exogonium (Choisy) Griseb. (Convolvu- laceae). Hise 28:41 —87. pe ian arta S. J. VAN. 1934. A monograph of the genus Evolvulus. Meded. Bot. us. Herb. Rijsk Univ. Utrecht 14. eGo Ga K. R. 1971. A revision of the genus ea aa (Convolvulaceae) in as - Pan America and the West Indies. Ph.D. dissertation, Washington ouis, Misso SHINKERS - H. 1965. ies lacunosa (Convolvulaceae) in Arizona. Leaflets W. Bot. 10(10): 162. THARP, B. C. and M. C. JOHNSTON. 1961. Recharacterization of Dichondra (Convolvu- laceae) and a revision of the North American species. Brittonia 13(4):346 — 360. TRYON, R. 7 - 1939. The varieties of Convolvulus es and of C. sepium. Rhodora 41:415— YATSKIEVYCH, G. and C. T. MASON, JR. 1984. A taxonomic study sities tenutloba Torr. (Convolvulaceae), with notes on related species. Madrofo 3 1(2):102 — 108 BOOK REVIEWS Goubey, CHRISTOPHER J. 1988. A Handbook of Ferns for Australia and New Zealand. Lothian Publishing Company. Order from: In- ternational Specialized Book Services, Inc., 5602 NE Hassalo Street, Portland, OR 97213-3640. $19.95 paper. 212 pp. This handbook of ferns is concerned with the horticultural aspect as opposed to an idencification manual. There are 3 chapters in Part 1. How to Identify Ferns; 5 chapters in Part 2. Propagation of Ferns; 2 chapters in Part 3. Pests and Diseases; 2 chapters in Part 4 Ferns in Cultivation; 3 chapters in Part 5. Where to See Ferns; 6 chapters (concerning hardiness, indoor, hanging baskets, and terrariums) in Part 6. Plant Lists. At the end of the text 1s a glossary, list of fern societies, and index. There are diagrammatic illustrations in Part | but the taxa are mostly photographs, black and white with some colored plates. An excellent book for the fern horticulturist. wfm Copy, WM. J. anp Donatp M. Brirron. 1989. Ferns and Fern Allies of Canada. Canadian Government Publishing Centre, Supply and Services Canada, Ottawa, Canada KIA 0S9. $38.50 CAN; $46.20 US (Check to Receiver General for Canada). This manual includes keys, synonymy, descriptions, cycology, habitat, range, remarks, and diagrammatic illustrations. The distribution maps are clustered prior to the glossary, references, and index. “It is hoped that the book will prove to be a useful tool, not only to individuals raking a first look at these interesting plants but also to the dedicated amateur and the professional botanist.” And I believe it will be very useful to all three groups. wfm Brown, ANTHONY H.D., MicHart T. Crecc, Arex L. KAHLER, Bruce S. Weir. Editors. 1989. Plant Population Shae oe Breeding, and Genetic Resources. Sinauer Associates, Inc. Sunderland, MA 01375-0407. $36.00 paper; $60.00 cloth. 449 a, This book is based upon the International oe on Po} pene Genetics and Germplasm Resources in Crop Improvement, held August 11 — 13, 1988 at the University of California, Davis. The articles or chapters are ae under 3 sections: Section 1: Genetic ae ar and Amounts G bee Section 2: Evolutionary Processes (7 chapters); Sect 3: Applications in Plant Breeding and Genetic Resources (6 chapters). The symposium nee was to honor eee Robert W. Allard, who founded experimen- tal plant population genetics as a scientific discipline. Allard wrote the first chapter: Fucure Directions in Plant Population Genetics, Evolution, and Breeding. An excellent resource text with a compilation of the literature citations of the individual chapters at the end near the Index. wfm Sipa 14(3):458. LOOL. TWO NEW VITIS (VITACEAE) FROM MOUNTAINOUS MEXICO BARRY L. COMEAUX Galveston College 4015 Avenue Q Galveston, TX 77550, U.S.A. ABSTRACT Two new species, Vitis bloodworthiana and V. jaegertana, are described and eee to the two most similar species in series Occidentales. Vitis bloodworthiana was found o high elevations (1820 — 2360 m) in the Sierra Madre Occidental in the states of see and Durango, and V. eee occurred in similar, high elevations in the Sierra Madre Oriental of San Luis Potos RESUMEN Dos especies nuevas, Vitis bloodworthiana y V. jaegeriana, son descritas y Comparadas con las dos especies mas similares en la serie pedis Se encontré a Vitis bloodworthiana solamente en altitudes mayores (1820 59 m) en la Sierra Madre Occidental de los estados de Sinaloa y Durango; asimismo Vitis jaegeriana, ocurrio en altitudes mayores, pero en la Sierra Madre Oriental del estado de San Luis, Pot Two new species of Vitzs (Vitaceae) were found in mountainous regions of central and western Mexico. The closest species morphologically to these listed by Standley (1924) for Mexico appears to be V. arizonica Engelm. These species belong to series Occdentales Munson, which ts characterized by having leaves with small stipules (1 — 3 mm long), small fruit (4— 11 mm dia.) and flowering during mid season when grown along with other species (Munson 1909). Series Occidentales includes western North American species: V. arizonica Engelm., V. californica Bentham, V. girdiana Munson and V. ¢releasei Munson. Table | provides a comparison between the new species and the two most similar species in series Occidentales, V. avizonica and V. treleaser. No other species of North American Vitis, except V. rotundifolia Michx.., V. munsontana Planchon ex Munson and V. monticola Buckley are known to bear fruit with lenticels (Munson 1909). Fruit with fewer, less conspicuous lenticels were observed on some individuals of V. treleaser and V. arizonica during studies (Table 2). Both new species bear fruit with lenticels. Fruit of V. bloodworthiana observed from numerous vines in the field, consistently were covered with small, circular, tan lenticels that were less obvious as the fruit ripened and became black. Similar lenticels occurred on fruit of V. jaegeriana, although in some vines the lenticels were not conspicuous. Sipa 14(3):459 — 466. 1991. 460 NEW TAXA — VITIS BLOODWORTHIANA Comeaux, sp. nov. Fig. Caules angulatescens teres, glabrescentes ad interdum puberuli, striati, sine lenticellis; apices et folia immatura manifeste colorata cum pigmentum rubrum. Folia plerumque interdum sine lobata, longa- pare ad fere delroidea, longa-acumminata, cordata ad fere truncata lobi laterales interdum divergences acutati ad acuminati; lamina matura glabrata limitata ad venae primarias et axillares. Baccae ae ae, oa 6— 11 mm diam.; tacrae cum lenticellis circulares fulvae; semina 3.5 — m longa, 3— 3.5 mm lata. Vines to.10 m, stems on current season growth siebeceon or occasional- ly pubescent then turning glabrescent, striated; branchlets angled, becom- ing terete; internodes 3— 16 cm long; nodes rarely encircled with red pigmentation; pith interrupted at nodes by a diaphragm 2— 3 mm thick; bark brown, shredding during second season growth; lenticels absent; growing tips glabrous to glabrescent, occasionally pubescent, with white to tan trichomes, not enveloped by young leaves, ordinartly tips and imma- ture leaves prominently colored with red pigmentation; bud scales glabrous to pubescent, 3—4 mm long, brown. Leaves long-cordiform to nearly long- deltoid, flat, usually 3-lobed, with lateral lobes acute to acuminate, often divergent, apex long accuminate, base cordate to nearly truncate, lateral sinuses acute (rounded on ground shoots); margins serrate to nearly crenate, with teeth 0.5 — 3 mm long, oriented perpendicularly to margin, towards apex or base, triangular or with concave or convex sides, occa- sionally ciliate, with veins extending beyond teeth, midrib with 4—7, usually 6 pairs of prominent veins; /amina glabrous on both surfaces of mature leaves, except for simple, straight, pointed trichomes and arachnose ichoaies on primary veins and vein axils, 7— 13 cm wide, 9-17 cm long; petioles glabrous to puberulent, striated, 3 — 10 cm long; stipules brown, glabrous to pubescent, 1—1.5 mm wide, 1—2.5 mm long, caducous; pubescence white to tan, consisting of straight, pointed, simple trichomes and arachnose trichomes. Tendrils and inflorescences absent every third node, tendrils bifurcate or trifurcate, to 20 cm long. Inflorescences 1—8 cm long, peduncles 1.2—4.5 cm long, shoulder 1 —4.5 cm long. Flowers not observed. Fruit a berry, black, glaucous wth small, tan, circular lenticels, 0.6 — 1.1 cm in diameter; skin thin; pulp clear, greenish to purplish. ae brown, irregular in shape, ovate to nearly pyri- form, 3— 3.5 mm wide, 3.5—5 mm long. Tyee: MEXICO. DuRANGO: 16.5 km W of Del Diablo and 35.5 km E of Tropic of Cancer via Hwy 40, 2,300 m, 1 Jul 1986, Comeawx 4219 (HOLOTYPE: SMU; isoryprs: MEXU, a ATYPES. DURANGO: 39 km W of ee Diablo and 13 km E of Tropic of Cancer, via ra 40, 2,174 m, | Jul 1986, Comeaux 4214 (SMU); 37.4 km W of Del Diablo and 14.6 461] mt rrr a FIG. 1. Type specimen of Vitis bloodworthiana (Comeaux 4219). 462 km W of Del Diablo and 1 ), 2,236 m E a of palace via Hwy 40, 2,143 m, | Jul 1986, C aya (SMU), 33.4 km km E of Tropic of Cancer via Hwy m, 1 Jul 1986 Comeaux 4216 (SMU); oe km W of Del Diablo and 29.4 km ; of Tropic of Cancer via 40, 2,113 m, | Jul 1986, Comeaux 4 and 0.2 km W of Tropic of Cancer via Hwy 40, 1,820 m, Jul 1, W of Del Dib ey 50 m W ae Tropic Cancer via Hwy a, and 4211 Soa 52.0 km 1,990 m, 1 Jul 1986, Comeaux 4212 (SM Tropic of Cancer via Hwy 40, 2,150 m, 218 (SMU). Sinaloa: 52.3 km W of D 1986, Comeaux 4209, 421 el eee 9.6 km W of Del Diablo and 2.4 km E of Fi i. 1986, Comeaux 4213 (GA, SMU). Taste |. Comparison of four species of series Occrdentales Munson. Character V. bloodworthiana V. a V. trelea (11 Vines Sampled)(11 Vines Sampled) (12 ee veer 15 Vines Sampled) V. jaegerian 1. Leaves lobed usually *rarely usually usually 2. Leaves pubescent abaxial surface gs' gt, pr’, pb* gs, gt, pr, pb gs adaxial surface gs gs, gt 2S, pl gs Leaf in a he mean 0.7 0.7 0.9 0.9 range 0.5-0.9 0.6-0.9 0.8— 1.1 0.7—1.1 4. Teeth number (for 1 side of leaf) mean 27 20) 21 30 range 20-41 15 — 27 14— 26 4— 30 S. Midrib pairs per lea mean 5.4 4.3 4.0 4.0) range 4-7 4-5 3-5 55 6. Basal sinus width (in degrees) mean 107 : 79 range 70— 170 30 — 150 ** 30-175 -1— 150 7. Stems pubescent gs, gt, pr, pb pr, pb pr, pb gS, gt, pr 8. Stem tips pubescent gS, gt, pr pb gt, pr, pb gt, pr, pb 9. Stem tip (gy, Cw)’, re’, 8 (w), ¢, ru” w, (re) w, 2, pubescence color 10. Stem tip enveloped by leaves nl? fu n, f f, sl’ n,f,sl,st! I 1. Leaf and stem pubescence color w, t w, ¢, ru Ww w Q = rarely observed 5 -g = green 10-n = negative 1 - ys = glabrous 6-w = white ll -f = faintly 2-gt = glabrescent 7-re = red 12-sl = slighel 3.- pr = puberulent 8-c¢ = tan 13 -se = sctrongly 4 - pb = pubescent 9-ru = rufescenc *Only two leaves were observed from different vines out of many 1 d in the field. **Negative values relate to cases where leaf bases overlapped. 463 & 2. Specimens of Vitis arizonica and V. treleaser examined in comparison with bloodworthiana and jaegeriana. Viris arizonica Engelm. ARIZONA. Cochise Co.: 9.6 km S of Sierra Vista on Carr spe Rd., I, ee 5 J 1986, Comeaux 4236, 4237, 4238, 4239, 4240 and — (SMU). Santa Cruz Co.: 28 wy 83, 1,500 m, 5 Jul 1986, egos U); 28.5 km S of Sonoita via Hwy , 1,580 m, 5 Ju fi 1986, Cees 4243 and 4244 (SMU), 1.0 km me of Nogales via Hwy 82, 1,200 m, e ul 1986, ou (SMU); 3 km W of E. city eee in Nogales via Hwy 82, 1,175 m, 5 Jul 1986, Comeaux 4246, 4247 and 4248 (SMU). Vitis TRELEASEL Lee ARIZONA. Maricopa Co.: Fish Creek Hill bridge via Hwy 85, NW of Lost sae: St. Park, 700 m, 6 Jul 1986, Comeaux 4249, 4250, 4251, 4252 ap 4253 (SMU). NEW XICO. Catron Co.: 4 km NE of Aragon via si y 12, 2,100 m, 4 Jul 1986, Comeaux ea 4227 3 4228 (SMU); 3 km NE of Aragon via Hwy | ies m, 4 Jul 1986, plein 4229 (SM Socorro Co.: 1.5 guy downstream from Water Canyon Campground between stream and road, oa National os 2,180 m, 4 Jul 1986, Comeaux 4220, , 4222, 4223, 4224 and 4225 (SMU). This species is named in honor of P. J. Bloodworth (1950-), grape bree- der and fellow student under the direction of the late W. B. Nesbitt. Jeff ts acknowledged for his assistance in the author's research, willingness to share his great knowledge, and for his devotion to the vine. Also, the epithet bloodworthiana seems appropriate as the dark-red growing tips and young leaves that characterize this species are distinctive in the series. Vitis bloodworthiana was found only at high elevations (1,820 — 2,359 m) in pine forests within the Sierra de las Ventanas mountains, Sierra Madre Occidental, in Durango and Sinaloa. Annual recipitation is 40 — 80 cm and minimum temperatures range from -10° to 0° C (Rzedowski & Huerta 1978). This species occurs in dry or relatively moist, but well- drained sites, without sympatric species of Vitzs. More field studies are nee- ded to ascertain the overall distribution of V. bloodworthiana. The long-cordiform leaves, as indicated by the small leaf index values (Table 1) for V. bloodworthiana and V. jaegeriana, easily separate these from the two similar species in series Occidentales. Vitis bloodworthiana differs from V. jaegeriana in having more teeth and pairs of lateral veins per leaf, consistently broader basal sinuses, and the dark red-colored pigmentation in growing tips and young leaves. All of the nearly 70 seedlings grown in containers outdoors at Galveston College from one collection of V. blood- worthiana (Comeaux 4213) easily were differentiated within two months after germination from seedlings of numerous other species of Vztzs, in- cluding V. treleaset and V. arizonica, by their brilliant red growing tips and long-cordiform leaves. Four seedlings grown for one season, each with stems over 2 m in length, were distinguished readily from about 300 in- dividuals of similar size, representing numerous species and hybrids, by their red growing tips and various degrees of red pigmentation in all ma- ture leaves. 464 ViTIS JAEGERIANA Comeaux, sp. nov. Fig. 2. Caules angulatescens teres, puberuli obscura ad conspica, striati, sine lenticellis. Folia sine lobati plerumque, longa-cordiforma, long-accuminata, cordata, lamina matura supra glabras ad glabrescentes, infra puberuli leviter ad modice, alinquando glabrescente, tricho- mata fulva and ferruginea, vel alba. Baccae nigrae, glaucae, 6 — 11 mm diam., tectae cum lenticellis circulares fulvae; semina 3.5 —5 mm longa, 3—5 mm lata. Vines to 7 m, stems on current season growth puberulant to pubescent; striated; branchlets angled becoming terete; internodes 3— 10 cm long; nodes faintly to conspicuously encircled with red pigmentation, pith in- terrupted at nodes by a diaphragm 2 — 3 mm thick; bark brown, shredding during second season growth, lenticels absent; growing tips pubescent, tan or rufescent, occasionally white, not enveloped by young leaves; bud scales pubescent, 2—3 mm long, brown. Leaves long-cordiform, flat, rarely lobed, then lobes acute, apex /ong-acuminate, base cordate, lateral sinuses acute (when present); margin serrate, with teeth 0.5—3 mm long, oriented perpendicular to margin, towards apex or base, triangular or with convex sides, ciliate, with or without veins extending beyond teeth, midrib with 4 to 7 pairs of prominent veins; lamina with glabrous to glabrescent adaxial surfaces on mature leaves, abaxial surfaces pubescent to puberulent, occa- sionally glabrescent on mature leaves, not glaucous, with or without tufts of trichomes in axils of major veins, 4— 10 cm wide, 6— 15 cm long, petioles pubescent to puberulent, faintly striated, 1.3—7.5 cm long; stipules brown, pubescent to puberulent, |1— 1.5 mm wide, 1—3 mm long, caducous; pubescence tawny, rufescent or white, consisting of straight, pointed, simple trichomes or arachnose trichomes. Tendrils and inflorescences absent every third node, bifurcate, to 20 cm long. In- florescences 1.2 —5.8 cm long, peduncles 0.6—4.7 cm long, shoulders 0.2—2.3 cm long, occasionally replaced by a tendril. Flowers not observed. Fruit a berry, black, glaucous, with small, tan, circular lenticels, 0.6— 1.1 cm in diameter; skin thin pulp clear to purplish. Seeds brown, irregular in shape, ovate to nearly pyriform 3—5 mm wide, 4—5 mm long. Type: MEXICO. San Luis Porost: 86.7 km W of jet. Hwy 70 and 69 in a Verde, 2,150 m, 24 Aug 1987, Comeaux 4681 (HOLOTYPE: SMU; isorypes: MEXU, PH). PARATYPES: SAN Luis Porost: 94.6 km W of jct Hwy 70 and 69 in Rio Verde, 2 Sere m, 27 Jun 1986, Comeaux 4176, 4177 and 4178 (SMU); 75.7 km 1.c., 1,938 m, 28 Jun 1986, Comeanx 4206 (SMU); 92.6 km l.c., 1,815 m, 24 A Aug 1987, Comeaux 4674 —7 (SMU); 86.7 km l.c., 2,150 m, 24 Aug 1987, Comeaux 4679, 4680 ( This species 1s named for the grape breeder, Hermann Jaeger (1844 — 1895’), Neosho, Missouri (Smith 1962). Thomas V. Munson (1843 — 1913), the world renowned grape breeder, referred to Jaeger as 465 FIG. 2. Type specimen of Vites saegertana (Comeaux 4081). 466 “my esteemed co-worker” who “for more than twenty years hunt{ed] and hybridize{d} grapes” (Munson 1900). A grateful French government awarded Jaeger the Cross of the Legion of Honor in 1889 for his con- tributions towards saving the French wine industry, previously devastated by the phylloxera root louse (Smith 1962). Hermann and his brother, John Jaegar sent millions of grape cuttings to France as phylloxera resistant rootstocks for the native French varieties. Vitis jaegertana was found only at high elevations (1815 — 2400 m) in the Sierra de Juarez mountains, Sierra Madre Oriental, in San Luis Potosi. Veg- etation at the collection sites is dominated by Janiperus spp. Minimum temperature and annual precipitation are similar to those reported for V. bloodworthiana (Rzedowsik, J. and L. Huerta 1978). Vitis jaegeriana occu- pies relatively drier sites than the former species, and also has no associated sympatric species of Vitzs. Additional field work is needed to clarify the distribution of V. jaegeriana. Leaves of V. jaegeriana examined during field work consistently were without any lobing, except for a few isolated leaves observed only on two vines. This character easily separates if from V. bloodworthiana, the only species in series Occrdentales similar long-cordiform leaves. Another distinc- tive feature of V. jaegeriana is the generally tan pubescence on growing tips, and young stems and leaves. ACKNOWLEDGEMENTS The author ts grateful for the assistance of C. O. Foerster, Jr. and Dr. W. L. Hagerman during field studies in August 1987, and also for the assist- ance of Dr. PR. Fantz in the preparation of this manuscript. REFERENCES MUNSON, ay 1900. American grapes. Texas Agriculcural Ext. Sta. Bull. 56. —_________.. 1909. Foundations of American grape culture. T.V. Munson and Son, Deni- son, Tees 1S. RZEDOWSIK, J. and L. Huerta 1978. Vegetacién de México. Editorial Limus, Mexico, D.E SMITH, G. 1962. A man who ae into ea air after revolutionizing an industry. Evening Press, Carthage, Miss ):7. STANDLEY, PC. 1920 — 1926. ane ye cane of Mexico. Contr. U.S. Natl. Herb. 23. NOTEWORTHY PLANTS FROM NORTH FLORIDA. V. LORAN C. ANDERSON Department of Biological Science Florida State University Tallahassee, FL 32306-2043 U.S.A. ABSTRACT The following appear to be first reports for the state of Florida: Amthemis arvensts, Artemisia vulgaris, Carex leavenworthii, Cerastium semidecandrum, Cucuta maculata, Cladium mariscoides, Curcuma zedoaria, Hedyotis purpurea, Paspalum minus, Ranunculus ihc ae Silene virginica. Some additions to the Florida panhandle are documented here also, and several significant range extensions, particularly for rare or endangered taxa aes our area, are given This is the fifth installment of a series (Anderson 1984, 1986, 1988a, 1989) to update our knowledge of the flora of the Florida panhandle and Clewell’s (1985) guide to the flora. The area of coverage is from the Suwannee River west to the Alabama state line. New discoveries — 1.e., taxa not listed by Clewell — and range exten- sions of selected rare or senenuce noteworthy taxa are given here. Exotics that appear to be adventive or naturalized are also listed. Collections at FLAS, , USE and the Gholson Herbarium were consulted in addition to pertinent literature. Wilhelm (pers. comm.) provided updated distribu- tional data for his 1984 study, and Wunderlin (pers. comm.) shared his Florida checklist of vascular plant species. Voucher specimens for this re- port are at FSU unless noted otherwise. TAXA NEW TO THE AREA ACACIA ANGuStTISsIMA (P. Miller) Kuntze var. HirtTA (Nutt.) B. L. Robinson. Dixie Co.: frequent near junction rtes 358 and 361, juse NE of Jena, 2 Jun 1989, Anderson 12045 (FLAS, FSU). Jefferson Co.: Monticello, Jun 1931, J. K. Small s.n. (FLAS), new to Florida panhandle. AMORPHA HERBACEA Walt. var. HERBACEA. Dixie Co.: frequent in cut- over flatwoods bordering rte 361, ca. 6 air mi S of Steinhatchee, 2 Jun 1989, Anderson 12050; new to Florida panhandle. See map in Wilbur (1975) for previously known range. ANTHEMIS ARVENSIS L. Jackson Co.: weed in garden area, 1 mi W of Grand Ridge, 11 and 18 May 1980, A. K. Gholson 8298, 8302 (Gholson Herbarium); naturalized, new to Florida. Sipa 14(3):467 — 474. 1991. 468 ARTEMISIA VULGARIS L. Alachua Co.: roadside by wet woods N side of Gainesville, 11 Nov 1980, K. D. Perkins 855 (FLAS); Escambia Co.: Pensacola near Pensacola Bay, 17 Sep 1980, J. K. Burkhalter 7210 (FLAS); abundant along sandy roadside of Hollywood Avenue near Pensacola, 29 Oct 1989, J. R. Burkhalter 11717; naturalized, new to Florida. BOTHRIOCHLOA ISCHAEMUM (L.) Keng. var. SonGARICA (Fish. & Mey.) Celarier & Harlan. Escambia Co.: Naval Air Station, SW of Pensacola, 17 Sep 1988, J. R. Burkhalter 11138; vacant field S of Pensacola, 24 Sep 1988, J. R. Burkhalter 11149; W end of Santa Rosa Island, 28 Oct 1989, JOR. Burkhalter 11715; Santa Rosa Co.: hiway 98 E of Gulf Breeze, 6 Nov 1988, J. R. Burkhalter 11310; Washington Co.: beside hiway 90 in Chipley, 10 May 1990, L. C. Anderson 12714; native, new to Florida panhandle. CAREX LEAVENWORTHIL Dewey. Gadsden Co.: locally established near Marion Street, Chattahoochee, 10 May 1990, L. C. Anderson 12681, 26 Aug 1985, A. K. Gholson 11305 (Gholson Herbarium); Jackson Co.: Neal's Landing, Lake Seminole, under Quercus above floodplain, 18 May 1982, A. K. Gholson 9729 (Gholson Herbarium); native, new to Florida. CERASTIUM SEMIDECANDRUM L. Escambia Co.: abundant on sandy roadside of Saufley Field Road near Pensacola, 9 Mar 1990, J. R. Burkhalter IISI1; naturalized, new to Florida. Cicura MAcuLATA L. Jackson Co.: frequent along open border of pine-oak woodland on S side of I-10, ca. 5 air mi SSW of Sneads, 26 May 1990, 12 Jul 1990, L. C. Anderson 12846, 13093; apparently native, new to Florida. CLADIUM MARIScOIDES (Muhl.) Torr. Santa Rosa Co.: abundant in ditch E of Gulf Breeze on S side of hiway 98, 16 Jul 1989, J. R. Burkhalter 11500, native, new to Florida. Kiikenthal (1942) listed Florida as part of this species’ range, but recent workers have found no documented collec- tions for the state—see Bridges and Orzell (1989) for notes on the ecology and geography of this species in the eastern United States. CuURCUMA ZEDOARIA (Christm.) Rosc. Leon Co.: locally established along margin of Freeman Creek Cove of Lake Talquin, ca. 17 air mi W of Tallahassee, 24 May 1989, L. C. Anderson 12007; naturalized, new to Florida. Cyperus ECHINATUS (L.) Wood. Gulf Co.: edge of wet flatwoods 5.5 air mit NW of Wewahitchka, 15 Jun 1989, L. C. Anderson 12095; native, new to Florida panhandle. ELBOCHARIS ROSTELLATA Torr. Taylor Co.: common in roadside ditch beside tidal marsh near mouth of Fish Creek, ca. 2.5 air mi SSE of Keaton Beach, 2 Jun 1989, L. C. Anderson 12060; Wakulla Co.: St. Marks, 1843 469 or 1845, F Rugel 281 (FLAS); not listed by Clewell (1985) for the Florida panhandle. HepyorTis PURPUREA (L.) T. & G. Jackson Co.: alluvial area below Neal’s Landing, Lake Seminole, 24 Apr 1972, Gholson 3039, 3040 (Ghol- son Herbarium), Neal’s Landing, 17 May 1978, Gholson 7100 (Gholson Herbarium); native, new to Florida. INDIGOFERA SPICATA Forskal. Leon Co.: frequent in newly sodded lawn, ES.U. campus, Tallahassee, 21 May 1990, Anderson 12851; fallow field, Tallahassee, 3 Dec 1990, Anderson 13342. Taylor Co.: frequent in dry sand of disturbed site along Alvarez Street in Perry, 21 Jun 1989, Anderson 12067 (FLAS, FSU); naturalized, new to Florida panhandle. Morton (1989) reports this species 1s widespread in southern Florida, where it is a hazard to grazing animals (some horses have been fatally poisoned). MurDANNIA KEISAK (Hassk.) Hand.-Mazz. Jackson Co.: extensive mats along shoreline of Lake Seminole near Parramore Landing, 13 Sep 1976, Gholson 3553 (Gholson Herbarium); naturalized, new to Florida panhandle. This Asian species has been spreading in the southeastern United States relatively recently (Dunn and Sharitz 1990). PasPpALUM MINUS Fourn. Escambia Co.: near Perdido River, N of hwy 90 and NW of Pensacola, 7 Aug 1990, Burkhalter 12223; native, new to Florida. RANUNCULUS MARGINATUS D’Urville var. TrRAcCHYCARPUS (Fischer & Meyer) Azn. Washington Co.: moist sandy loam of shaded floodplain of Holmes Creek at roadside park beside rte 79 just N of Vernon, 4 May 1990, Anderson 12631; native, new to Florida (see Keener and Hoot 1987). SILENE VIRGINICA L. Bay Co.: frequent with Carex baltzellii on shaded steep slopes of ravine E of Hammond Lake, ca. 3.5 air mi WNW of Fountain, 4 May 1990, Anderson 12679; new to Florida. This species has been found sporadically in other southern states (Moore 1956), and Kral (1966) suggests this Pleistocene relict has been able to persist by inhabit- ing ravine banks where cold air drainage provides a suitable niche in an otherwise inhospitable environment. SOLANUM PsSEUDOCAPSICUM L. Jefferson Co.: persisting shrub in mesic hardwoods of Avalon Plantation, S of Capps, 26 Apr 1989, Gholson 12126 (Gholson Herbarium), | Mar 1989, Godfrey 83004; naturalized, new to Florida panhandle. ADDITIONAL RANGE EXTENSIONS The following collections represent additional counties of record for taxa listed by Clewell (1985) from only one county, or they are significant range 470 extensions for rare or otherwise noteworthy species. ARNOGLOSSUM DIVERSIFOLIUM (T. & G.) H. Robins. Holmes Co.: 4 mi S of rte 2 on rte 177, 26 May 1967, Smith 1324 (FLAS). Jackson Co.: Chipola River, 6 mi S of Marianna, 6 Jun 1957, Kral 4813 (FLAS). Walton Co.: shaded floodplain of Choctawhatchee River just S of rte 20, 25 May 1990, Anderson 12814. Washington Co.: Holmes Creek at rte 280, 30 Jul 1954, E. S$. Ford 3779 (FLAS); upper Holmes Creek, ca. 5.5 air mi SW of Chipley, 31 May 1985, Anderson 8207, 15 Jun 1989, Anderson 12111; Hightower Spring, 3.5 air mi WSW of Vernon, 25 May 1990, Anderson 12820; Brunson Landing, Holmes Creek, ca. 3 air mi WSW of Vernon, 15 Jun 1990, Anderson 12941; Live Oak Landing, Holmes Creek, 7.5 air mi WSW of Vernon, 22 Jun 1990, Anderson 12958; new counties of record for this threatened species (state listed, Wood 1990). ASCLEPIAS RUBRA L. Escambia Co.: 3.5 mi E of Muskogee, 7 Jun 1962, E. S. Ford 6312a (FLAS); Spanish Mill Creek at Gonzalez, 26 Jul 1980, Burkhalter s.n. (FLAS). Santa Rosa Co.: edge of mesic thicket, Paquette Camp, Blackwater State Forest, 29 Jun 1990, Anderson 13013. Walton Co.: bayhead 6.5 mi SE of DeFuniak Springs, 1 Jun 1954, West & Arnold s.n. (FLAS); boggy area 3.5 mi S of DeFuniak Springs, 29 May 1967, Smith 1335 (FLAS). CAREX BALTZELLU Chapm. ex Dewey. Escambia Co.: wooded slope on Univ. W. Fla. campus near Pensacola, 2 Apr 1978, Burkhalter 5748 (FLAS). Okaloosa Co.: 6.5 air mi N of Ft. Walton Beach, 20 Mar 1968, Ward 06603 (FLAS); near Rogue Creek, 7 mi NW of Niceville, 20 Mar 1968, Ward 6599 (FLAS). Santa Rosa Co.: Weaver Creek, 5 mi N of Holley, 21 Mar 1968, Ward 6637 (FLAS). Washington Co.: frequent on shaded slopes of sinkhole ravine ca. 12 air mi S of Chipley, 26 May 1990, Anderson 12830. This endangered species (Wood 1990) is nearly endemic to the Florida panhandle; Muller et al. (1989) give the previously known range of this species. Carex seEORSA Howe. Liberty Co.: common on floating islands in beaver-dammed swamp of Pittman Creek just N of rte 20, 0.8 mi W of Ochlockonee River, 22 May 1990, Anderson 12741. CONOPHOLIS AMERICANA (L.f.) Wallr. Okaloosa Co.: oak-beech- magnolia woods SW of Laurel Hill, 3 Mar 1990, Burkhalter 11807; new to western panhandle (not listed by Wilhelm 1984). DioctEA Muttiflora (T. & G.) Mohr (= Galactia mohlenbrockii Maxwell). Gadsden Co.: mesic hardwoods of Chattahoochee Nature Park below Morgan Avenue near Apalachicola River on SW side of Chatta- hoochee, 4 June 1990, Anderson 12858. Washington Co.: head of Blue 471 Springs just E of Econfina Creek, ca. 14 air mi SE of Vernon, 24 May 1990, Anderson 12708. DroseRA FILIFORMIS Raf. Washington Co.: with D. intermedia on open, peaty shore of Lucas Lake, 8 Jun 1990, Anderson 12875, edge of Rattlesnake Lake, 5 Jul 1990, Anderson 13042; with D. tracyi (no signs of intergradation) in small seepage bog on S side of Gully Lake, 5 Jul 1990, Anderson 13048. HeEpyoris NUTTALLIANA Fosberg. Washington Co.: frequent in dry sand of open turkey oak woodland ca. 12 air mi S of Chipley, 26 May 1990, Anderson 12831. The only other collection in Florida from Walton County was listed as quite atypical by Terrell (1959). Juncus GyMNocarpus Coville. Washington Co.: sphagnum seepage in mixed hardwoods of steephead 5 air mi SE of Vernon, 31 May 1985, Anderson 8200; loamy sand of mesic woodland E of Gap Lake, ca. 15 air m1 S of Chipley, 11 May 1990, Anderson 12721; with Kalmia latifolia along White Oak Creek, NE of Gap Lake, 8 Jun 1990, Anderson 12897. Lupwicia erecta (L.) Hara. Walton Co.: marshy border of Fuller Lake on Coffeen Nature Preserve near Four Mile Village just E of Sandestin, 21 Oct 1989, Anderson 12487. The species was first reported for the Florida panhandle by Anderson (1986); this collection extends its range westward significantly. LupwiGIA LANCEOLATA El]. Walton Co.: Pine flatwoods between coas- tal dunes and Fuller Lake in Four Mile Village (Coffeen Nature Preserve), 21 Oct 1989, Anderson 12479. This is a range extension westward from Franklin County (Peng 1989). MATELEA FLAVIDULA (Chapm.) Woodson. Washington Co. : infrequent in hardwoods on upper slopes of small sinkhole ravine just N of Wash- ington Blvd, 12 air mi S of Chipley, 8 Jun 1990, Anderson 12891. Neprunia pupescens Benth. Taylor Co.: Frequent along edge of tidal marsh at N edge of Keaton Beach, 2 Jun 1989, Anderson 12062. PHYSOSTEGIA GODFREY! Cantino. Walton Co.: swampy ditch SW of Bruce, 18 Jun 1971, H. A. Davis 15829 (FLAS); locally common in mesic roadside depression along rte 20 just W of Black Creek bridge, 7.7 mi E of Freeport, 24 May 1990, Anderson 12773. These represent a significant range extension to the west for this Florida panhandle endemic (Cantino 1D). PINCKNEYA BRACTEATA (Bartr.) Raf. Washington Co.: edge of Mag- nolia-Liriodendron thicket bordering Boggy Branch, 1.9 air mi W of rte 77 and Greenhead, 26 May 1990, Anderson 12836; listed as threatened in Florida (Wood 1990). 472 PLUCHEA OBLONGIFOLIA Nash. Dixie Co.: 4 mi N of Shired Island, 10 Jul 1989, Godfrey 83348. Taylor Co.: mesic woodland near Fish Creek, 3.5 air mi SE of Keaton Beach, 2 Jun, 1989, Anderson 12059. RHEXIA SALICIFOLIA Kral & Bostick. Okaloosa Co.: E of Destin along Four Prong Lake, 11 Jul 1990, A. F Johnson 8747; new county record for this rare species (Bounds 1987). Its center of distribution appears to be the karst lake region of Washington County, where it is usually associated with endangered Hypericum lissopbloeus and Xyris longisepala (Wood 1990), and the following collections further document its presence there: Wash- ington Co.: Chain Lakes, W of Greenwood, 6 Jul 1963, Godfrey 62951 (FLAS); Sparkleberry Lake, 15 Jun 1990, Anderson 12935, Piney Lake, 6 Jul 1990, Anderson 13077; Crystal Lake, 6 Jul 1990, Anderson 13082, 3 Aug 1990, Anderson 13153; Sand Lake, 3 Aug 1990, Anderson 13164, Whitewater Lake, 3 Aug 1990, Anderson 13170; Porter Lake, 3 Aug 1990, Anderson 13177; Major Lake, 1 Oct 1990, Anderson 13296. RHYNCHOSPORA CRINIPES Gale. Santa Rosa Co.: banks of Sweetwater Creek, ca. 3 air mi S of Munson, 5 Aug 1989, Johnson 8368; Big Coldwater Creek, ca. 0.6 mi above rte 191 bridge, 29 Jun 1990, Anderson 12996. These collections increase the known range considerably for this very rare species (Anderson 1988b). STACHYS HyssopirOLiA (Michx.) var. LYTHROIDES (Small) J. B. Nel- son. Jefferson Co.: periodically wet pine-hardwood stand on Norias Plan- tation, NNE of Lake Miccosukee, 19 Jul 1990, Godfrey 83917, 27 Jul 1990, Anderson 13098. This Florida endemic was previously known only from a few collections in Leon County (Nelson 1981). Uvu LaRIA SessiLirotia L. Walton Co.: Knox Hill, 22 Mar 1968, Ward 6643 (FLAS). Washington Co.: 26 May 1990, Anderson 12827. XYRIS LONGISEPALA Kral. Walton Co.: 3.5 mi SSW Mossy Head, 7 Aug 1969, Ward 7216 (FLAS); 3 mi SW Mossy Head, 10 Sep 1979, Judd 2255 (FLAS). Washington Co.: sandy shores of the following karst lakes: Parrish Pond, 8 Jun 1990, 5 Jul 1990, Anderson 12883, 13028; Boat Lake, 8 Jun 1990, Anderson 12885; Fox Pond, 8 Jun 1990, Anderson 12902; Sparkleberry Lake, 15 Jun 1990, Anderson 12934, 12936; Hammock Lake, 5 Jul 1990, Anderson 13037; Rattlesnake Lake, 5 Jul 1990, Anderson 13044; Gully Lake, 5 Jul 1990, Anderson 13047; Piney Lake, 6 Jul 1990, Anderson 13079; Crystal Lake, 3 Aug 1990, Anderson 13156; Sand Lake, 3 Aug 1990, Anderson 13163; Whitewater Lake, 3 Aug 1990, Anderson 13171; Porter Lake, 3 Aug 1990, Anderson 13173, 13176; unnamed lake W of Porter Lake, 3 Aug 1990, L. C. Anderson 13180; Mule Pond, W of Sunny Hills, | Oct 1990, Anderson 13293; Major Lake, 1 Oct 1990, Ander- son 13.298; new counties of record (Clewell 1985) for this Florida endange- 473 red species (Wood 1990). A Leon County collection (Lake Iamonia, 21 Aug 1989, L. C. Anderson 12198) is particularly interesting because at that loc- ation the plants occur in peaty muck of floating islands in the lake rather than in coarse sands bordering karst ponds as in all other known popula- tions. ACKNOWLEDGMENTS My field work in 1990 was sponsored in part by the Florida Natural Areas Inventory. James R. Burkhalter, A. K. Gholson, and R. K. Godfrey are acknowledged for their continuing contributions to our knowledge of the flora of the Florida panhandle. REFERENCES ANDERSON, ic 1984. Noteworthy plants from north Florida. Sida 10:295 — 297. 5. Noteworthy plants from north Florida. I. Sida 11:379 — 384. o 88a. Noteworthy plants from north Florida. HI. Sida 13:93 — 100. ide Status of endangered Rhynchospora crinipes (Cyperaceae). Syst. Bot. 3:405 — 41 ie ono: Noteworthy plants from north Florida. IV. Sida . oe BOUNDS, R. R. 1987. Rare species of Rhexia L. Castanea 52:304 — BRIDGES, E. L., and S. L. ORZELL. 1989. Additions and notewor ee vascular plant sane from Texas and Louisiana, with historical, en and geographical ( Phytologia 66: 209: CANT INO, PD. 1979. a godfrey: (Lamiaceae), a new species from northern Florida. Rhodora 81:409 — 417. CLEWELL, A. E 1985. Guide to the vascular plants of the Florida panhandle. Florida State University ae Presses of Florida, Tallat DUNN, C. P, and R. R. fe 1990. The history of Srdoni ketsak (Commeli- naceae) in the ee United States. Castanea 55:122— 129. KEENER, C. S., and S. B. HOOT. 1987. Ranvnnlns section tes (Ranunculaceae) in the southeastern United States. Sida 12:57 — 68 KRAL, R. 1966. Seer on the flora of ie out henna United States with special eference to northern Louisiana. Sida 2:395 — 408. KUKENTHAL, G. 1942. Vorarbeiten zu einer Monographie der Rhynchosporideae. XII. age Repert. Spec. Nov. Regni. Veg. 51:1— 17, 1 _ A. 1956. Ph virginica in the Gulf states. Rhodora 58: 27 = 29, oon J. E 1989. eping indigo ( oe spicata Forsk.) (Fabaceae) — a hazard to herbivores in Mr Econ. Bot. 43:314— MULLER, J. W., E. D. HARDIN, D. - JACKSON, S. E. GATEWOOD, and N. CAIRE. 1989. ae report on the vascular plants, animals, and plant communities endemic to ue Florida Game and Freshwater Fish Commission, Nongame Wild- life uae Tech. Rep. no. 7, Tallahassee. NELSON, J. B ee eee (Labiatae) in southeastern United States. Sida 9:104 — PENG, C. 1989, The systematics and peers of Ludwigia sect. er — Ann. Missouri Bot. Gard. 76 — 402. TERRELL, E. E. 1959. A revision of the ieee purpurea group (Rubiaceae). Rhodora Panera 188 — 207. AT4 WILBUR, R. L. 1975. A revision of the North American genus Amorpha (Legumi- ee Rhodora 77:337 — 409. WILH ELM, G.S. 1984. Vascular flora of the Pensacola region. Unpubl. doctoral disserta- tion, Southern [Illinois University, Carbo WOOD, D. A. 1990. Official lists of seein and potentially endangered fauna and flora in Florida. Florida Game and Fresh Water Fish Commission (1 August), Talla- hassee, A REPORT OF CYPERUS GRAYIOIDES AND CYPERUS RETROFLEXUS (CYPERACEAE) NEW TO MISSOURI AND NOTES ON OTHER SELECTED MISSOURI CYPERUS RICHARD CARTER Herbarium (VSC), Department of Biology, Valdosta State College Valdosta, GA 316098, U.S.A. CHARLES T. BRYSON USDA, ARS, Southern Weed Science Laboratory Stoneville, MS 38776, U.S.A. ABSTRACT Field work in southeastern Missouri during 1989 and 1990 has resulted in a number of noteworthy Cyperus records. Cyperus grayivides and Cyperus retroflexus are reported new to Missourt. Also, additional records of Cyperus croceus and Cyperus * mesochorus and two previ- ously unknown Cyperus hybrids are reported. INTRODUCTION Our investigation has been centered on a system of dry sandy ridges and rises in Mississippi, New Madrid, and Scott counties of southeastern Mis- souri. This part of Missouri is located in the Mississippi Embayment, a northward extension of the Gulf Coastal Plain (Fenneman 1938, Walker and Coleman 1987). Soils of the Scotco series [previously classified as Crevasse series} occur on and along these prominent sandridge formations, which rise as much as 30 feet above the surrounding floodplain. Scotco soils are excessively drained and droughty, coarse sands (Brown 1977, Fes- tervand 1981). Many of the natural Scotco sandridge habitats are highly disturbed or have been destroyed by row-crop agriculture and construction of buildings or roads. The presence of Cyperus grayioides and certain of its associates on the Scotco sandridges of southeastern Missouri indicates a floristic affinity with sandridges of eastern Texas and adjacent Louisiana and sand prairies of central and northern Illinois (Bowles et al. 1986, Bridges and Orzell 1989). Furthermore, it is interesting to note that prairie vegetation origi- nally occurred to a limited extent in southeastern Missouri on these coarse sandy soils (Brown 1977). Sipa 14(3):475 — 481. 1991. 476 CYPERUS GRAYIOIDES NEW TO MISSOURI Cyperus grayioides Mohl. is an obscure member of Cyperus section Lax- iglumi, which until recently was poorly known and infrequently collected. It was originally described from sand-prairies of northwestern Illinois (Mohlenbrock 1959), and its occurrence in eastern Texas and Louisiana on xeric sandridges was subsequently documented in a thorough taxonomic treatment of Cyperus section Laxiglumi by Marcks (1972). More recently, additional populations in eastern Texas were reported (Bridges and Orzell 1989). Cyperus graytides is listed among “candidates for possible addition to the List of Endangered and Threatened Plants,” category 2, by the U. S. Fish and Wildlife Service (1990). It was thought that the Texas and Louisiana populations were disjunct by a distance of more than 850 km rom the nearest Illinois sites. Cyperus grayioides has not been previously recorded from the state (Yatskievych and Turner 1990). It is locally abundant on open, Scotco sands in southeastern Missouri, where it is associated with the following species: Cenchrus longispinus (Hack.) Fern., Conyza canadensis (L.) Cronq., Croton glandulosus L., Cycloloma atriplicifolium (Spreng.) Coulter, Cyperus lupulinus (Spreng.) Marcks ssp. /upulinus, C. lupulinus ssp. macilentus (Fern.) Marcks, Diodia teres Walter, Eragrostis cilianensis (All.) Vign. ex Jan- chen, Exphorbia dentata Michaux, Froelichia floridana (Nutt.) Mog. var. campestris (Small) Fern., Helranthus petiolaris Nutt., Heterotheca subaxillaris (Lam.) Britton & Rusby, Monarda punctata L., and Opuntia humifusa Raf. A distribution map based upon examination of specimens (ILL, MO, NLU, TEX-LL, VSC) and other data (Mohlenbrock 1959, Marcks 1972, Bowles et al. 1986, Bridges and Orzell 1989) is shown in Figure 1. Collec- tion data for Cyperus grayioides in Missouri are given below. MISSOURI. Mississippi Co.: 0.2 mi E of jct of county roads 408 and 433, S of county road 408 by about 0.2 to 0.4 mi, T26N R1I4E 826, rim of sandy ridge around pine thicket and along old fence row, open, highly disturbed area, 26 Sep 1990, Bryson 10472 (ctb- personal herbarium of C.T. Bryson, MO, VDB, VSC); 0.5 mi W of jct of county road CC and hwy 1-57/US 60, just N of hwy I-57, near Scott-Mississippi county line, 26 Sep 1990, Bryson 10474 (ctb, VSC). New Madrid Co.: ee i in floodplain, E of hwy 1-55 front- age road, 1.0 mi S of Sikeston city limit, T25N RI4E NW % S83, locally common on loose sand, 27 Aug 1989, Carter 8263 (BE, MO, a Tor. VSC); sandy rise in floodplain, i of hwy I-55 frontage road, 0.8 mi S$ of Sikeston city limit, T26N R 14E SW 4 $34, 36° 50! 41" N, 89° 31' 52” W, locally abundant on loose sand, 27 Aug 1989, Carter 8267 ee MIC H, MO, NY, NYS, eee SMU, US, VDB, VSC, WIS); 6.6 to 7.1 mi N of jet of hwy MO 80 and county ro¢ id AA, S of Sikeston city limits, t ou I- ee an RI4E NE ve open sandy area, 26 Sep on ae 10460 (ctb, IBE, MICH, MO, SMU, SWSL VSC); 4.7 mi N of jet of hwy MO 80 and county ae AA, S of seni T25N,. “ ee NW, $11, open sandy area, 26 Sep 1990, Bryson 10462 (ctb, VDB, VSC); 6.6 mi N of jet of hwy MO 80 and county road AA, S of Sikeston; T25N R14E S3, along E side of county 95° +-27° 350km FIG. 1. The distribution of Cyperus grayioides. road AA, open gentle slope at base of clones sandy soil, 27 Sep 1990, es 10509 (ctb, VDB, VSC). Scott Co.: 0.2 mi W of jct of hwy MO 77 and county road 514, T27N RISE SEM% a sandy soil in ditch he county road 514, 26 Sep 1990, Bryson one (ctb, MO, VDB, VSC),; 2.1 mi E of Blodgett city limit and jct of county roads 514 and H, T27N RISE SW y SG, open areas along roadside, crest and slopes of sandy ridge N of county road 514, sandy soil, 26 Sep 1990, Bryson 10481 (ctb, MO, SMU, VDB, VSC) and 10483 ee IBE, MICH, MO, NLU, NYS, SMU, SWSL, TAES, US, VDB, VSC); 1.8 mi SSE of hw I-55 and county road H by dead end, unnumbered access road, T27N RI4E SW $34, gentle slope near road, open sand, 26 Sep 1990, Bryson 10498 (ctb, VDB, VSC). 478 CYPERUS RETROFLENUS NEW TO MISSOURI Cyperus retroflexus Buckley [= Cyperus uniflorus Torrey & Hooker, non Thunberg (fide Tucker 1987)} ranges from northern Mexico into New Mexico and throughout much of Texas and eastward into Oklahoma, Arkansas, and Louisiana (Carter, in prep.). Recently, it has been reported from Mississippi and Alabama (Carter, Bryson and Lipscomb 1987). Dur- ing September 1990, an extensive population of Cyperus retroflexus was dis- covered growing on a Scotco sandridge in Mississippi County, Missouri. This species has not been previously reported from Missouri (Yatskievych and Turner 1990). Collection data for C. retroflexus in Missouri follow. MISSOURI. Mississippi Co.: 0.2 mi E of jet of county roads 408 and 433, S of Sa road 408 about 0.2 to 0.4 mi, T26N RI4E 826, along top of sandy ridge, er pin thicket and along old fence row, most of area highly disturbed and open, 26 Sep 1590, Bryson 10473 (ctb, IBE, MICH, MO, NLU, SMU, SWSL, TAES, VSB, VSC). ADDITIONAL RECORDS OF CYPERUS X MESOCHORUS During 1989, plants with sharply angled, scabrid culms; ascending bracts; and multiple, pedunculate inflorescence rays were located in New Madrid County, Missouri, along an open roadside and edge of an adjacent field in coarse sandy soil of the Scotco series (Brown 1977, Festervand 1981), only about one-half mile from the aforementioned Cyperus grayioides Bite, Initially, these plants were placed with Cyperus schweinitzii Torrey, how- ever, a critical examination indicates they are actually Cyperus & mesochorus, a hybrid between C. schweinitz1i and C. lupulinus (Spreng.) Marcks ssp. lupulinus (Marcks 1974). Cyperus X mesochorus Geise is rare in Missouri, and has not been previously reported from the southeastern quadrant of the state (Steyermark 1963, Yatskievych and Turner 1990). Cyperus X mesochorus was found growing with only one of its putative parents, Cyperus lupulinus (Spreng.) Marcks ssp. /upulinus. However, its sharply angled, scabrid culms and ascending spikelets leave little doubt that C. schwernitzi is its other parent. Although it is disconcerting that C. schweinitzi was not found at any of these sites, this kind of problem is not unprecedented in Cyperaceae (Cayouette and Morisset 1985). Moreover, further investigation may document the occurrence of C. schweinitzii in the vicinity, which would be similar to a situation in Carex described by Cat- ling et al (1989). These plants also exhibit reduced fertility (ca 50% mean seed set) when compared with Cyperus schweinitzi1 (294%), which indicates a hybri origin. The low fertility observed in this southeastern Missouri population is consistent with observations of Marcks (1974). Additionally, a number 479 of the specimens are intermediate and difficult to place taxonomically, which suggests that introgression, as documented by Marcks (1974), has occurred. Thus, we hypothesize that (1) Cyperus schweinitzii is rare in southeastern Missouri and was overlooked in our brief field work; (2) the intermediate plants have been formed by backcrossing between F1 hybrids and either or both parents; and (3) C. X mesochorus is the result of hybridization or intro- gression between Cyperus schweinitzii and Cyperus lupulinus ssp. lupulinus. Collection data for Cyperus X mesochorus follow. MISSOURI. Mississippi Co.: 0.2 mi E of jct of county roads 408 and 433, S of county road 408 about 0.2 — 0.4 mi, T26N RI4E 826, rim of sandy ridge around pine thicket and along old fence row, highly disturbed open sand, 26 Sep 1990, Bryson 10471 (ctb, MO, VDB, VSC). New Madrid Co.: sandy rise in floodplain, E of hwy I-55 frontage road, 0.4 S of Sikeston City limit, T26N RI4E NW S34, locally abundant in loose sand, 27 hee 1989, Carter 8276 IBE, MICH, MO, NY, NYS, SMU, US, VDB, VSC); 0.5 mi N of jct of hwy MO 80 and county road AA and 0.25 mi E of county road AA. T25N R14E 835, open well drained sandy ridge, 27 Sep 1990, Bryson 10510 (ctb, VSC). A RECENT COLLECTION OF CYPERUS CROCEUS The correct name for the species long known as Cyperus globulosus Aublet is Cyperus croceus Vahl (Carter and Kral 1990). In the United States Cyperus croceus Vahl is distributed from New Jersey southward throughout Florida then westward into eastern Texas and Oklahoma. It is common in the Atlantic and Gulf coastal plains and occurs sporadically inland into Tennessee and Missouri (Carter, in prep.). Cyperus croceus was collected in Missouri in the late 19th and early 20th centuries but apparently has not been collected there since 1910. During 1989, a population of Cyperus croceus was located in New Madrid County, Missouri, where it was growing in sandy loam soil in a poorly kept lawn. This collection is the first of its species from Missouri in nearly 80 years. Data for all Missouri specimens of C. croceus, which we have examined, are given below. URI. Dunklin Co.: “sandboden,” without locality, 27 Jul 1893, H. Eggert s.n. (MO 759399); without locality, 18 Sep 1893, Bush 140 (NY); Kennett, 27 Jul 1895, Bash 683 (NY); Campbell, woods, 7 Sep 1910, Bush 6222 (NY, US); Malden, sands, 8 Sep 1910, Bush 6270 (NY, US). New Madrid Co.: New Madrid, poorly kept churchyard about a block W of levee, at corner of Russell and pg gees streets, T23N R14E S34, locally common, 2 Aug 1989, Carter 8201 (M B, VSC). St. Louis Co.: Allenton, Jul 1889, G hie ee s.n. (MO 795915); a a Louis: without locality, 12 Jul 1884, G. W. ees sn. (NY). PREVIOUSLY UNREPORTED CYPERUS HYBRIDS Cyperus lupulinus (Spreng.) Marcks ssp. lupulinus X Cyperus strigosus L. — Hybrid plants of low fertility (<< 1%) were found in Scott County 480 growing with Cyperus lupulinus ssp. lupulinus and Cyperus strigosus. The hybrids were found along a gentle slope between a swale and well drained sandy rise. Furthermore, this site was artificially watered by irrigation runoft. Cyperus strigosus was located in the swale, and Cyperus lupulinus ssp. lupulinus on the sandy rise. Presumably, the artificial water source has produced an intermediate habitat favorable for survival of hybrids. This situation appears to be analagous with the hybridized habitat described by Anderson (1949). The reduced fertility of these hybrids is not surprising since the parents are not closely related. Following are collection data for this hybrid. MISSOURI. Scott Co.: NW of jct of hwy US 1-55 and US 62 and between veges Inn and I- ae Pe soil, 25 Sep 1990, Bryson 10453 (ctb, MICH, MO, SMU, SWSL, VDB ni SSE of jct of hwy US 1-55 and an road H, by dead- ans unnum- area access ae "SW Y1 of SW of Sec. 34, T27N, RI4E, open sandy soil, gentle slope near road, 26 Sep 1990, Bryson 10500 (ctb, VSC &; WS Cyperus grayioides Mohl. X Cyperus lupulinus (Spreng.) Marcks ssp. macilentus (Fern.) Marcks. — A single completely sterile hybrid plant was cae growing with Cyperus grayioides, C. lancastriensis Porter in Gray, and C. Lupulinus ssp. lupulinus. This plant is apparently an Fl hybrid between Coprras graytotdes and Cyperus lupulinus (Spreng.) Marcks ssp. macilentus (Fern.) Marcks and is intermediate between these taxa with respect to spikelet posture and anther length. Following are collection data for this hybrid. MISSOURI. Scott Co.: 0.2 mi W of jet of Hwy MO 77 ane county road 514, T27 RISE, SEM, Sect. 5, sandy ae in ditch along county road 514, 26 Sep 1990, Bryson 10479 (ctb). ACKNOWLEDGEMENTS We are grateful to George Yatskievych (MO) for thoroughly reviewing the manuscript; to Tony Reznicek (MICH) for pointing out references deal- ing with similar hybrid situations in Carex; to Bill Harrison and Cary Norquist, U.S. Fish and Wildlife Service, and Bill McClain, Illinois Department of Conservation, for providing information about the official status of Cyperus grayioides; and to Clayton Robertson, USDA-SCS, New Madrid, MO, for sending a soil survey of New Madrid County. _ REFERENCES ANDERSON, E. ee Introgressive aaa John Wiley & Sons, Inc. New York. BOWLES, M.L., D.R. KURZ, R. NYBOER, and J.E. SCHWEGMAN. 1986. a report on Capers coy Unpublished report to the U.S. Department of Interior, Fish and Wildlif vice BRIDGES, E.L. and an ORZELL. 1989. Additions and noteworthy vascular plant collec- 481 tions from Texas and Louisiana, with historical, ecological and geographical notes. Phytologia 66:12 — 69. BROWN, B.L. 1977. Soil survey of New Madrid County, Missouri. U.S.D.A. Soil Con- servation Service in cooperation with Missouri Agricultural Experiment Station. U.S. Government — Office: 1977 — 204/697/15. CARTER, R. in prep. A systematic treatment of Cyperus section Umbellati in North America. , C.T. BRYSON and B.L. LIPSCOMB. 1987. Cyperus uniflorus (Cyperaceae) east of the Mississippi River. Sida 12:250 and R. KRAL. 1990. Cyperus echinatus and Cyperus croceus, the correct names for North American Cyperus ovularis and Cyperus globulosus (Cyperaceae). Taxon a9 32232). CATLING, PM., A.A. REZNICEK, and K. DENFORD. 1989. Carex lacustris XC. alll iCipe eraceae), a new natural — Canad. J. Bot. 67:790 CAYO E, J. and P MORISSET. 1985. omosome studies on natural hybrids oe maritime species of Carex (sections Pha and acon gees in northeastern North America, and their taxonomic implications. Canad. J. Bot. 63:1957 — 1982. FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill Book Co., Inc., New York. FESTERVAND, D. E 1981. Soil survey of Cape Girardeau, Mississippi, and Scott Counties, Missouri. U.S.D.A. Soil Conservation Service in cooperation with Missouri Agricultural Experiment Station. U.S. Government Printing Office: 1981 — 337-909/ 1020. MARCKS, B.G. 1972. Population studies in North American eats section Laxiglumi (Cyperaceae). Unpublished Ph.D. dissertation, University of Wisc —____. 1974. Preliminary reports on the flora of Wisconsin 767. SipA Book Reviews Botanical Research Institute of Texas S509 Pecan Street Fort Worth, TX 76102-4059, U.S.A. Arpirti, Josepn, Ed. 1990. Orchid Biology, Reviews and Perspec- tions. V. Timber Press, Portland, Oregon. Hardbound $58.00. 450 pp. In this fifth volume, Joseph Arditti continues his excellent task of gathering con- tributions to orchid biology and culture. First, Frits W. Went, discoverer of auxin, writes “Orchids in My Life.” It is an adventure in the history of botany, starting with his auxin discovery, CO, studies, through thermoperiodic studies with mention of many explorations and friends in botany, Chapter 2. The Western Australian Fully Subterranean Orchid Rhizanthella gardneri, by Kingsley W. Dixon, John S. Pate and John Kuo covers history, habitat, biology and morphology, endophyte and nutrition, seed morphology and germina- tion, comparison to other Australian Achlorophyllous orchids, and conservation sug ges- tions. Chapter 3: Water Relations in Orchids, by Russell Sinclair covers Structure and Function, Water Relations of Tissues, Transpiration, Crassulacean Acid Metabolism in Orchids, Epiphyce Distribution Patterns, and Special Cases. Chapter 4: Auto-Pollination in Orchid’s, by Paul M. Catling covers Recognition of Self-Pollination and the Use of Terms, Characteristics of Auto-pollinating Orchids, Methods of Auto-Pollination Degree of Auto-Pollination, Phenotypic and Genotypic Control, Occurrence in the Orchid Family, Geographic Aspects, Evolutionary Implications and Taxonomic Implication. Chapter 5: A review of the Genus Dactylorhiza by Leonid V. Averyanou covers Anatomy and Morphology of Dactylorhiza, flowers, pollen, seeds, chromosomes, taxonomy of Genus Dacty/orhiza, Hybridization, and Natural History of the Genus Dactylorhiza and its species. Chapter 6: Power and Passion: The Orchid in Literature by Martha W. Hoffman Lewis covers orchids in Nineteenth Century England, Orchids in England before World War I, French and German Orchids 1884-1921, Orchids in United States: The Nineteenth Century, Orchids in United States before World War II, and Orchids in Contemporary Literature. Chapter 7: Perspectives of Tropical Orchids In Space Research by Tatyana M. Czerevczenko and Irene V. Kosakovskayoc covers materials and Methods and Results and Discussion. Appendix: Flowering Month of Orchid Species under cultivation by Robert M. Hamilton contains a list of synonyms and their month of flowering months of species under cultivation, a list of 5,594 species. Dottie M. Woodson Bates, Davin M., RicHarp W. ROBINSON, CHARLES JEFFREY. 1990. Biology and Utilization of The Cucurbitaceae. Cornell Univer- sity Press, 124 Roberts Place, Ithaca, New York, 14850. Hardbound. $69.50. 485 pp. — This text covers the interrelations of studies on the biology and utilization of cucurbits. It ts divided into five parts dealing with systematics and evolution, comparative morphol- Oy, Sex expression, utilization and crop improvement and protection. There are 36 chapters by reseachers from around the world. This book is well written and includes an appendix of the classification of the Cucurbitaceae including 118 genera and 825 species. Dottie M. Woodson Stipa 14(4):568. 1991. TYPIFICATION OF VERNONIA TENUIFOLIA SMALL AND V. JAMESII TORREY & GRAY (COMPOSITAE) DAVID E. BOUFFORD Harvard University Herbaria 22 Divinity Avenue Cambridge, MA 02138, U.S.A. ABSTRACT Jones, ina study of the fasciculate group of Vernonia, designated the single collection C, Wright 242 as the lectotype for both V. jamesim Torrey & Gray and V. tennifolia Small. Vernonia jamesti Torrey & Gray is based on the type of V. altissima Nutt. B marginata Torrey and as such is typified by the specimen, Dr. E. P. James “On the Arkansa?,” {sic} cited by Torrey. Although Small cited no specimens in his description of V. tenuifolia in 1898, he did annotate the collection Wright 242 (NY), which can serve as the lectotype for that name. In 1827 John Torrey prepared an account of the botanical specimens collected by Dr. E. PR James during the 1820 expedition to the Rocky Mountains commanded by Major Stephen H. Long. Among the taxa described as new by Torrey was Vernonia altissima B marginata. Torrey ques- tioningly placed this variety under V. a/tissima, a plant that he admittedly had not seen. Rafinesque (1832) raised Torrey’s variety to specific rank as V. marginata (Torrey) Raf., and referred back to Torrey’s original description. ater, Torrey and Gray (1841), without mentioning V. marginata (Torrey) Raf., proposed the name V. jamesti Torrey & Gray. In so doing they placed Torrey’s V. altissima B marginata in synonymy and also gave unmistakable reference to the same collection, “On the Arkansas? Dr. James!,” on which Torrey’s B marginata was based. Since Vernonia marginata (Torrey) Raf. (1832) and V. samesiz Torrey & Gray (1841) are based on the same type specimen the later name, V. pamesti Torrey & Gray, is superfluous. 1898 Small named a new species of Vernonia, V. tenuifolia, from western Texas, but did not cite specimens or designate a type. Gleason (1922) recognized Small’s V. tenuifolia, but again did not designate a type; he merely stated “Type locality: Texas. Distribution: Texas.” Shinners (1950) also failed to typify V. tenwifolia when he reduced it to a variety of V. marginata. Jones (1972; Jones & Faust 1978) placed Vernonia altissima vat. marginata Torrey, V. jamesii Torrey & Gray, and V. fenwifolia Small in synonymy under V. marginata (Torrey) Raf. In his 1972 paper on fasciculate vernonias Jones Sipa 14(4):569—S571. 1991. 570 also designated lectotypes for all three names. He correctly designated the James collection from the Long Expedition as the lectotype for V. a/tissima var. marginata, but incorrectly designated the C, Wright 242 specimen as the lectotype for both V. jamesii and V. tenuifolia. Since Torrey and Gray based their Vernonia jamestt on V. altissima B marginata Torrey, the James collection, the only specimen cited in their description, must also serve as the type for V. jamesiz. According to Jones (1972), however, Torrey later annotated another specimen, Wright 242, as V. gamesi?. This same sheet was later annotated by Small (Jones 1972), as V. tenutfolia. Jones (1972) reasoned that since Small had annotated Wright 242 as V. tenuifolia Small over Torrey’s annotation of the collection as V. jamesti Torrey & Gray that he (Small) was providing another name for V. james. This is, however, not the case since the situation is one of taxonomy and not of nomenclature. As long as Small did not annotate the James collection, the type of V. a/tissima B marginata Torrey, as V. tenuifolia then it cannot be assumed that he was including it in V, tenuifolia. Also, Wright 242 was collected in 1849, 22 years after Torrey named V. altissima B margmata and 17 years after Torrey and Gray’s V. jamesit was published, and Wright's collection could not have been among the material on which those names were based. Furthermore, because Small did not annotate the James collection as Vernonia tenuifolia, that name cannot, as Jones con- cluded (1972), be considered superfluous. The typification for Vernonia altissina Nutt. 8 marginata Torrey, V. marginata (Torrey) Raf., V. jamesti Torrey & Gray, and V. tenuifolia Small, should be as follows: Vernonia altissima Nutt. B marginata Torrey, Ann. Lyceum Nat. Hist. New York 2:210. 1827. — Tee: the specimen collected on Long’s First Expedition, “Dr. James, On the Arkansa” (NY). Vernonia marginata (Torrey) Raf., Atlantic J. 1:146. 1832. — Based on Vernonia altissima Nutt. B marginata Torrey. Vernonia jamesii Torrey & Gray, Fl. N. Amer. 2:58. 184 1, nom. superfl. et illeg. — Based on the same type as Vernonia altissima Nutt. B marginata Torrey. Vernonia tenuifolia Small, Bull. Torrey Bot. Club 25:145. 1898. No specimens cited, typified by C. Wright 242, the specimen at NY bearing the annotation V. tenuifolia in I's) handwriting. — (Lecrory pe (as designated by Jones 1972): NY: possible isoLecroty pes: GH, > 3 sheets.) Vernonia marginata (Torrey) Raf. var. tenuifolia (Small) Shinners. — Based on Vernonia tenuifolia Small. ACKNOWLEDGEMENTS I wish to thank James L. Luteyn for providing information on annota- tions of the James and Wright specimens at NY and for commenting on the manuscript, Norton G. Miller for his discussion on an earlier draft of the manuscript, and two annonymous reviwers for their helpful comments. REFERENCES GLEASON, H. A. 1922. Vernonieae. North Amer. Fl. 33(1):47 — 110. JONES, S. B. 1972. A systematic study of the fasciculate group of Vernonta (Compositae). Brittonia 24:28 — 45. WZ. FAUST. 1978. Compositae tribe Vernonieae. North Amer. FI. I, 10: 180 — 196 RAFINESQUE, C. 8. 1832. Twenty new genera of plants from the Oregon Mountains & c. Aclantic J. 1:144— 146. SHINNERS, L. H. 1950. Notes on Texas Compositae. Field & Lab. 18:25 — SMALL, J. K. 1898. aie in the botany of the southern United States. XIIL. Bull. cee Bot. Club 25:1 15 TORREY, J. 1827. ee account i; a collection of plants made during a journey to and ae dhe Rocky Mountains in the summer of 1820, by Edwin P. James, M. D. Assistant Surgeon U. S. Army. Ann. Lyceum Nat. Hist. New York 2 1oL= 254, & A. GRAY. a ea Fl. N. Amer. 2:57 —59. Stipa Book Reviews Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4059, U.S.A. Cox, PA. AND S.A. BaNack (eds.). 1991. Islands, plants, and Polyne- sians. An introduction to Polynesian Ethnobotany. Dioscorides Press, 9999 S.W. Wilshire, Portland, Oregon. Hardbound. $34.95 plus $3.75 shipping. 228 pp. This interesting book is the Proceedings of a Symposium Sponsored by the Institute of Polynesian Studies, Brigham Young University- -Hawaii Campus Laie, Hawai. Ethnobo- tany is the study of the use of plants by indigenous peoples and in the case of this book the Polynesians. Quoting from chapter 1, by R. Ré aymond Fosberg, “Polynesia includes the islands scattered over a vast triangular area in the Pacific with Hawaii, Easter Island, and New Zealand as its apices, and with a scattering of outlying islands westward into Melanesia and Micronesia. All the islands are tropical or snberopica! except New Zealand and its outlying islands and groups, which are temperate or even ¢ old.” The book has ten chapters and includes and index to scientific names and an index to Polynesian words. Stipa Book Reviews Botanical Research Institute of Texas S09 Pecan Street Fort Worth, TX 76102-4059, U.S.A. Brown, Craup L., L. KarHerine KirKMAN. 1990. Trees of Georgia and Adjacent States. Timber Press, Inc., 9999 S.W., Wilshire, Portland, Oregon 97226. Hardbound. 292 pp. This manual includes summer and winter keys and descriptions of 205 native taxa with ‘ color photographs of leaves, flowers, fruit, and bark and 95 black and white photographs of winter twigs. After a thorough introduction, the Summer Key to genera then family and Species Descriptions are divided by angiosperms and conifers. There is a list of Introduced and Naturalized Species and a glossary. This manual could be used by naturalist, horticulturist and gardeners. Dottie M. Woodson Cater, Marir, JosepH K. Mertzwetter. 1988. The Louisiana Iris. PO. Box 9005, Waco, Texas 76714. Hardbound. 225 pp. Louisiana irises are rhizomatous and beardless. There are five species in Louisiana and surrounding states. This text published by The Society of Louisiana Irises covers Chapter | History of Louisiana Irises, Chapter 2: Classification and Species, Chapter 3: Diversity and Adaptability, Chapter 4: Natural Hybrids and Collecting, Chapter 5: Propagation of Louisiana Irises, Chapter 6: Diploids and Tetraploids, Chapter 7: Hybridizing Louisiana Irises, Chapter 8: Culture of Louisiana Irises, Chapter 9: Diseases and Pests, Chapter 10: Irises Shows, Judging and Awards, Chapter 11: Uses and Other Activities. There is a glossary and many line drawing, painting, and photographs. This book is a must for — — everyone interested in irises. Dottie M. Woodson Oaks, ALBERT. 1990. Ornamental Grasses and Grasslike Plants. Van Nostrand Reingold, New York, N.Y. Hardbound $64.00. 614 pp. An illustrated text on grasses, bamboos, grasslike plants, rushes and sedges and their ornamental uses. Chapter 1: covers types, adaptations, and uses of ornamental grasses with selected species described by botanical name, synonym, common name, origin, habitat, description, uses and cultivation. Chapter 2: describes bamboo; Chapter 3: describes grass- like plant; Chapter 4: describes rushes and sedges with the same format as Chapter 1. There are 5 appendixes for quick reference and a glossary. There are 150 illustrations. This text is a good reference for any one interested in using or identifying grasses. Dottie M. Woodson Stipa 14(4):572. 1991. EUPHORBIA JOHNSTONIT (EUPHORBIACEAE), A NEW SPECIES FROM TAMAULIPAS, MEXICO, WITH NOTES ON EUPHORBIA SUBSECTION ACUTAE MARK MAYFIELD Department of Botany University of Texas at Austin Austin, TX 78713, U.S.A. ABSTRACT Euphorbia johnstonii, a newly recognized species from northern Tamaulipas, is descri- y resembles E. acuta, from which it is distinct in its less — bed and illustrated. It most close upright habit, shorter, wider leaves, shorter, appressed vestiture, and geographical dis- tribution. Euphorbia iasee® is a member of Boissier’s subsection Acwtae, a group of hia which are morphologically intermediate between Eaphorbia subgenus Agaloma nd Euphorbia subg. Chamaesyce. Three species considered here to belong to subsect. Acutae are mapped and compared. RESUMEN Se describe e ilustra Euphorbia johnstonii una especie nueva del norte deTamaulipas. Esta a es semejante a E. acuta, de la cual se distingue por su habito menos erecto, menor tamaho, hojas mas anchas y cortas, indumento adpreso y distribucion geografica diferente. Euphorbia johnstonii es miembro de la subseccion Acwtae de Boissier, un grupo de Exphorbia morfol6gicamente intermedio entre Euphorbia es Agaloma y Euphorbia ibg. Chamaesyce. Se mapean ycomparan las tres especies consideradas dentro la subsecci6n Acutae. Boissier (1862) placed Euphorbia acuta Engelm., E. angusta Engelm. and E. lata Engelm. in his subsection Acawtae of the section Anzsophyllum Roeper. The section Anisophy/lum is now recognized as the genus Chama- esyce S. E Gray by some recent Euphorbia specialists (Webster 1967 Koutnik 1987, 1984; Hassall 1976) or as Euphorbia subgenus Chamaesyce Raf. (Oudejans 1989, Carter 1988, Johnston 1975) by those who prefer a broader concept of Euphorbia. Three synapomor phies cited as evidence of monophylesis in subgenus Chamaesyce are terminal differentiation of the apical meristem with the formation of the first pair of leaves (Hayden 1988), C4 photosynthesis (Webster et al. 1975), and the possession of obvious, nonglandular, interpetiolar stipules (Koutnik 1987). Members of the subsect. Acutae resemble subg. Chamaesyce in their entirely opposite, asymmetrical leaves and four-glanded cyathia but (excluding E. /ata) are Sipa 14(4):573 —579. 1991. 574 aberrant in their C,; photosynthetic pathway (Webster et al.1975) and glandular stipules. Exphorbia johnstonii Mayfield sp.nov., in common with subg. Chamaesyce, has opposite, asymmetrical leaves and four-glanded cyathia, but, like members of the subsect. Acwtae, has linear, glandular stipules and no organized bundle sheath (pers. obs. at X 400 without staining) indicating C, photosynthesis. Within Exphorbia, this combina- tion of characters is unique to the subsect. Acwtae supporting a close relationship between E. johnstonii and these taxa. Cytological evidence suggests a base number of X = 14 for the subsect. Acutae (Urbatsch et al 1975), however E. johnstonii is yet to be counted. Euphorbia lata possesses persistent, interpetiolar stipules and C, photosynthesis and, in spite of a chromosome number based on X = 14 (2n = 28I], Keil 1976), is not part of the subsect. Acwtae as defined above. Therefore, Euphorbia subsect. Acutae includes only those three species here shown to possess glandular stipules and C; photosynthesis, and which, in these features, depart from other members of the subgenus Chamaesyce. Euphorbia johnstonii Mayfield, sp. nov. (Fig. 1). Euphorbiae acutae Engelm. similis sed Bea subprostrato, foliis brevioribus lati- ortbusque, et caulibus vestimenco strigoso diffe Perennial herbs with minute, oe pubescence; stems arching to nearly horizontal or prostrate; vestiture mostly sparse to canescent on young growth, white, trichomes not more than 0.20 mm in length. Roots tuberous, fusiform, to ca. 8 cm long and 1.5—2.0 cm wide, 2—8 cm below the soil surface. Underground stems persistent, produced singly, ca. 1 — 15 cm long, often thickened and branching at ground level to produce | —‘5 aerial stems. Aerial stems articulated, few to numerous, ew aa: from the underground stem apex, to ca. 15 cm long, 0.9 — 1.4 thick, stramineous at maturity; internodes (2-) 4 — 10 (-20) mm ce. oe 2 per node, glandular, caducous (rarely evident), subulate, basally canescent, 0.9— 1.5 mm long. Leaves opposite; petioles brief, usually 0.4 — 0.6 mm long; blades broadly ovate, (3-) 5 — 8 (-13) mm long, (4-) 6-8 (-12) mm wide; abaxially pubescent with evenly-distributed, outcurved trichomes ca. 0.2 mm long, these reaching the margins of the adaxial surface which is otherwise glabrous, or sometimes sparsely beset with similar, though scattered trichomes; base asymmetric, rounded, or less often cordiform; apex obtuse, produced into a shortly acuminate point. Cyathia solitary at the nodes on the distal-most 1/3 to 1/2 of the stems, strigulose, the orifice slightly constricted, ca. 2.0 mm high and 2.3 mm wide just below the glands; peduncles 0.8 — 1.2 mm long; glands 4,sessile, oblong to narrowly elliptic, slightly convex, burgundy to red-brown, 0.4 —0.6 mm in width Dp) Re cS, Mas | uy ‘4 ’ geen ny y “bg SL ph ys / eed SEY FIG. 1. A. Habic (1 cm bar); B— D. Cyathium at various stages of development (1 mm bar, upper ht); B. at onset of female pedicel elongation; C. late in male flowering phase; D. after capsule dehiscence; E—E Seed, ventral and dorsal sides, respectively (1 mm bar, lower middle). Drawn from isotype (Mayfield et al. 762, URY). 576 (in the radial plane of the cyathium), 1.1 — 1.5 mm long (tangential to the rim of the cyathium); appendages basally the same width as the glands, slightly wider at their apex, ochre-white, 0.2 —0.6 mm long, the margins erose to crenulate, rarely deeply parted. Staminate flowers 25 — 35 per cyathium. Pistillate flowers, at anthesis, borne on pedicels ca. 2 mm long, ovary densely appressed-tomentose; styles 3, distinct from the base, ca. 0.5 mm long, bifid for 1/3 — 1/2 their lengths, stigmas as wide as or slightly wider than the styles. Capsules 2.6—2.8 mm long, sparsely appressed- puberulent, pedicels to 6 mm long at dehiscence. Seeds (2.1-) 2.2 — 2.3 (-2.6) mm long, 1.6— 1.7 mm wide, ovate in outline, obscurely 4-angled in Cross-section; seed surface mostly pale with broad, light to dark-brown, obscurely transverse, shallow depressions. Type: MEXICO. Tamautipas: 47 mi (76 km) § of the bridge at ae on the San Fernando Hwy (Mex 97), 29 mi (47 km) N of the jet. with 2 , between the towns of Alfredo V. Bonfil and Pedro J. Mendez, caliche cuesta with d vee loamy soils, elev. 59 m, N 25° 26'25” W 98° 13'22”, 11 Jul 1991, aoe “al. 762 (HOLOTYPE: TEX; tsorypes: MEXU, URV, US). Additional collections examined: MEXICO. Tamautipas: 10 mi E of Abasolo on the road to [La] Pesca, near Los Afejos, 900 ft, caliche upland of the Goliad Cuesta, 6 Feb 1960, pees and} ne ae (TEX); 13 mi E of the Abasolo turnoff on the Santandar Jimenez-Pesca road, 15 Dec 1960, . cei and} Johnston 61406 (TEX); 3 mi of Morales, 19 mi E of the arcane Victe wy on the road to Loreto, calichified ha upland wich short brush and prairie openings on the deeper sand, frequent perennials, Apr 26 1960, [Crutchfield ae i 53406 (TE X); km 164 on Matamoros-Victoria Hwy 1O1, rocky roadside, 7 dominant, 22 Apr 1971, Richardson 1415 (TEX); 48 mi from Reynosa on the San Fernando road, 27 mi from Matamoros-San Fernando Hwy turnoff, brush on caliche spot, 19 Oct 1959, Johnston and Graham 4381 (TEX); 47 mi S of ue on the road to San Fernando, short brush on caliche outcrop, snes perennial herbs, 2 Apr 1960, [Cratchfield and} olan 5333 (TEX). Nuevo Leon: Monterrey Hwy, 65 km : of Nuevo Laredo in hard, sandy marl and silt, 7 Apr 1962, Dominguez M. ae 8243 (TEX). Euphorbia johnstonii most closely resembles E. acuta, from which it is dis- tinguished by its shorter pubescence and shorter, wider leaves. In vestiture, E. johnstoni is nearly identical to E. angusta, a plant strikingly different in ics lance-linear to linear leaves and strong, woody taproots. In the field, the low stature and short, arching (sometimes prostrate) stems of E. johnstonti (Fig. 1) are very different from the longer, decumbent to ascending stems of E. acuta and E. angusta. Euphorbia acuta often has red leaves in the late part of the growing season, a character not observed the either of the other species. Mature seeds offer the best characters for differentiation of these three species. A tabular comparison of morphological differences between the three species is given in Table 377 Taste |. Morphological distinctions between E. johnstonit and its nearest relatives. E. jobnstonit E. acuta E. angusta Vestiture appressed; trichomes — spreading; trichomes fares trichomes 0.3 mm long 0.3 mm long 0.3 mm long Leaf shape Ovate Lance-ovate linear to ee L:W ratio 1.3:1 or less 1.3—4.0:1 greater than 5. Aerial stems arching strongly to decumbent to always ascending; prostrate; up to 15 ascending; 20-35 cm 30-40 cm long : long Seeds shallowly alveolate; smooth; concolorous — obscurely depressions brown, transversely rugose; ridges pale concolorous Cyathium campanulate; ca. 2.0 turbinate; ca. 1.5 funnelform; ca. 1.0 mm wide mm wide mm wide Male flowers 2-02 20-25 5 The new species is allopatric with respect to its nearest congeners (Fig. 2) and is almost exclusively confined to the state of Tamaulipas, Mexico. This was the initial clue to its distinctiveness. The only botanist to have systematically collected in this vicinity was Marshall C. Johnston and associated collectors in the late 1950's and early 1960's. Because his collec- tions account for the majority of exsiccatae, the specific epithet is in recognition of his efforts. Lack of botanical exploration in addition to the ephemeral nature of the above ground parts may partially explain the paucity of specimens of E. johnstonii. Widespread habitat destruction in northern Tamaulipas since the 1950's may also be a factor. No specimens were seen from Texas, even though the nearest collection is within about 25 air miles of the border at San Ignacio in Zapata County, where similar habitat is found. The Rio Grande may provide a natural barrier for this plant which has its center of distribution farther south and east. Future collections will probably extend the range southward in Nuevo Leon and perhaps northwest into Coahuila Euphorbia acuta has the most westerly distribution of the subsect. Acutae with stations reaching northwestern Chihuahua and south-central New Mexico (Fig. 2). It seems to prefer calcareous or gypseous clayey soils of the Chihuahuan Desert. Current collections place the center of E. angusta’s dis- tribution in the limestone uplands of the Edwards Plateau and adjacent Coahuila, but its occurrence far to the south in Coahuila’s Sierra de la Gavia indicates a possible hiatus of collections from Coahuila. Exphorbia johnstoni appears to be restricted to the thornscrub of Tamaulipas, which extends into south Texas, northern Nuevo Leon, and extreme eastern Coahuila. To 579 the north and west, Tamaulipan thornscrub blends into the Chihuahuan Desert and Edwards Plateau vegetation, where the other two species of the subsect. Acwtae occur. The new species can be found in open areas on low, caliche-hills of the Tamaulipan coastal-plain and Rio Grande drainage in calcareous, sandy loam with Asclepias prostrata, Leucophyllum frutescens, Guaiacum ie en Turnera diffusa, Melochia tomentosa, Macrosiph lanuginosa, Heliotropium confertifolium, and Evolvulus sp. ACKNOWLEDGEMENTS I thank Guy Nesom and B. L. Turner for their review of the manuscript and Guy Nesom, who prepared the Latin diagnosis. Luis Hernandez prepa- red the Spanish translation of the abstract. Thanks to Sheila Hayden for her efficient and expert preparation of the illustrations. REFERENCES BOISSIER, P. E. 1862. a (In) DeCandolle’s Prodromus Systematis Universalis Regni Beco pars 15, sect. 2. pp. 3— CARTER 1988. Ree (part 2) } Tribe Ans horbieae. In R.M. Polhill. Flora of copied os Africa. A.A. cema, Rotterdam. S ; 1976. crt and cytotaxonomic evidence for generic delimitation in Australian Baghomicse Aust. J. Bot. 24:633— 640. HAYDEN, W. J. 1988. Ontogeny of the cotyledonary region of Chamaesyce maculata (Euphorbiaceae). Amer. J. Bot. 75:1701— 1713. JOHNSTON, M. C. 1975. Studies of the se species of the Chihuahuan Desert region and adjacent areas. Wrightia 5: 120 — KEIL,- Dy J 1976: Caen eee for ue (Euphorbiaceae) from western 405 — 408. — 88 North America. Madrono 05 — 408 KOUTNIK, D.L. 1987. A taxonomic revision of the Hawaiian species of the genus Chama- esyce Euphorbiaceae. Allertonia 4:331— 388 KOUTNIK, D. L. 1984. ear wie horbinceae) —a newly recognized genus in southern Africa. S. African J. Bot. 3:262 — 264. OUDEJANS, C. H. M. 1989. New names and new combinations in the genus Exphorbia. Phytologia 67:43 — 49. URBATSCH, L. E., J. D. BACON, R. L. HARTMAN, M. C. JOHNSTON, T. H. ATSON, Jr., and G. peace R. 1975. Chromosome numbers for North American nae ee Amer. J. fee 494 — 500. WEBSTER, G. L., W. ROWN a _N. SMITH. 1975. Systematics of photosy- nthetic carbon ate ee in ane Taxon 24:27 —33. WEBSTER, G. L. 1967. The ae Euphorbiaceae in the southeastern United States. J. Arnold Arbor. 48:303 — Stipa Book Reviews Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4059, U.S.A. Upron T. Warrer. 1989. Dendrobium Orchids of Australia. Timber Press, Portland Oregon. Hardbound. 237 pp. The genus Dendrobium is one of the largest in Orchidaceae. In Chapter 1; Walter Upton briefly describes the genus and with text and fine illustrations explains the subgenera and sections. Chapter 2: History of the genus ee in Australia. Chapter 3: Descriptions of Dendrobium species in Australia describes each Australian species by synonyms, descrip- tion chromosome #, habitat, te and brs with illustrations and photographs by Upton. Chapter 4: Natural hybr rring in Australia. ay ae History of hybridiza- ion 3 tion, Chapter 6: Pollination and seed raising. Chapt : Culture Appendix 1: Main months of flowering in native habitat. Appendix 2 eee hybrids using only ‘A seea: lian indigenous parents registered to February 1988. Appendix 3: Dendrobium hybrids includi ing Phalaenopsis and D. schroederiamum in their parentage to March 1987. Appendix Reclassifying the genus Dendrobium. Appendix 5: Authors of plant names and publica- changes of Dendrobinm orchids as proposed oe tion abbreviations. Appendix 6: November 1988. The text and illustrations will be helpful to anyone studying Dendrobium. Dottie M. Woodson Wacner, WARREN L., Derrat R. Hersst, S.H. Sonmer. 1991. Manual of The Flowering Plants of Hawaii Volume I and II. University of Hawaii Press, Bishop Museum Press. Hardbound. 1854 pp. This manual provides keys and descriptions for 146 families, 649 genera and 1, 817 species of native and naturized plants. Eight years and fifty contributors make this study of The Flowering plants of The Hawaiian Archipelago the most complete since Hillebrand’s a century ago. The first volume includes chapters on Methods and Scope, Geology, Climate, Vegetation, Important Collections, and Abbreviations. The listings are a eran cally arranged within dicots and monocots. The glossary, literature cited, ‘lusecanion vouchers, and index are in volume I. This Fiaeual would be excellent s any studying plants and plant distributions as well as any One wanting to grow plants from these islands. Dottie M. Woodson SIDA 14(4):586. 199 | REVISION OF THE GENUS CINNA (POACEAE) DAVID M. BRANDENBURG! and WILL H. BLACKWELL’ Department of Botany, Miami University Oxted, OF 2090, US A JOHN W. THIERET Department of Biological Sciences, Northern Kentucky University Highland Heights, KY 41076, U.S. A ABSTRACT Variational patterns and discontinuities were studied in Cina (Poaceae). Nearly 4000 herbarium specimens of the genus were examined; four species were cae on the basis of features of the spikelets. Crmua arundinacea inhabits moist forests in eastern North America; C. /atifolia occupies similar sites in circumboreal regions; C. a occurs in mountainous areas from Mexico to Venezuela and Bolivia; and C. bolanderi i is a central montane Californian endemic previously considered conspecific with C. /atefolia. Keys, descriptions, illustrations, and distribution maps are provided for all species. on Cinna L. is a small but widely distributed genus of perennial grasses. It was originally described by Linnaeus (1753), who recognized C. arundi- nacea L., mostly a woodland species of eastern North America. Subsequent authors referred this species to Agrostis and Muahlenbergia, both of which differ from Cinna by several characters. Fernald and Griscom (1935) descri- bed C. arundinacea vat. inexpansa Fern. & Grisc. as a southern Coastal Plain native supposedly differing from typical C. arundinacea by its more ascend- ing panicle branches and its smaller spikelets. A second species, the circumboreal C. /atifolia (Trevir. ex Gopp.) Griseb. in Ledeb., was first described in 1830 as a species of Agrostis, A. latifolia Trevir. ex Gépp., by Treviranus (Géppert 1830). Trinius (Bongard 1833), however, considered the taxon to be a species of Muhlenbergia, M. pendula Trin. in Bong.; later he (Trinius 1841) transferred it to Canna as C. pendula (Trin. in Bong.) Trin. The combination Cinna latifolia was made by Grisebach (Ledebour 1853). For several decades the epithet pendula — not latifolia — was misapplied to the species. Gray (1856) treated the taxon as C. arundinacea var. pendula A. Gray. Other authors named several varieties ‘Current address: The Dawes Arboretum, 7770 Jacksontown Road S.E., Newark, OH 43055. ‘To whom reprint requests should be sent. Sipa 14(4):581—596. 1991 582 of C. pendula, and it was not until the late 19th century that the earlier specific epithet, /atifolia, came into widespread use. A third member of the genus, C. poaeformrs (H.B.K.) Scribn. & Merr., is a montane species ranging from Mexico south to Venezuela and Bolivia. Originally described by Humboldt, Bonpland, and Kunth (1815) as Deyeuxia poacformis H.B.K., it has also been treated as belonging to Calamagrostis or Poa. The combination Cinna poaeformis was eventually made by Scribner and Merrill (1901). Cinnastrum of Fournier ( 1886), based on Deyewxia poaeformis, was established apparently because of Linnaeus’ failure, in the original description of Cinna, to note the prolonged rachilla. Fournier characterized Crnnastrum by “spiculus bifloris, lore superiore ad pedi- cellum sterile redacto.” A collection of Czyna from central California was recognized by Scribner (1884) as distinct from C. latifolia, he named it C. bolanderi Scribn. in honor of H.N. Bolander, who collected it in 1866. The species was later reduced to the synonymy of C. /atifolia by Hitchcock (1935). We are rein- stating it to specific rank; it is quite distinct from C. latifolia (see discus- sion under C. bolandert). Behrens (1877) gave a brief account of nervature of the palea of C. arundinacea and C. latifolia, Brandenburg et al. (n.d.) more fully described that of C. arundinacea. Chase (191 1a) published a short paper on subter- ranean organs of “Cynna ead she later discovered that the specimens she studied were Arrhenatherum elatius (L.) Pres] (Chase 191 1b). Stérmer (1949) observed that just Below the glumes of Cinna is a slight collar that, upon disarticulation of the spikelets from the plant, persists as a small cupule at the apex of the pedicel; he regarded this cupule to be of diagnostic value in differentiating from Cinna those Norwegian grasses likely to be confused with it. Several writers have noted that grains of Cinna contain semt-liquid endosperm (Dore 1956; Martin 1946; Terrell 1971). Cina Clearly belongs to the subfamily Pooideae on the basis of spikelet morphology, in conjunction with evidence obtained from root-hair development (Row and Reeder 1957), features of the embryo (Reeder 1957), leaf anatomy (Brown 1958), and chromosome number (Bowden 1960; Davidse and Pohl 1978). Hitchcock (1950) included Cinna as a member of the Agrostideae, a tribe not recognized by most later authors (e.g., Stebbins and Crampton 1961), who assigned the genus to Aveneae. Recently, Macfarlane and Watson (1982) reexamined the relationship between these two tribes; in their final analysis Cinna is placed in a recir- cumscribed Agrostideae. However, Clayton and Renvoize (1986) classified Cima in tribe Aveneae, subtribe Alopecurinae, the latter consisting of grasses with ° ‘spikelets I-flowered; glumes commonly enclosing florets; palea not gaping. TAXONOMY We define Cinna as having the following generic characters:(1) spikelets 1-flowered, (2) disarticulation below the glumes,(3) awn (when present) subterminal, (4) rachilla typically prolonged behind the palea as a small glabrous or scaberulous stub or bristle, and (5) palea 1-nerved, or 2-nerved and the nerves very close together. Two characters traditionally employed to define the genus should be mentioned. The first of these, the prolonged rachilla, varies from a minute stub in C. arundinacea to a slender bristle half the length of the lemma in C. poaeformis. However, this prolongation is often difficult to detect (especially in C. arundinacea), occasionally absent, and is therefore a poor character to use as the principal basis for identifying Czana in a generic key (cf. Hitchcock 1950). The second feature associated with the genus ts the pres- ence of monandrous flowers. Although this holds true for C. arundinacea and C. latifolia, C. poaeformis and C. bolanderi have diandrous flowers. Because there is such a clear-cut distinction among the species of Czzna on this basis, there is a temptation to recognize two sections within the genus (especially when one considers that C. arundinacea and C. latifolia also have stipitate florets and 3-nerved lemmas, while C. poaeformis and C. bolandert have more or less sessile florets and normally 5-nerved lemmas). However, even though C. bolanderi agrees with C. poaeformis in these characters, on the basis of macromorphology it is markedly similar to the other two species rather than to C. poaeformis. Also, lemmas of both C. arundinacea and C. Jatifolia on rare occasion have five nerves. Hence we decided against subdivision. This revision is based on the study of about 4000 sheets of Czmna. To save space, specimens are not cited; such citations are given in Brandenburg (1980). on CINNA L., Sp. Pl. 1:5. 1753; Gen. Pl. 1:6. 1754. Abola Adans., Fam. Pl. 2:31, 511. 1763. Based on Cina L. Blyitia Fries, Novit. Fl. Suec. Mant. Alt. 2:2. 1839 (fide Berg 1966). Cinnastrum Fourn., Mex. Pl. 2:90. 1886. Based on Deyeuxta poaeformis H.B.K. Tall perennials with solitary or cespitose culms, sometimes tinged with purple. Nodes and internodes glabrous or rarely somewhat scaberulous. Blades flat, the margins scabrous; adaxial and abaxial surfaces scabrous or smooth. Ligules scarious. Sheaths open, glabrous. Panicle branches spreading or ascending, the axis and pedicels scabrous to smooth. Spikelets aterally compressed, 1-flowered or very rarely with a second rudimentary or fertile floret above the first. Disarticulation below the glumes. Rachilla prolonged behind the palea as a minute stub or slender bristle, this smooth — or scaberulous at tip, occasionally absent. Glumes acute, I- or 3-nerved, 584 sometimes minutely awn-tipped; keel upwardly scabrous, body smooth or occasionally scaberulous, margins hyaline; first glume somewhat shorter than or equalling the second. Floret sessile or stipitate. Lemma similar to glumes, 3- or 5-nerved (the nerves parallel and often faint or obscure), with a short, straight, upwardly scabrous awn just below the apex (mostly awn- less in C. poaeformis, sometimes awnless in the other species). Palea hyaline, mostly smooth, I-nerved, or 2-nerved and the nerves very close together, upwardly scaberulous along the keel(s). Grain yellowish-brown, often beaked by the persistent style. Stamens | or 2. x = Tyee species: Cinna arundinacea L. KEY TO THE SPECIES OF CINNA 1. First glume 3-nerved; spikelets + obtuse .................... 4. C. poaeformis |. Firse glume 1-nerved; spikelets acute. Stamens 2; lemma 5-nerved; florer + sessile ................ 5 Stamen |; lemma mostly 3-nerved; floret raised on a 0.1 —0.65 mm 3. C. bolanderi Dh NO Stipe 3. Second glume prominently 3-nerved; spikelets typically 4—6 mm C. arundinacea 3. Second glume I-nerved (very rarely 3-nerved); spikelets typically LEELA GESTURES Dire EEE eS Mc MON — 2.5—4 mm in length Spikelets of the four species of Cinna are shown in Fig. 1; diagnostic features of the species, in Table 1. Taste 1. Diagnostic features of the species of Cinna. C. arundinacea C. latifolta C. bolanderi C. poaeformis Length of sptkelets, mm* (3.5)4 — 6(7.5) (2)2.5 — 4(5) (3.6)4 — 5.5(6.3) 1.9— 3(3.5) Number of nerves on glume | | I l 3 Number of nerves on glume 2* 3 1(3) lor 3 3 Floret stipitace or sessile stipitate stipitate + sessile + sessile Number of nerves on lemma* 305) 4(5) 5 > Number of stamens l | 2 2 Anther length, 0.8— 1.9 0.4—1.1 1.2—2.6 0.5-—1.2 *The word “usually” should be understood here. 585 Cinna arundinacea L., Sp. Pl. 1:5. 1753. — (Puororype: Linnaean Herbarium, IDC No. S-3-8! We designate this specimen as the lectotype, as a material, IDC No.S-3-6!, also exists.) — Type Loca- _ second specimen of origina Liry: C DA Agrostis cinna Lam., Tabl. Encycl. 1:162. 1791. Based on Cinna arundinacea L. Agrostis cinna Pursh, Fl. Amer. Sept. 1:64. 1814. Based on Cinna arundinacea L. Cinna agrostidea Beauv. ex Steud., Nom. Bot. 1:20, 198. 1821. Based on Agrostis cinna Lam. (C. “agrostoides’ according to Hitchcock). Muhlenbergia cinna Trin., Gram H. 191. 1824. Based on Agrostis cnna Lam. Cinna arundinacea L. var. inexpansa Fern. & Grisc., Rhodora 37:135, pl. 334, fig. 1 1935 Plants 2.8— 18.3 dm tall, somewhat bulbous at base. Nodes 5 — 13. Blades to 34.5 cm long, 3 — 19 mm wide. Ligule 2— 10 mm long. Panicle green, gray-green, or purplish, 6.5—55 cm long, 1-22 cm broad, loosely to densely flowered; branches ascending to spreading. Spikelets acute, (3.5)4— 6(7.5) mm long. Floret raised on a 0.25 — 0.65 mm stipe. First glume somewhat shorter than lemma, l-nerved, (2.7)3.5 — 5(6. 1) mm long; second glume equal to or slightly longer than lemma, 3-nerved, (3.5)4— 6(7.5) mm long. Lemma 3-nerved, occasionally with an addition- al nerve along one or both sides, (2.7)3.5 — 5(6.4) mm long; awn 0.2 — 1.5 mm long, rarely absent. Palea 1-nerved, 2.4—4.6 mm long. Grain 2.1—2.8 mm long. Prolonged rachilla 0.1 —0.4 mm long, sometimes absent. Stamen 1, anther 0.8 — 1.9 mm long. 27 = 28 (Bowden 1960), 40 (Avdulov 1928, fide Fedorov 1969. The report of 2” = 40 ts suspicious because all other available counts for Cina are on a base number of x = 7). Fig. General range: eastern North America (Fig. 2A). Habitat: most commonly found in moist woodlands, in swamps, along streams, and in upland woods, less commonly in wet meadows, marshes, and waste ground and along roadsides; elevation ca. 0 — 850 m. Flowering and fruiting time: late summer and fall. Discussion. Cinna arundinacea may be distinguished from C. latifolia primarily by its strongly 3-nerved second glume and secondarily by its lar- ger spikelets. Inflorescence characters commonly employed in floras — C. arundinacea. panicle dense, the branches ascending versus C. latifolia: pan- are not reliable, as it 1s not un- icle loose, the branches spreading common for C. arundinacea to have very open panicles and drooping branches. Two collections seen of C. arundinacea have not been mapped. The first 1s an August 1890 collection by Sandberg (PENN 25045) labeled “Isanti Co., Idaho.” There is no Isanti County in Idaho, a state west of the range of 0.5 —20 cm broad, glume (1.8)2.5 —4(4 586 the species (the specimen may have come from Minnesota, where there /s an Isanti County). The second sheet (She/don 268, MU), labeled “Deschampsia caespitosa, “1s from Clear Creek Co., Colorado. As this state is also west of the range of C. arundinacea, it is piorcble that somehow a mix-up of label data occurre Cinna wae was attributed to Montana and northern North Dakota by McGregor et al. (1977). The voucher specimen (Stephens 67806, KANU) for the Montana report is a species of Calamagrostis. We were un- able to locate any voucher for the North Dakota report. 2. Cinna latifolia (Trevir. ex Gépp.) Griseb. in Ledeb., Fl.Ross. 4:435. — Tree tocauiry: EUROPE. Agrostis latifolia Trevie ex Gopp., Beschr. Bot. ‘Gait: Breslau 82. 1830. Sct. Math. bia oes Trin. in Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Zee. 33. Cinna expansa Link, Hort. Berol. 2:236. 1833. Agrostis suaveolens Blytt ex Sommerf., Kongl. | ae es 1837:256. 1838. a Suaveolens Fries, Novit. Fl. Suec. Mant. 2:2. 1839. Based on oo suaveolens Blyte ex Sommerf. Cinna cient Rope eX a FI. 1853. Based on oe suaveolens Bly Cinna peidile _ a Be St. pes Sér. 6, Sci. Nat. 4:28 134 1. The earlier ies pendula Trin. not me anaes L. var. pendula A. Gray, M (Trt n Bong.) Trin. Cina on (Trin. in Bong.) Trin. var. saan Scribn., Proc. Acad. Nat. Sci. Phila. 884:290. 1884. (Lecrorype here ignated: Gedy 664, US!) pies pendula (Trin. in Bong.) Trin. var. A alan Macoun, Cat. Ca Pl. 2(V):393. 1890. Epithet ascribed to “Scribn.”; error for var. glomerula Scribn. Cinna pendila (Trin. in Bong.) Trin. var. acutiflora Vase sey ex Macoun, Cat. Canad. PI. 2(1V):203. 1888, nom. nud.; then, in the same Macoun work (p. 393. 1890), C. pendula var. acutiflora was soblicned as a synonym of C, penale ¥ var. omer ‘la Scribn. (glomerata’). (Lecvory pr here designated: Macoun ee US!; IsoLec Cinna pendula (Trin. in Bong. ) Trin. var. muti 202, 1888, nom. nud.; name validly ublished in Contr. U.S. Natl. Hed, 3:57. 1892. (Lecrorype here designated: Cusick s.n., US!: isolectotype: NY!). Cinna lee (Trevir. ex GoOpp.) Griseb. in Ledeb. var. glomerata Beal, Grasses N. Am 319. 1896. Epithet ascribed to “Scribn.”; error for var. glomerula Scribn. Plants 2— 19 dm tall. Nodes 4 —9. Blades to 28 cm long, |— 20 mm wide. Ligule 2—8 mm long. Panicle green or purplish, 3 —46 cm long, loosely to densely flowered; branches spreading or sometimes ascending. Spikelets acute, (2)2.5—4(5) mm long. Floret raised on 0.1 —0.45 mm stipe. Glumes + equal, longer than to shorter than lemma, each I-nerved (second glume very rarely 3-nerved); first .7) mm long; second glume (1.9)2.5 —4(5) mm long. Lemma 3-nerved (rarely faintly 5-nerved), the lateral nerves often Ross. 4:435. rin., Mém. Imp. yned. Cinna an. ed. 2. 545. 1856. Based on meee ~~ in ne on ee FIG. 1. Spikelets of Cinna. A. C. arundinacea. B.C. latifolia. C. C. bolandert. D. C. poaeformis. 588 FIG. 2. Generalized range of Cinna arundinacea. C. latifolia (New World), solid line. Cinna bolanderi, California, triangle 589 PIG. 3. Generalized range of Cinna latifolia (Old World). Adapted from Hultén and Fries 1986. obscure, 1.8 — 3.8 mm long; awn 0.1 — 2.5 mm oie or absent. Palea 2- nerved, the 1 nerves very close together, or l-nerved, 1.8 —3.4 mm long. Grain 1.8 — 2.8 mm long. Prolonged rachilla slender, . 3 molone: sometimes sap Stamen |, anther 0.4 — 1.1 mm long. 2” = 28 (Bow- den 1960; Ehrenberg 1945). Fig. 1B. General range: circumboreal (Fig. 2B, 3). Habitat: moist to wet soil in woodlands, swamps, thickets, bogs, and streamsides; elevation ca. 0 — 2600 m. Flowering and fruiting time: late summer and fall Discusston: Morphologically, this 1s the most ere le of the species of Cinna, a fact doubtless accounting for the several varietal names proposed. There are no consistent differences between the Eurasian plants and those from the New World. A collection from the Aleutian Islands (Eyerdam 1791, UC, US) is peculiar with its abnormally large (to 5.5 mm) and often 2-flowered spikelets. These measurements are not included in the above description of C. latifolia. Cinna latifolia was attributed to northeastern Montana and northwestern North Dakota by McGregor et al. (1977). We were unable to locate any vouchers to verify this report. 590 3. Cinna bolanderi Scribn., Proc. Acad. Nat. Sci. Philadephia 1884:290. 1884 ee TOTYPE here . Bolander 6090, US(#323939)!; Isotecrorypes: DS! GH! MO! NY! US!) — Tyee central montane California. Cinna pendula var. Sci ) Vasey, Contr. U.S Natl. Herb. 3:57. 1892. Based on Cinna bolanderi Scri Plants 8.5 — 20.3 dm tall. Nodes 4—8. Blades to 40 cm long, 2— 19 mm wide. Ligule 3.5—7 mm long. Panicle green to golden green 7.5—43 cm long, 3— 18 cm broad, loosely to densely flowered; branches spreading to loosely ascending. Spikelets acute, (3.6)4—5.5(6.3) mm long. Floret + sessile. First glume longer than to shorter than lemma, l-nerved, (3.3)3.5 —5.2(6) mm long; second glume longer than or equal to lemma, I- or 3-nerved, (3.6)4 —5.5(6.3) mm long. Lemma 5-nerved, the lateral nerves sometimes faint or obscure, (2.7)3.2 —4.6 mm long; awn 0.2—1.5 mm long or absent. Palea 2-nerved (the nerves approxi- mate), (2.7)3 —3.5(3.8) mm long. Grain 2—2.9 mm long. Prolonged rachilla slender, 0.4—0.9 mm long, sometimes absent. Stamens 2, anthers 1.2 — 2.6 mm long (rarely at a da and to 0.7 mm long). No chromosome number available. — = General range: central montane a. (Pig, 2c). Habitat: meadows and streamsides; elevation ca. 1900 — 2400 m. Flowering and fruiting time: lace summer and fall. Duscussion: Cinna bolanderi is endemic to central montane California (Fresno, Mariposa, and Tulare counties); all collections we have seen are from Sequoia National Park, Kings Canyon National Park, and the southern tip of Yosemite National Park. Cryna latifolia is more northern in range in California, the southernmost record being from just north of Yosemite National Park. Cznna bolanderi, with its two stamens, is easily distinguished from the monandrous C. /atifolia. Another noticeable differ- ence is the length of the anthers, those of C. bolandert being much the lar- ger. Diagnostic characters for fruiting specimens of C. bolander? are the lar- ger spikelets, the sessile florets, and the 5-nerved lemmas (though the nerves are often faint). 4. Cinna poaeformis (H.B.K) Scribn. & Merr., Bull. U.S.D.A., Div. Agrost. 24:21. 1901. — Tyee tocatrry: MEXICO. Deyenxta poaeformis H.B.K., Nov. Gen. Sp. 1:146. 1815. (Type: P!). Poa subuniflora Kunth, Révis. Gram. 115. 1829. Based on Ape gti neh #8 H.B.K ex Sceud., Nom. Bot. ed. 2. 251 . Based on Poa pe ia nth. Cranastram poaeforme CH.B.K) 5: Foun. Mes, “a ei 1886. Based on Dense poaeformis BLK. Calamagrostis poaeformis (Fourn.) Beal, Grasses N Calamagrostis poaeoides Trin. Based on Cinnastrum poaeforme Fourn. Cinnastrum milraceum Fourn., Mex. Pl. 2:91. 1886. 591 FIG. 4. Documented distribution of Cinna poaeformis. Plants 2.8— 22.2 dm tall. Nodes 4—9(11). Blades to 33 cm long, 1— 18 mm wide. Ligule prominent, (1)5 — 12 mm long. Panicle green or purplish, 7—47 cm long, 1—18 cm broad, many-flowered; branches spreading. Spikelets + obtuse, 1.9—3(3.5) mm long. Floret + sessile. Glumes thick in texture, + equal, longer than to shorter than lemma, 592 each 3-nerved; firsc glume 1.8—3(3.4) mm _ long; second glume 1.9 — 3(3.5) mm long. Lemma thick in texture but tearing easily length- wise, S-nerved (one or both pairs of lateral nerves often obscure), (1.6)2 — 2.8(3.4) mm long; awn normally absent, to 0.4 mm long when present. Palea + thick in texture, tearing easily lengthwise, 2-nerved (the nerves approximate), (1)1.7 — 2.4(2.9) mm long. Grain 1.3— 1.8 mm long. Prolonged rachilla slender, 0.3 — 2 mm long, rarely absent. Stamens 2, anthers 0.5 — 1.2 mm long. 2” = 28 (Davidse & Pohl 1978; Pohl & Davidse 1971). Fig. 1D. General range. Mexico south to Venezuela and Bolivia (Fig. 4). Habitat. mountains, in moist or dry soil of woods, meadows, and paramos; elevation ca. 2200 — 4000 m Flowering and fruiting time: late summer and late fall in Mexico; July through May farther south. Discussion: Cinna poaeformis is the most dissimilar in outward appearance among the four species in the genus. Its spikelets are small, more or less obtuse, and normally awnless; they have a prominent prolonged rachilla. However, the species agrees wholly with the characters used to circum- scribe the genus. DOUBTFUL AND EXCLUDED SPECIES Agrostis Cinna — , Observ. Bot. 5:18. 1789. Originally as synonym for Cinna arundinacea .., but 2 years later Retzius i Bot. 6:22. 1791) concluded that his A. cinna was really a species of the genus now known as Muahlenbergia. sitet poaeformis (Fourn.) M.E. Jones, Contr. West. Bot. 14:9. 1912. Based on Cinnas- n poaeforme Fourn. as to name but not as to description. Cinna? alba Nees ex Steud., Syn. Pl. Glum. 182. 1855. (Tyee: ie me Medoza. In US! is asheet marked “ Type” that has but one spikelet ina packet. The lemma has a long, ved awn and callus hairs, which does not agree with oe pera “Hosculi ia inferiore nuttica acuta. Cinna ples Kunth, Révis. Gramin. 1:67. 1829 = Muhlenbergia expansa (DC.) Trin., fide , Man. Grasses U.S. 900. 1950. Based on Agrostis arachnoidea Poit. Cinna as a ee , Fl. Bor. Amer. 2:238. 1840 (non L., 1753). Listed in Index , but no ne combination made by pe Cina Cee Retz. ex Steud., Nom. Bor. ed. 2. 1:365. 1841 (non L., 1753) = Mu piel mexicana (L. ) Trin., fae Hitchcock, -. Grasses U.S. 903. 1950. As onym of Cinna mexicana Beauy Cinna ey wit Rupr., Beitr. nzen Rass, Reich. 2:66. 1845 = Arctagrostis latifolia (R. 3r.) Griseb. in Ledeb "hide Nash, N. Amer. Fl. 17:498. 1937. Based on Colpodtum latifolinm R. Br. Cinna yet Trin., Fund. Agrosct. 118. 182(¢ ae. L., which = Dre . Species 1:202. 1962. Cinna decipiens Kunth, Révis. Gramin. 1:67. 1829. Based on Agrostis decipiens R. Br. (Vilfa 3 Beauv.) = Deyeuxita decipiens (R. Br.) Vickery, Contr. New South Wales Natl. Herb. 1:70. 1940. ~~ OS = Di eee crintta (L.) Hook. Based on / ; _ fide Chase and Niles, Index to 295 Cinna filiformis (Willd.) Link, Enum. Pl. 1:70. 1821 = inion mexicana (L.) Trin fide Hitchcock, Man. Grasses U.S. 903. 1950. Based on Agrostis filiformis Willd. iii le eee Hitchc., fide Secon, Contr. U.S. Natl. Herb. 34:148. 1967. Based on Agrostis lanata H.B.K. Cinna pba Walt., Fl. Carol. 59. 1788 = Andropogon virginicus L., fide Hitchcock, Man. Grasses U.S. 817. 1950. Cinna ee (Michx.) Kunth, Révis. Gramin. 1:67. 1829 = Muhlenbergia frondosa (Poir.) Fern. , fide Hitchcock, Man. Grasses U.S. 901. 1950. Based on Agrostis lateri- flora Michx. Cinna macroura (H.B.K) Kunth, Reévis. ieee 1:67. 1829 = ee macroura H.B.K.) Hitchcock, fide N bea . Amer. Fl. 17:468. 1935. ed on Crypsis macroura H.B.K. (“C. macronra Kunth” misapplied by Thurber in Wats., aoe Calif. 2:276. 1880 to bahia rigens (Benth.) Hitchcock). Cinna? scan ) Beauv., Ess. Agrost. 32, 148, 158. 1812 = Mable mexicana (L.) n., fide Hitchcock, Man. Grasses U.S. 903. 1950. Based on Agrostis mexicana L. a mexicana (L.) Link, Enum. Pl. 1:70. 1821 = Muahlenbergia mexicana (L. ‘Thin: Based Agrostis mexicana L., which = Muahlenbergia mexicana, fide Hitchcock, Man asses U.S. 903. 1950. Cinna pos Kunth, Révis. Gramin. 1:67. 1829 = Echimopogon sp. Kunth Lists in ey ostis fe Forst., Echinopogon ovatus Beauv., Echinopogon asper Cinna Hho (H.B.K. Cane h, Révis. Gramin. 1:67. 1829. Based on Co pilieiies = Mublenberia aff. | oe TR. Soderstrom per Cinna? Le (H.B.K. Ré 57. 1829. Mublenbergia pubescens B.K.) Eizch cock, - oe on 1S: a Herb. 34:148. 1967. paced on Agrostis Jeg H Cinna? purshit Kunth, Révis. Gramin. 1:67. 829 = Calamagrostis canadensis (Michx.) Beauv. fide Hitchcock, Man. Grasses U. : 839. 1950. Based on Arundo agrostoides ursh. Cinna racemosa (Michx.) Kunth, Révis. Gramin. 1:67. 1829 = Muhlenbergia racemosa (Michx.) B.S.P., fide Hitchcock, Man. Grasses U.S. 904. 1950. Based on Agrostis racemosa Michx Cinna setifolia (Presl) Kunth, Révis. Gramin. suppl. XVI. 1830 = Muahlenbergia macroura 594 4.B.K. : is fide Nash, N. Amer. Fl. 17:468. 1935. Based on Crypszs set- oi @ Pre Cinna oe oie ) Link, Enum. Pl. 1:71. 1821 = Muahlenbergia sobolifera (Muhl.) ide Hitchcock, Man. Grasses U.S. 906. 1950. Based on Agrostis sobolifera Willd. Cinna? stricta (Humb. & Kunth) Kunth, Révis. Gramin. 1:67. 1829 = Muhlenberg ta Aes (Presl) Kunth, fide Chase and Niles, Index to Grass Species 1:507. 1¢ I n Crypsis stricta Humb. & pow Cinna oni (Willd.) Link, Enum. PI. O. 1821 = Muahlenbergia tenuiflora (Willd.) Saad Hitchcock, Man. Grasses U.S. ae 1950. Based on Agrostis tenuiflora z= Cinna ae wana Phil., Anal. Univ. Chile 1873:563. 1873. From description not a Cinna: 0 brevi pilosa’ 5 “pedicellus pilosus rudimentum secundi floris sistens tertiam partem paleae inf vioris aequat’, lower glume “wnivervia, vix dimidiam paleam aequante is eas Griseb., Abh. Kénigl. Ges. Wiss. Gottingen 19:208 — 209, fig. 7. 1874. Incorrectly listed as synonymous with Cinna L. by Willis (1973). Grisebach’s account of a grass with unisexual spikelets, hairs on the rachilla, and articulation above the glumes is descriptive of a genus other than Cinna. Muahlenbergia baicalensis Trin. ex Turez., Bull. Soc. Imp. Naturalistes Moscou 24(1):21. 1856. Published as synonym of Cinna latifolia (Trevir. ex Gépp.) Griseb. in Ledeb. ACKNOWLEDGEMENTS We express appreciation to Robert Brandenburg for his fine line drawings; to Dr. James R. Estes, Dr. Thomas R. Soderstrom, and Dr. John L. Vankat for comments; and to the curators of the following herbaria for the loan of specimens: A, ALU, ASC, CO U, NCU, B, OKLA, PENN, PH, POM, RSA, SDU, SMU, TENN, TEX, UC, UNA, US, VDB, VEN, WVA, YU. REFERENCES BEHRENS, a 1877. Notiz zur Kenntnis der Gramineen Blithe. Bot. Zeitung (Berlin) S20 = BERG, R.Y. ok On the discovery and distribution of Cinna latifolia in Norway, with remarks on ecology and migration. I. Blyttia 24:145 — 160. [In Norwegian, with Engl- a H.G. 1833. Observations sur la végétation de l'tle de Sitcha. Mém. Acad. Imp. Sci. Sc. Petersbourg, Sér. 6, Sci. Math. 2 BOWDEN, W.M. 1960. Chromosome epee and taxonomic notes on northern grasses. II. Twenty-five genera. Canad. J. Bot. 38:541 BRANDENBURG, D.M. 1980. A synopsis of the genus Crna (Gramineae). Master's thesis, Miami aeons Oxford, Ohi R ES, $.D. RUSSELL, and J.W. anne n. d. One- nenves paleas in Sie arundinacea L. (Poaceae). Trans. Kentucky. Acad. BROWN, W.V. 1958. Leaf anatomy in grass systematics. Bot. — "(Crawfordsvil le) 119:170— 178. 595 CHASE, A. 191 la. Subterranean organs of Cinna arundinacea. Rhodora 13:9 — 10, 19 Arrhenatherum elatius tuberosum in America. ae 13:207 — 208. CLAYTON, W.D. and S.A. RENVOIZE. 1986. Genera graminum. Grasses of the world. Kew Bulletin Additional Series XH. Her Majesty’s —— Office, London. DAVIDSE, G. and R.W. POHL. 1978. Chromosome numbers of tropical American grasses (Gramineae): 5. Ann. Missouri Bot. Gard. aie 19. DORE, W.G. a4 Some grass genera with liguid endosperm. Bull. Torrey Bot. Club 83:335— a re L. 1945. Kromosomralen hos nagra Karlvaxter. Bot. Not. 1945:430 — 437. FEDOROV, A.A. (ed.). 1969. Chromosome numbers of flowering plants. Academy of Sciences of the U.S.S.R., V.L. Komarov Botanical Institute, Leningra FERNALD, M.L. and L. GRISCOM. 1935. Cinna arundinacea L., vat. inexpansa, vat. nov. See 37:135, pl. 334 FOURNIER, E. 1886. Meacanes plantas, pars secunda, Gramineae. Ex Typographeo Republi, aris GOPPERT, H.R. 1830. Decca ian = Botanischen Gartens der Konighchen Univer- sitat ee Josef Mar und Komp. eslav GRAY, A. 1856. Manual of the botany i - conte United States (2nd ed.). George P. Putnam and Co., New York. HITCHC A.S. 1935. Manual of the grasses of the United States. U.S.D.A. Misc. Publ. 1950; Seg of the . of the United States (2nd ed., revised by Agnes Chase). U.S.D.A. Misc. Publ. HULTEN, E. and M. eer 1986. a as . on European vascular plants north of the Tropic of Cancer I. Koeltz Scientific Books, Koenigstein, eS Republic of Germany. HUMBOLDT, A. de, A. BONPLAND, ae C.S. KUNTH. 1815. Nova genera et species plantarum, vol. 1. Sumtibus Librariae enn er SG a Lutetiae Partsi- orum. KOVALEVSKAYA, S:S. (ed.). 1968. a rastenit srednei Azii. "FAN,” Tashkent. {In rae Pes karataviensis, p. 98} LEDEBOUR, C.E 1853. Flora Rossica, vol. 4. Sumtibus Librariae E. Schweizerbart, No) LINNAEUS, C. 1753. Species plantarum. Laurentit Salvi, Holmiae. MACFARLANE, T. and L. oe ae 1982. The classification of Poaceae subfamily Pooideae. Taxon 31:178 — MARTIN, A.C. 1946. The « comparative internal morphology of seeds. Amer. Midl. Naturalist 36:5 13 — 660. McGREGOR, R.L., et al. 1977. Atlas of the flora of the Great Plains. Iowa State Univer- sity Press, Ames. PAVLOV, N.V. 1956. Flora Kazakhstana, vol. 1. Akademiya Nauk Kazakhskoi SSR, Alma-Ama {sic}. [In Russian} [Civna, p. 176-177] 1958. Flora Kazakhstana, vol. 2. Akademiya Nauk Kazakhskoi SSR, Alma-ata. [In Russian} [Cinna karataviensis, p. 277] POHL, R.W. and G. DAVIDSE. 1971. Chromosome numbers of Costa Rican grasses. Brittonia 23:293 — 324. REEDER, J.R. 1957. The embryo in grass systematics. Amer. J. Bot. 44:756— 768 ROW, H.C. and J.R. REEDER. 1957. Root-hair development as evidence of relationships 596 among genera of agen Amer. J. Bot. 44:596— 601. SCRIBNER, EL. 1884. Observations on the genus Cima, with i of a new species. Pise. ee Nat. Sci. Philadelphia 1884:289— 291, pl. 7. and E.D. MERRILL. 1901. Studies on American ee Bull. U.S.D.A. Div. Agrostol. 24:2 STEBBINS, G.L. and B. € RAMPTON. 1961. A tpee revision of the grass genera of iB — 145. temperate North America. Recent Advances Bot 13 4 STORMER, P. 1949 [“1948"]. — on Cinna me in Lier and Modum (south eastern Norway). Blyctia 6:62 . {In Norwegian, with Englis TERRELL, E.E. 1971. Survey es occurrences of liquid or soft endosperm in grass genera. Bull. Torrey Bor. Club 98:264 — 268. TRINIUS, C.B. 1841. Gramina Aare, II. Callus ae nar Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pr. . 6:247 — 390. TZVELEV, N.N. 1976. Bosc Ones. Editio ’ ‘Nauka,” fen nen. ot Russian} [Cinna karataviensis, p. 329— WILLIS; J.C. 1973. A dictionary of the flowering plants and ferns (8th ed., revised by H.K.A. Shaw). Cambridge, London. Nn ion Sipa Book Reviews Botanical Research Institute of Texas S09 Pecan Street Fort Worth, TX 76102-4059, U.S.A. GOODLAND, Roser. Ed. 1990. Race to Save the Tropics: Ecology and Economics for a Sustainable Future. Island Press, Box 7, Covelo, CA 95428. Price unknow. 219 pp. This seminal book promotes the field of applied ecology as it relates to humid tropical ecosystems and their varied problems. Robert Goodland is to be commended for assembling the ee array of active monica applied ecologists who authored the nine chapters of the | d its role in the design of agricultural projects, applied book. cs include agroec ology an aie ets and natural forests Management, agricultural pest Management, dams and development in the humid tropics, the teaching of applied ecology to nationals in develop- , and the integration of applied ecology into national conservation and ing countries ise i setae) plans. owing awareness of the plight of global ee rainforests indicates that the ent resource text for the applied el — a ee lessons of this book are certainly on target. An ecologists and a valuable information source for non-scientists. Joe Fo Kuban. THE BLUE-EYED-GRASSES (SISYRINCHIUM: IRIDACEAE) OF ARKANSAS KATHLEEN L. HORNBERGER Department of Biology, Science Division Widener University Chester, PA 19013, U.S.A. ABSTRACT Data obtained from morphological characters and chromosome number indicate that for Arkansas: §. albidum Raf., S. angusti- jan seven species of Sisyrinchiwm should be recognizec foltum Mill., S. atlanticum Bickn., S. campestre Bickn., S. langlossiz Greene [S. praimosum Bickn.}, S. rosvlatum Bickn. [S. exile Bickn.], and S. sagittiferum Bickn. These seven ta are different from the eight previously recognized by Smith in that S. sagittiferum Bickn. was not listed and S. dangloisi? Greene and 8. pruinasum Bickn., plus S. rosulatum Bickn. and S. exile Bickn., were listed as four distinct species. Distribution maps were produced using Xa herbarium voucher specimens. INTRODUCTION The genus Sésyrinchium consists of both herbaceous perennials and annuals with simple or branched stems that may or may not be con- spicuously winged. The flowers are epigynous, subtended by a spathe (made of two overlapping bracts), and have undifferentiated actinomor- phic perianths. The mucronulate to aristulate tepals range in color from a purple eye-ring to more typically on white or yellow with purple stripes anc blue, purple, or white with a yellow eye-ring. No major revision had been done on this genus in the southeastern United States since Small (1933), which did not include the state of Arkansas. Smith (1978) recognized eight species for the state: S. a/bidum Raf., S. angustifolium Mill., S$. atlanticum Bickn., 5. campestre Bickn., S. exile Bickn., S. langloisii Greene, S. pruinosum Bickn., and S. rosulatum Bickn. Therefore, as part of a taxonomic revision on this genus in the SE U-S. for the Southeastern Flora Project (Massey & Radford 1981), which includes Arkansas, special attention was paid to this state's species in order to update Smith’s Atlas (1978). This study was part of a dissertation completed at the University of Arkansas in Fayetteville in January 1987. Sipa 14(4):597 — 604. 1991. 598 SYSTEMATIC STUDY A) Morphology Several hundred herbarium specimens of Blue-eyed-grass for the state were examined for 24 different characters (Hornberger 1987a). Discrimi- nant analysis indicated that of these 24 characters, the following provided the best separation for Arkansas species: paired vs. single spathes, length of outer to inner spathe bracts (equal to subequal vs. unequal), connation of outer spathe bract, stem width (includes wing on either side of raised cen- ter portion), stem wing width (average value of both wings), and dried capsule color and shape. Flower color and shape are also important characters, but do not preserve well, and are, therefore, not readily avail- able from herbarium specimens. However, they are included, along with the characters mentioned above, in the taxonomic key which follows this section. Essentially there are two basic morphological groups, one with simple stems (S. alhidum Raf., 8. campestre Bickn., and S. sagittiferum Bickn.) and one with branched stems (S. angustifolium Mill., 8. atlanticum Bickn., S. langloisii Greene, and S. rosulatum Bickn.). The species with simple stems are arranged in one of two ways: 1) the stems have either single spathes ona scape that has no cauline leaf, hence, there is no node or 2) the stems have paired sessile spathes (sometimes a single one) subtended by a cauline leaf. This latter type of stem is considered to have one node. Some of the simple- stemmed species may, however, occasionally branch, particularly S. sagit- ferum Bickn. (Hornberger 1987a). Species that typically have branched stems have one or more nodes, each of which has a cauline leaf and one or more pedunculate spathes. B) Chromosome Number Chromosome reports in the literature indicate that the genus is based on x =8, with most of the species being tetraploids (Oliver & Lewis 1962; Oliver 1966; Goldblatt 1982: and others). Among these chromosome reports were counts of n= 16 from flower bud material for only two species collected in Arkansas, S. campestre Bickn. and S. langloisii Greene (the latter taxon reported as S. pruinosum, Oliver & Lewis 1962; Lewis & Oliver 1961). I was able to confirm this number for S, campestre Bickn. from two different Arkansas populations, one in HorSpr- ing Co. and the other in Washington Co. (Hornberger 1987a). I also obtained the count of n= 16 from flower bud material for S. langlo- wit Greene, S. rosulatum Bickn., and S. sagittiferum Bickn., all collected, however, in Louisiana (Hornberger 1987a, 229 Sisyrinchium albidum Raf. was reported as n= 16 from Louisiana by Oliver & Lewis (1962). This number was originally reported by Bowden (1945) for a population collected in Virginia. It was further supported by Ingram (1964) for a population in Tennessee. Even though I have not had the opportunity to count the number for this species, it appears to be one of the tetraploid blue-eyed-grasses. Sisyrinchium angustifolium Mill. has been reported as n =48 from Louisiana by Oliver & Lewis (1962); they also reported this number for several populations in Texas. I have seen these specimens and agree with their identification. Hill (1984) recorded this same number for a Virginia population. However, Ingram (1964, 1967) reported n=40, 44, 45 for populations in North Carolina, Tennessee, and Virginia. Goldblatt (1982) feels that these conflicting reports for the same species are probably more a reflection of misidentification or incorrect counts than cytological diver- sit Sisyrinchinm atlanticum Bickn. has more reported diversity in chromo- some number than the preceding taxon. Numbers range from n = 8 (Oliver 1966) to n= 16 (Ingram 1964; Oliver 1966; Hill 1984) to n= 48 (Oliver & Lewis 1962). I have seen the specimens collected by Oliver (1966) and I agree with the identifications. However, I have seen the herbarium speci- men (Oliver 253, ASTC) of one of the two populations collected by Oliver & Lewis (1962) from Texas that was identified as $. atlanticum Bickn.; this specimen represents S. biforme Bickn., a species seemingly restricted to the Gulf Coast and offshore islands. C) Synonymy The genus Sisyrinchinm has been misunderstood taxonomically for more than a century. Because of subtle differences in morphology, disagreements among botanists on recognition of legitimate taxa, synonymy, of the proper epithet for a taxon have led to a plethora of species’ descriptions 1n the literature. Nomenclatural considerations, then, became a major task of the SE U.S. revision. Holotypes were requested for all taxa, and when they were determined lost or nonexistent, neotypes and lectotypes were designated. The only type specimen not seen was the one for S. rosulatum Bickn. because it was unavailable for this study. A complete discussion o types and synonyms is included in Hornberger (1987a). Synonyms will only be listed in this paper if they are different from Smith (1978) and would cause confusion if not included. KEY TO THE SPECIES IN ARKANSAS A. Outer spathe bract not connate at base, or only slightly so (1.0 mm or LOSS). gas sc ieceta te ote weit PD enh, cede al ue Sete RE GG ee eR Re B 600 B. Spathes paired at top of stem; bracts of outer spathe slightly unequal, stem with a single cauline leaf subtending the spathes ....... S. albidum Raf. B. Spathe single at top of stem; bracts unequal with outer one | 1/2—5 X length of inner one; stem with no cauline leaf subtending the spathe > - Outer spathe bract connate at base (1.1 mm or more) C. Spathe bracts equal to subequal 60 chica eg wa eatse e444 eee D Outer spathe bract connate at base up to 2.0 mm .... S, sagittiferum Bickn. D. Outer spathe bract connate at base for more than 2.0 mm .... E i. Stems 2.5 mm or more wide, wings 0.9 mm or more wide S. angustifolium Mill. 4 — 2. Stems less than 2.5 mm wide, wings less than 0.9 mm wide F. Capsules pale beige with purple or brown sutures, globose to subglobose; tepals white or yellow with purple stripes and purple eye-ring; flowers urceolate ». rosulatum Bickn. F. Capsules brown to black, globose to obovate; tepals light blue to purple, sometimes white, wit! 1 yellow eye-ring; flowers rotate G G. Inner spathe bract distinctly mucronate; spathes often deflected at base; capsules oblong-subglobose to obovate Auees ee Oy eee areata eur its eure toast S. atlanticum Bickn. G. Inner spathe bract not mucronate; spathes not deflected at base; capsules subglobose C. Spathe bracts noticeably unequal ~ eae a oe cei S. langlotsit Greene A a ag eh a eh a Be H H. Outer bract connate at base up to 2.0 mm; outer bract up to 3 lene CH OP IGAGl DICE acess dpee we ae eee caigeeie a's . Sagittiferum Bickn. H. Outer bract connate at base more than 2.0 mm: outer bract only 1 e205 Lene OF WMGr BRACE: gi anu cig dose hagden ce o< 4a J. Stems 2.0 mm or more wide, wings 0.9 mm or more wide; tepals light blue to white with yellow eye-ring; flowers rotate; capsules Gat OW .bb.a0 ce eee eee sien aeee bade S. angustifolium Mill. J. Stems less than 2.0 mm wide, wings less than 0.9 mm wide; tepals white or yellow with purple stripes and purple eye-ring; flowers urceolate; capsules pale beige with purple or brown su- CURES: pening a eye seetens Me 4 ese tan cock aioe ae oe a ett S. rosulatum Bickn. DISCUSSION Seven species were recognized in this study and will be briefly discussed in alphabetical order. Sisyrinchium albidum Raf. is a simple-stemmed perennial with paired sessile spathes at the node, where a large cauline leaf is found. Flowers are usually white with yellow eye-rings and the globose capsules dry pale beige to a straw color. Populations bloom late March to April and are found in prairies, woods, and roadsides. Sisyrinchium angustifolium Mill. is the most common and most to- bust species of Blue-eyed-grass in the state, with fairly wide leaves and 601 stems with conspicuous wings. This perennial produces stems that typical- ly have | node with a cauline leaf where two peduncles emerge. Flowers are light blue in color with yellow eye-rings and the globose to subglobose capsules dry dark brown. Plants are found in fields, woods, or along roadsi- des in April and May. This taxon has a very confusing nomenclatural his- tory which is presented elsewhere (Hornberger 1987a), but several manuals currently in use have names considered synonyms: S. bermudianum L. emend. Fern. and S. graminoides Bickn. (Gleason & Cronquist 1963; Steyermark 1963). Sisryinchium atlanticum Bickn. is a branched species found in scattered prairie areas in the state. Stems are terete to slightly flattened, with 1—2 nodes, and are narrowly winged ( 1 mm wide). Spathes are small, often deflected at the base; bracts are equal to subequal, and the inner bract is distinctly mucronate. The oblong-subglobose to obovate capsules dry dark brown to black. Flowers are generally light blue with yellow eye-rings, but the tepals are sometimes dark blue to purple. This perennial taxon blooms from March to April. Sisyrinchium campestre Bickn. is commonly found in prairies, meadows, and grassy areas along roadsides in April and May. This peren- nial has a simple stem with a single spathe at the top of the scape. The bracts of the spathe are very noticeably unequal, with the outer one at least 1 — 1/2 to 2 times or more the length of the inner, gibbous one. Sisyrinchium langloisii Greene is a branched perennial found along grassy roadsides, prairies, and disturbed areas in March and April. Its spathe bracts typically have a purple coloration which 1s often restricted to the base of the spathe. This taxon is similar in morphology to S. pruinosum Bickn., although the purple coloration of the spathe bracts is generally not present in the latter. Both taxa have the same chromosome number, 2n = 32 (Lewis & Oliver 1961; Oliver & Lewis 1962), and have been reported to hybridize in areas of overlapping range (Correll & Johnston 1970). Comparison of flavonoid spot profiles between a population from Texas and another from Louisiana showed similar patterns (Hornberger 1987a). Morphology, chromosome number, and flavonoid chemistry suggest that these various populations may actually be variations of one large species complex; therefore, I have synonymized S. prainosum Bickn. under the older name, S. /ang/oisiz Greene. Sisyrinchium rosulatum Bickn. ts the only annual Blue-eyed-grass found in Arkansas, seemingly restricted to several southern counties, plus Polk County. Flowers are yellow to white with purple to maroon stripes and purple to maroon eye-rings. These flowers are urceolate, instead of rotate as displayed by the flowers of the other six taxa. Spathes are slender 602 and foliaceous, with the outer bract slightly falcate at the apex and often | — 1/2 times longer than the inner bract. Globose to subglobose capsules dry pale beige with purple to brown stripes along the sutures. Populations can be found in disturbed areas of roadsides and lawns, prairies, river bottoms, and pine woods. Synonyms include S. exi/e Bickn, (Smith 1978). Sisyrinchium sagittiferum Bickn. is represented only from Miller County, with possible hybrids (sagittiferum X langloisii) collected in Union County. This taxon is usually represented by simple, leafless stems often with fibrous bases. Spathes are single (or sometimes paired) at the top of the scapes, being conspicuously broader than the stems. Spathe bracts are equal or the outer one can be three times the length of the inner one. Flowers are blue to purple with yellow eye-rings, bloom in March to April, and produce dark brown, globose to subglobose capsules, often with sub- marginal veins. This latter taxon is most similar in morphology to S. campestre Bickn., buc differs in several important respects: 1) S. sagzttiferum has outer spathe bracts that are connate for several mm: S, campestre has non-connate outer bracts (less than 1.0 mm); 2) spathes of S. sagittiferum are conspicuously wider than the stems and dry brownish, often mixed with purple...s, campestre has bracts that dry green in color and are not conspicuously wider than the stems; and 3) S. sagittiferum often has fibrous leaf bases attached to the stems, S. campestre seldom does. Sisyrinchium sagittiferum Bickn. had been recorded for Arkansas by Demaree (1943), but not by Smith (1978). My study indicates that it should be considered part of Arkansas’ flora. SUMMARY Based on data collected in this study from observation, investigation, and literature review, seven species of Susyrinchium are recognized for the state of Arkansas: S. albidum Raf., S. angustifolium Mill., S. atlanticum Bickn., 8. campestre Bickn., 8. langloisii Greene, S. rosvlatum Bickn., an S. sagittiferum Bickn. This information is presented in Smith (1988). Dis- tribution maps were prepared for each taxon based on herbarium voucher specimens. A dot indicates that at least one specimen exists for a particular taxon in a particular county (Fig. 1). (Note: the dot in Union County for S. sagittiferum Bickn. represents possible hybrids between it and S. langloistt reene). ou ACKNOWLEDGEMENTS I thank E. B. Smith for serving as my dissertation advisor. Many thanks to Jerry Shell and his wonderful dog, Mittens, for helping me collect ower uds. I would also like to thank the Arkansas Native Plant Society for §, albidum Raf. S. rosulatum Bickn. FIG. 1. Documented distribution of Arkansas Sisyrinchium. awarding me the Aileen McWilliams Scholarship in Botany tn 1986 which provided encouragement and financial support to finish my dissertation. Special thanks are also extended to the curators of the following herbaria for lending me specimens that made this study possible: ASTC, BH, BLH, BM, FLAS, FSU, GA, GH, LAE MA, MIN, MO, NCSC, NCU, ND, NDG, NLU, NY, PH, SMU, TENN, UAM, UARK, US, and WIS. 604 REFERENCES BOWDEN, W. M. 1945. A list of chromosome numbers in higher plants. [. Acanthaceae to aa Amer. J. Bot. 32(2):81 —92. CORRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the baer ra of Texas. Texas Research Foundation, Renner. 1881 pp. (Sisyrinchium, pp. 425 — 428). ae D. 1943. ee of the vascular plants of pee Taxodium I(1): : ae a eee p. GLEASON, H. A. and A. C cue 1963. Manual of vascular plants of northeastern United States a adjacent Canada. D. Van Nostrand Co., NY. 810 pp. Szsyrinchium, pp. 219— 220). GOLDBLATT, P1982. Chromosome cytology in relation to suprageneric systematics of neotropical i Syst. Bot. 7(2):186— 198. ILL, L. M. 1984. A floristic and chromosomal study of Sisyrinchium (Iridaceae) in Virginia. Cave 49(2):62 —68. HORNBERGER, K. L. 1987a. Hanae of the genus Szsyrinchium (Iridaceae) in the southeastern ae States. PhD , Univ. of Arkansas. 328 pp. —_________.. 1987b. In A. Lov pene number reports XCV. Taxon 36:497. ecg R. 1964. The taxonomy one cytology of the genus Sane (Iridaceae). PhD » Univ. of Durham. 105 pp. ——_____—.. 1967. On the identity of the Irish populations of Sésyrinchinm. Watsonia 6(5): 283 — 289. LEWIS, W. H. and R. L. OLIVER. 1961. Meiotic chromosomes in six Texas and Mexican nea and Svsyrinchinm (Lridaceae). Southw. Naturalist 6(1):45 — 46. MASSEY, J. R. and L. S. RD. 1981. Revised contributors’ for the vascular ee of ie southeastern United States. Univ. of North Carolina. 21y OLIVER, R. 1966. In chromosome numbers of phanerogams. I. ne ve Bot. Gatd. 52: 1 and W. H. LEWIS. 1962. Seb as numbers of Szsyrinchium _ in eastern North America. Sida 1(1):43 — SMALL, J. K 33. Manual of the southeastern flora. His a of North Carolina Press, Chapel Mil ee t pp. Sasyrinchinm, pp. 327 — 330) MITH, E. B. 1978. Anatlas and annotated list of the oun plants of Arkansas. Student Union een i. of Arkansas, Fayetteville. 592 pp. [OUT OF PRINT]. (Svs- yrinchium, pp. 481— 482). fe LOS An atlas and annotated list of the vascular plants of Arkansas. 2nd ed. Kinko's, 653 West Dickson, Fayetteville, AR, 72701. 489 pp. ~ (Stoyrinehiuy pp. 381—382 STEYERMARK, J. A. 1963. Flora of as The Iowa State University Press, Ames. 1728 pp. Sayrinchium, pp. 466 - NOTES LYCIANTHES ASARIFOLIA (SOLANACEAE), NEW TO NORTH AMERICA — In November of 1989 a colony of Lycianthes asarifolia (Kunth & Bouché) Bitter, was discovered growing in City Park, New on Orleans, Louisiana. The plants are stoloniferous, creeping herbs with sparsely shaggy-pubescent internodes to 7 cm long. The leaves are solitary at each node, the petioles slender, to 10 cm long, laterally pubescent or glabrous, the leaf blades cordate to reniform, to9 X 8 cm, obtuse to roun- ded at apex, cordate-auriculate at base, glabrous to subciliate. The flowers are solitary at each node, the pedicels slender, 4— 7 cm long, subglabrous, nodding at apex, subtended by a small auriculate bract at base. The calyx 1s cupular, to 4 X 5 mm at anthesis, ca. 10-costate, appressed-pubescent, 5-toothed or occasionally also with minute apiculations alternating with the teeth. The corolla is rotate-campanulate, the limb usually reflexed, 16 — 20 mm broad, 5-lobed, glabrous, white. The 5 stamens are equal, the anthers 2.5 —3 mm long, apically dehiscent. The ovary is 1-2 mm in diameter, the style 5 — 6 mm long, slender, the stigma truncate to subcap1- tate. The voucher collection ts Fesbelman 107 (MO, NO, NY, US), compris- ing stems, leaves, and flowers taken from a colony covering roughly 450 sq. meters beneath Quercus virginiana. The colony appears to be expanding vegetatively, as no fruits have been seen at the City Park location or on plants propagated from cuttings. We surmise that the colony developed from a single introduction and is a self-sterile clone. The species is well adapted to mowing, and our plants withstood 72 hours of freezing temperatures (as low as 11° F) in 1989 without noticeable damage. Lyctan- thes asarifolia thus appears to be well adapted to the Gulf Coast climate, at least in shaded lawns, where it makes a vigorous and attractive ground- cover. Lycianthes is a genus of 150 — 200 species, mostly of tropical America, but with a dozen or more species in Asia and the South Pacific. It is usually distinguished from Solanum yx with 10 small teeth — —_— oy the 10-nerved ca appearing as enations below the truncate apex, but the teeth are sometimes absent. A good discussion of generic characters is given by D’Arcy (Ann. Missouri Bot. Gard. 60: 631. 19753). On account of its unusual habit, Lycranthes asarifolia was, with L. repens (Sprengel) Bitter, placed by Bitter (Abh. Nat. Ver. Bremen 24:422 — 426. 1920) in Lycianthes sect. Asaropsis. Both species are South American, L. Sipa 14(4):605. 1991. 606 dsavifolia reported from Venezuela, Colombia, Bolivia, Paraguay, and Argentina, while L. repews is apparently restricted to southeastern Brazil. Bitter distinguished the more variable L. asarifolia from L. repens on the basis of its usually longer petioles, as well as larger calyces, anthers, and ruits, among other features of indument and flower color (corollas reported as pale blue to violet in L. repens). Our plants clearly fall within Bitters concept of L. asarifolia, and we have compared e from Venezuela (A/ston 6105, USF), Colombia (Snerdern 31216, GH), Bolivia (Steinbach 6212, GH; Rusby 1875, GH; Solomon 13467, ee Nee 35104, NY), and Beil (Dusén 11315, GH). Under cultivation, our plants show considerable variability in pubescence as well as leaf blade shape and dimensions. Should the two species be united, the name Lyctanthes repens (Sprengel) Bitter, based on Boldoa repens Sprengel [Syst. Veg. 1: 179. 1824} would have priority. An excellent illustration of L. repens (as Solanum violaefolinm Schott) can be found in Martius, Fl. Brasil. 10:52, tab. 4, fig. 44 — 46; tab. 12. 1846. A note with Nee 35104 states that the orange, rather pleasant-tasting fruits of “motojo- bobo” are edible and made into preserves We thank Dr. M. Nee (NY) for invaluable assistance with the identifica- tion and nomenclature of this species, and Dr. M. Molvray (NO) for help with Bitter’s German. The curators of GH, MO, NY, SMU, US, and USF searched their collections for North American records of Lycianthes asari- folia, or made other material available for our examination. — Steven P Darwin and Toby Fetbelman, Dept. of Ecology, Evolution, and Organismal Biol- ogy, Tulane University, New Orleans, LA 70118, U.S.A. — eon PALIURUS SPINA-CHRISTI (RHAMNACEAE) NEW FOR NORTH AMERICA IN TEXAS — Paliurus spina-christi Miller, an Eurasian rhamnaceous shrub not previously reported as naturalized in North America, has been found growing on the Edwards Plateau of central Texas. The plant is known only from Gillespie County where it was apparently introduced ca. LOO years ago. It is well established along the flood plains of two crecks and the Pedernales River and has become a pernicious weed. In 1986, a rancher brought to the attention of Gillespie County agri- cultural extension agent Duery Menzies the presence of an unusual spiny shrub that was invading his pastureland along Dittmar Creek 21 km west of Fredricksburg in Gillespie County. The plant was taken to Texas A&M University in College Station by Roger Landers, Range Specialist, Texas Sipa 14(4):606. 1991. 607 Agricultural Extension Service, where it was identified by Kancheepuram N. Ghandi as Paliurus spina-christi, known in the vernacular as Christ- thorn. Another specimen was taken to the University of Texas in Austin where Marshall Johnston also identified it as P. spina-christz. Ghandi (pers. comm.) stated that there has been a Palimrus on the TAMU campus at College Station for many years but that it has not produced other plants. Johnston (1969 and pers. comm.) who has authored publications on Rhamnaceae and traveled internationally to study the family, notes that Paliurus is not known in the wild as an escape in Texas or the United States. Kartesz & Kartesz (1980) does not list this taxon from the United States or Canada. James B. Phipps (pers. comm., Western Ontario Univ.) says that this plant has not been found in Canada. In addition a check of the MEXU herbarium revealed no specimens from Mexico and Rogers McVaugh (pers. comm. Univ. North Carolina, Chapel Hill) note its absence and any pub- lished report in Mexico. This plant would thus appear to be the first report of P. spina-christi for all of North America. Collection data are: Collections examined: TEXAS. Gillespie Co.:Dittmar Creek, 5.5 km N Hwy 290, 4 May 1988, peat 2766 (BRIT/SMU); Dittmar Creek at Hwy 290, 4 May 1988, O'’Kennon 2767 (BRIT/SMU); a Creek, 200 m S of Tivydale Rd, 17 Aug 1988, O’Kennon 3117 Rada U); Pedernales River, Boos Rd, 6.5 km § of Fredricksburg, 22 Aug 1990, O’Kennon 7613 (BRIT/SMU). Paliurus spina-christi is a deciduous multi-trunked shrub 3 —4(-5) m nigh. The stems are spreading and armed with curved spiny stipular thorns to | cm. The leaves are alternate and distichous or in two ranks, short petiolate, ovate, crenate-serrate, and 2—4 cm long. The bright yellow Howers are small but numerous in axillary cymes or terminal panicles and are striking in appearence when in full bloom. The greenish yellow fruits are flat disc-like capsules 2—3 cm in diameter. The dried capsules often remain on the plant until the following year’s howers are blooming on new growth. This Pa/iwrus is one of the legendary trees from which the Crown of Thorns was supposedly made. It has been in cultivation in Europe for over 300 years, and is sometimes cultivated in the United States (Everett 1981). Since the first discovery of Pa/iurus, an investigation has revealed the source of the plant’s introduction and the extent of its range. In the late 1800's a German homesteader planted seeds brought from Europe in order to form a spiny hedgerow along the west bank of upper Dittmar Creek 9 km north of highway 290 and 19 km west of Fredricksburg. Longtime residents of this area remember the 100 meter long hedge as having always been there. One 93 year old man who has lived on the site most of his life Sipa 14(4):607. 1991. 608 remembers not only the hedge but other younger thorny plants growing a short distance downstream. He states that as time passed more shrubs appeared downstream beyond sight of his ranch. Now, approximately 100 years after the plant’s introduction, the Christ-thorn has proliferated exten- sively along Dittmar Creek 9 km south to highway 290 and well beyond. Dittmar Creek feeds into Spring Creek which flows south 11 km more before emptying into the Pedernales River south of Morris Ranch. Paliurus is found in large thicket-forming populations along the entire length of these creeks and is widely scattered along the Pedernales for 13 more km until just south of Fredricksburg. In time it will undoubtedly be found farther east, perhaps into Blanco County and beyond. The capsules and their seeds appear to be carried solely by water and _— sprout primarily in the wide rich flood plains along the waterways. Occa- sional plants are found on flats just above the traditional flood plain. This can be explained by the changes in water level during torrential rains and resultant flash flooding typical of the deeply cut terrain of the central Ed- wards Plateau. Prior to 1978 the plants apparently remained ina restricted range from their point of introduction along upper Dittmar Creek to just north of Hwy 290 and had not yet become conspicuous pests. A 1978 flood which was associated with a stalled tropical depression, the remnants of hurricane Amelia, was most probably the force which generated the explos- ive proliferation of an entire generation of plants of similar size all along its present range and well outside the normal flood plain. Because Palimrus heavily infests prime fertile grazing land and habitats of native wetland flora, it is now considered a pernicious weed with the potential for being as disastrous as other old world invaders such as Lonicera japonica Thunb. (Japenese honeysuckle), Sapimm sebiferum (L.)Roxb. (Chinese tallow), and Pueraria lobata (Willd.) Ohwi. (kudzu). These plants are having devastating deleterious effects on many native plants as they take over more and more territory. Because of its rapid proliferation, a program to eradicate Pa/iurus has been initiated by the county under the direction of Mr. Menzies, and it is apparently effective. Thanks to Barney Lipscomb of the Botanical Research Institute of Texas (BRIT) and Mr. Duery Menzies for research, helpful comments and sug ges- tions - manuscript. — Robert J. O'Kennon, 30 Saint Laurent Place, Dallas, foe 2d PL AS Us REFERENCES EVERETT, T. 1981. Encyclopedia of Horticulture. Vol. 7. Garland Publishing, Inc., New York, NY Sipa 14(4):608. 1991. 609 JOHNSTON, M. C. eee A. JOHNSTON. 1969. Rhamnaceae, In: C.L. Lundell (ed.), een 7 Texas. Vol 2:35 — 392. Texas Research Foundation, Renne KARTESZ, J. T. and R. ae BSZ. 1980. A synonymized checklist of ie vascular flora of the a States, Canada, and Greenland. UNC, Chapel Hill. EUPHORBIA ~ LATHYRIS (EUPHORBIACEAE) NEW — FOR TEXAS — Euphorbia lathyris L., a European herbaceous euphorb not previously reported from Texas has been found naturalized on the Edwards Plateau of central Texas. The plant is known only from a single site in western Gillespie County. Euphorbia lathyris, caper or myrtle spurge, a European euphorb has been found growing on the banks of the Threadgill Creek in western Gillespie County, Texas. This spurge is native to the Mediterranean region of southern Europe and according to Marshall C. Johnston (pers. comm.) has not been previously reported from Texas. It is not listed in the more recent Texas checklists by Johnston (1988, 1990) and Hatch et al. (1990). Caper spurge is widely cultivated in Europe and ts occasionally cultiva- ted in the United States. It is known to have escaped cultivation in the Atlantic Northeast and in California. It is also known as “mole plant” because of its believed properties that repel moles from lawns. The seeds have cathartic properties. Euphorbia lathyris is distinct from other Texas euphorbs in its tall, somewhat conspicuous habit. It gets up to a meter tall with narrow leaves to 14 cm long arranged in four vertical rows along the stem. The infloresc- ence is umbellately branched with the floral bracts lanceolate to ovate. Its crescent-shaped glands are prolonged into short horns. The subglobose capsules are 1.0— 1.2 cm wide. The author previously observed non-flowering plants (vegetative) in Gillespie County, but were later eaten by animals and never positively identified. The plants are found ona cattle ranch inhabited by angora goats and white-tail deer, and it is likely that few of the plants ever reach maturity. The collection site is within a ean aed fence exclosure es- tablished for the purposes of native plant research and affords protection from these animals. Plants were first identified in May, 1990 when in full flower. Marshall Johnston visited the site with the author at that time and collected a single specimen from a colony of six plants along the creek bank. In August the author collected a fruiting specimen. The fruiting specimen was taken to SMU where the author and Wm. E Mahler deter- mined it to be E. /athyris matching European collections in the herbarium. Sipa 14(4):609. 1991, 610 Eight young seedlings had appeared by November in the vicinity and remained 20 — 25 cm tall through spring and summer of 1991; apparently this being the first year's growth of the biennial. At present there is no information as to the source of E. /athyris at this site. There 1s no report of it ever being cultivated in Gillespie County or anywhere else in Texas. It 1s possible that seeds were brought by the Germans who settled this area in 1846 and that the plants have escaped detection until now. Collection data are: Collections examined: TEXAS. Gillespie Co.: Threadgill Creek, . = N of Harper, | | 3 S of Doss on McGinley Ranch, 13 May 1990, O’Kennon and M. C. Johnston 6697 (T ; Threadgill Creek, 11 km N of Harper on McGinley Ranch, 3 a 1990, O’Kennon a pore U). — Robert J. O’Kennon, 30 St. Laurent Place, Dallas, TX 75225-8111, USA REFERENCES CORRELL, D. and M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, . HATCH, S.L., K.N. GANDH I, L. E. BROWN. 1990. Checklist of the vascular plants of Texas. Teas Agric. Exp. Sta. MP-1655, Texas A&M University, College Station, TX. JOHNSTON, M.C. 1988. The vascular plants of Texas: a list, up- Piet the “Manual of the vascular - ants of Texas. Published by the author, Austin, Texa JOHNSTON, 1990. The vascular plants of Texas: a list, up-dating the “Manual of the vascular ie of Texas. Second ed. ae hed by the author, Austin, Texas. — ULMUS PARVIFOLIA (ULMACEAE) NATURALIZED IN KEN- TUCKY — The exotic elm commonly naturalized in Kentucky and el- sewhere in the United States is the spring-flowering Siberian elm (U/mus pumila L.), often misnamed the Chinese elm. For several years we have noted many spontaneous individuals of another Asiatic species of U/mus, the fall-flowering “true” Chinese elm or lacebark elm (U. parvifolia Jacq.) in Loutsville, Jefferson County, Kentucky. Individuals of various sizes — seedlings through mature trees up to 35 cm DBH — can be found in empty lots, in fencerows, and along railroads. They are commonly associated with tree-of-heaven (As/anthus altissima (Mill.) Swingle), northern catalpa (Catalpa speciosa Warder), Osage-orange (Maclura pomifera (Raf.) Schneider), and Amur honeysuckle (Lonicera maackit Maxim.). e voucher cited below — from a small tree 10 cm DBH and in abundant fruit — was growing between the rails of an abandoned railroad track. These is no doubt that U. parvifolia is well naturalized in Louisville. Sipa 14(4):610. 1991. 611 FIG. 1. Ulmus parvifolia. Upper, frucing branch with cite elliptic fruits. Lower left, leaf non- fruiting oe lower right, nearly orbicular fruit. The vertical lines represent 1 cm. Drawings by Keith Book Stipa 14(4):611. 1991. 612 The species is not included in the standard northeastern U.S. manuals (Fernald 1950; Gleason and Cronquist 1963) or in floristic works of most states adjacent to Kentucky: Illinois (Mohlenbrock 1982), Indiana (Crovello et al. 1983), West Virginia (Strausbaugh and Core 1953), Virginia (Harvill et al. 1981), Tennessee (Sharp et al. 1960), and Missouri (Steyermark 1963, Recienicn and Turner 1990). Weishaupt (1971), however, listed it for Ohto. It is not accounted for by Elias (1980) and receives but passing mention, as a tree “in cultivation,” in Little (1979). In the account of U/mus for Vasewlar flora of the southeastern United States (Nicely, in press) the species is said to be “occasionally escaped and es- tablished locally” in Georgia, Illinois, Maryland, Ohio, and Virginia. It has also been noted to be “growing without cultivation” in California (Howell 1966). Though rather similar to U. pumila in leaves, U. parvifolia is dis- tinguished by its autumnal flowering and by its bark, which exfoliates in irregular scales and appears mottled, often beautifully so, in patches of brown, green, gray, and orange. Trees as small as 10 cm DBH may begin to show the mottling. For the winter interest of its bark alone, the tree is well worth growing. We were able to recognize individuals of the species from afar after leaf fall by the brown-hazy appearance of the crown brought about by the abundant fruits, which bedeck the branchlets and may persist until mid-winter. A voucher specimen (Thieret & Medley 567 Ms collected on 14 December L990 has been deposited in KNK and UNC. — Max E. Medley, Depart- ment of Biology, University of Louisville, oe KY 40292, U.S.A., and John W. Thieret, Department of Biological aii Northern Kentucky Univer- sity, Highland Heights, KY 41076, U.S./ REPERENCES CROVELLO, T.J., C.A. KELLER, and J.T. KARTESZ. 1983. The vascular plants of Indiana: a computer based checklist. Univ. of Notre Dame Press, Notre Dame, IN. 13 PI p. ELIAS, T.S. 1980. The complete trees of North America. Van Nostrand Reinhold Company, New York. 948 pp. FERNALD, M.L. 1950. Gray's manual of botany, 8th ed. American Book Company, New ° GLEASON, H.A., and A. CRONQUIST. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. D. Van Nostrand Company, Princeton, NJ. 810 pp. HARVILL, A.M., JR., T.R. BRADLEY, and C.E. STEVENS. 1981. Aclas of the Virginia Hora. Pare IH. Dicotyledons. Virginia Botanical Associates, Farmville. 148 pp Stipa 14(4):612. 1991. 613 HOWELL, J.T. 1966. Concerning two Asiatic elms {in California}. Leafl. W. Bort. 10:328- a2): mer E.L., Jr. 1979. Checklist of United States trees (mative and naturalized). Le ‘Aerie: Handbook. 541. 375 pp eer K, R.H. 1982. The illustrated flora of Illinois. Flowering plants. Bi Suen to spurges. Southern Illinois Univ. Press, Carbondale. 234 pp NICELY. _ n.d. Ulmaceae. Vascular flora of the southeastern United Grates, Univ. of Noth pene Press, Chapel ree In press. SHARP. A.J., R.E. SHANKS, H.L. SHERMAN, and D.H. NORRIS. 1960. A : a nary checklist of dicots in ate see. Univ. of Tennessee, Knoxville. Mime STEYERMARK, J.A. [1963]. Flora of Tckoar: | a State Univ. Press, Ames. 1725 pp. STRAUSBAUGH, PD., and E.L. CORE. 1953. Flora of West Vinotoia on : West Virginia a - Ser. 53(12 — 1):275 a WHISHAUPT. ©.G,, 197 1. ee vascalas plants of Ohio, 3rd ed. Kendall/Hunt Publishing Company, Dubuque, IA. YATSKIBVYCH, G;., aay ak 1990. Catalogue of the flora of Missourt, Monogr. Syst. Bot. 37 BRACHIARIA PLANTAGINEA, IMPERATA CYLINDRICA, AND PANICUM MAXIMUM: THREE GRASSES (POACEAE W TO LOUISIANA AND A RANGE EXTENSION FOR ROTTBOELLIA COCHINCHINENSIS. — Three tropical or subtropical grasses (Brachi- avia plantaginea (Link.) Hitche., Imperata cylindrica (L.) Beauy., and Panicum maximum Jacq.) not reported by Allen (1980) nor Thomas and Allen (1984) are now known to occur in Louisiana. Brachiaria plantaginea (Link.) Hitchc. (Plantain signal grass) is native from Mexico to Brazil and Bolivia. Hitchcock (1950) reported it as adven- tive in Georgia, New Jersey, and Pennsylvania. Collections from the southern part of Louisiana apparently represent the first for the State. Collection data are: Lafourche Par.: along beach and edge of marsh at the Gulf of Mexico S of the end of La. 3090 ‘ . Fourchon City § of Leeville; Sec _ 24, 7238, R22E, 7 Nov 1987, Thomas 103240 (NLU). St. Charles Par.: along road near Illinois Coagal Railroad and I-10 in the Bonnet Carre aie 21 Sep 1974, Montz 3/6/ (LSU, NO) Imperata cylindrica (L.) Beauv. (Cogon grass) has been reported from Florida and from ballast in Oregon (Hitchcock, 1950). Although Clewell (1985) considers the closely related taxon (Imperata brasiliensis Trin.) to be synonymous with this species; the two taxa are separated by Gabel (1932), Hitchcock (1950), Kartez and Kartez (1980), and Wunderlin (1982). Louisiana specimens of the two taxa can usually be separated by using the following key: Sipa 14(4):613. 1991. 614 Spikelets 3.5 mm or longer; anthers two; cauline blades wider than 5 mm eA aes Gahh ge ire ¥ a Da Mag oe ee et pale tA een dee Imperata cylindrica 5mm 1 Saisie eee tenng sign th aries acd nea eh aaa ced aE SI yee ee Imperata brasiliensis Imperata brastiensis Trin was reported new to Louisiana by Allen (1974) from Washington Parish. Current records indicate that it has spread to Ascension, Orleans, St. Tammany, and Tangipahoa parishes. Imperata cylindrica (L.) Beauv. has been collected from three sites in St. Tammany Parish including: west bank of La. 1129, 0.5 mi N of La. 40 E of Blond; Sec 22, TSN, RIIE, very large expanding population, 24 May 1979, Thomas 64869 (NLU). Panicum maximum Jacq. (Guinea grass) is a widely distributed grass of tropical areas and is native to Africa (Gould 1975). It has been reported from Florida and Texas, and a collection from New Orleans is apparently the first record of this grass from Louisiana. Collection data are: Orleans Par.: waste area along railroad at Bienville Street Wharf on the Mississippi River in New Orleans, 21 Jun 1979, Thomas 66207 (NLU). These three grasses new to Louisiana should be monitored to see if they become as widespread and noxious as Rottboellia cochinchinensis (R. exaltata L. f.) (itch grass) in the State. Rottboellia cochinchinensis (Lour.) W. Clayton is a noxious weed that was reported from Louisiana by Thieret (1963). Allen (1980) reported it from Lafayette, Pointe Coupee, St. Martin, and St. Tammany parishes. During the preparation of an atlas of the Monocotyle- dons of the State and of a revision of “Grasses of Louisiana”, the authors added records of R. cochinchinensis from twelve additional parishes (Acadia, Allen, Cameron, East Baton Rouge, Evangeline, Iberia, Lafourche, St. Charles, St. Landry, St. Mary, Terrebonne, and Vermilion). On September 6, 1991, Allen located a large population of this weed along U.S. 165 in the pine uplands of North Louisiana at least 65 miles north of any previous known locality. This population is significant in that it is not in an agri- cultural area and is only about 70 miles south of the Arkansas State line. The collection data are: Caldwell Par.: roadbank of U.S. 165 on hill in pine woods at La. 844 near Clarks, 3.9 mi S of Grayson; heavily herbicided roadside with a nearly pure stand of Sida spinosa L., population including several hundred plants, 10 Sep 1991, Thomas 125135 (NLU, and duplicates to be distributed). — Charles M. Allen and R. Dale Thomas, Herbarium, Dept. of Biology, North- east Louisiana University, Monroe, LA 71209, U.S.A. and Michel G. Lelong, Dept. of Biological Sciences, University of South Alabama, Mobile, AL 36688. mG Sipa 14(4):614. 1991. REFERENCES ALLEN, C. M. 1974. ara species of grasses (Poaceae) new to Louisiana. Proc. Louisiana Acad. Sct. 37:18 — ALLEN, C. M. 1980. Grasses i eoeres Univ. Southwestern Louisiana, Lafayette. CLEWELL, A. E 1985. Guide to the vascular plants of the Florida Pe pandic Univ. Presses a Florida, Gainesville. GABEL, M. L. 1982. A biosystematic study of the genus Imperata (Gramineae: Andropo- goneae). a dissertation, Iowa State Univ., S. GOULD, EW. 1975. The grasses of Texas. Texas A. & M. Univ. Press, College Station. are tee A. §. 1950. Manual of the grasses of the United States. U.S. Dept. Agric., Washing KARTEZ, r, T ‘AND R. KARTEZ. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Univ. North Carolina Press, Chapel Hill. THIERET, J. T. 1963. Additions to the flora of Louisiana. Castanea 28:169— 170 THOMAS, R.D. AND C. M. ALLEN. 1984. Checklist of the Prteridosperms, Gym- nosperms, and Monocotyledons of Louisiana. Contr. Herb. Northeast Louisiana. Univ. A= 55. WUNDERLIN, R. P. 1982. Guide to the vascular plants of Central Florida. Univ. Presses of Florida, Gainesville. ADDITIONAL RECORDS OF CYPERUS ENTRERIANUS (CYPERACEAE) IN THE UNITED STATES. — Cyperus entrertanus Bockeler was first reported in the United States by Carter (1990). Initially, it was cited from 15 counties in the Atlantic and Gulf coastal plains rang- ing from Georgia to eastern Texas. Subsequently, additional collections of C. entrerianus have been made in Florida, Georgia, and Texas, which are cited below. U.S.A. Florida. Holmes Co.: 1.8 mi W of Westville, mowed ditch along Hwy US 90, locally common, 4 Aug 1990, Carter 8393 (FLAS, FSU, VDB, VSC). Jackson Co.: 1.9 mi N of Round Lake, ditch along south-bound lane of Hwy US 231, locally common, 4 Aug a Carter 8392 (FLAS, FSU, IBE, MICH, MO, NY, US, VDB, VSC). Washington iryville, south side of town, ditch beside Hwy FL 279, between Hwys US I-10 and US a ell pee 4 — ee Carter 8397 (FLAS, FSU, GH, IBE, MICH, MO, NY, NYS, TAE , VD , VSC). GEORGIA. Echols Cae disturbed barrow pit aie dice; oe re icy a 94, 1.8 mi E of jet. with US 129 in Sratenville, flatwoods, 9 Jul 1991, Carter aa Jones 8806 (FLAS, FSU, GH, IBE, MICH, MO, NY. YS, TAES, US, VDB, VSC). TEXAS. Jackson Co.: jct. of Hafernick Rd. (C ounty Rd. 267) and US 59, NE of Ganado, open mesic roadside with black clayey soil, frequent, 30 Oct 1990, S. & G. Jones 6038 (TAES, VSC). Newton Co.: 150 mS on Hwy 87 from tts jct. with FR 2829, § of Trout Creek, open hydric roadside ditch with sandy mucky soil, fre- quent, 27 Jul 1990, S. & G. Jones 5645 & J.K. Wipff (TAES, VSC). OS While these new records do not extend the range of Cyperus entrertanus in the United States, they do fill distributional gaps in its known range. Each Stipa 14(4):615. 1991. 616 is anew county record. Thus, the number of counties where C. extrerianus is known in the United States is increased by 40% to a total of 21 counties. Appreciation ts expressed to Mr. W.K. George, Valdosta, Georgia, for supporting field work in Florida during 1990 and to the Valdosta State College Faculty Research Fund for meeting publication — ex- penses. — Richard Carter, Herbarium (VSC), Department of Biology, Valdosta State College, Valdosta, GA 31698, U.S.A, and Stanley D. Jones, S.M. Tracy Herbarium, Department of Rangeland Ecology and Management, Texas AGM University, College Station, TX 77843, U.S.A REFERENCE CARTER, R. 1990. C ne entrertanus (Cyperaceae), an overlooked species in temperate North pee Sida 14:69—77. A FIRST REPORT OF LEPTOCHLOA SCABRA NEES (POACEAE) FROM ALABAMA. — Recent examination of herbarium material of Lep- tochloa Beauv. has revealed the presence of Leptochloa scabra Nees in Alabama, which we report for the first time. It now appears to be es- tablished in south Alabama along the Tombigbee River, the Mobile River, and in the Mobile Delta. Commonly called Rough Sprangletop, this is first report for the species in the United States outside of Louisiana, where it apparently became adventive in the New Orleans area and was collected there by A. B. Langlois as early as 1884. It has since spread throughout much of Louisiana (Allen 1980), where it can form large populations. For example, L. scabra is common tn localized areas of the Bonnet Carre Spillway in St. Charles Parish, where it occurs with Leptochloa panicoides (Pres!) Hitchc. (N. Snow, pers. obsv. 1990) Leptochloa scabra can be easily confused with L. panicoides. The former is distinguished by its shorter, tightly imbricate spikelets, the keeled lemmas, its distinctly flexuous panicle branches, and culms that are somewhat flattened near the base. The earlier report by Lelong (1988) for L. pantcoides in south Alabama is here amended to L. scabra. Rough sprangletop is native to tropical America, occurring from Michoacan, Mexico, south to Argentina and Brazil, and in the West Indies (Hitchcock 1936). It seems likely that it will continue to spread slowly in the ee humid regions in the United States. Speci I : ALABAMA. Mobile Co. : by truck zl sass 98-90 across river from Mobile. nu ae area, 20 Sep 1975, Kral 56597 (MO): T2N, RIE, E boundary of Sect. 20, sandbar of small island along W bank of Tombigbee ite dineeely N of its confluence Sipa 14(4):616. 1991, 617 with the Alabama River, with Cyperus erythrorhizos, hs Sia glomerata, sr ae and Aster lateriflorus, 25 Oct “ap M. G. Lelong 11,732 (MO; USAM). Washingtc T2N, RIE, SE corner of Sect. 6, common along N banks of Tombigbee om a ea eaine, Spermacoce ae ae sates 25 Oct 1980, M. G. Lelong 759 (MO; USAM). Baldwin Co.: . ROE, 1/4 of Sect. 40, clay bluff along $ ee k of Alabama River cutoff, with cee cr Fimbristylis vablit, es tial halicacabum, 25 Oct 1980, M. G. Lelong 11,774 (MO; USAM), TIN, RIE E boundary of Sect. 54, wooded banks of Bottle C ae about 0.25 mi E of Indian ie Indian Mound island, Mobile Delta, with Leonotis Gee Apios americana, Matelea gonocarpa, \5 Nov 1980, M. G. Lelong 11,892 (MO; L — Neil Snow, Box 1137, Department of Biology, wae University, St. Louis, MO 63130, U.S.A. and Michel G. Lelong, Department of jee Sciences, University of South Alabama, Mobile, AL 36088, U.S.A REFERENCES ALLEN, C. M. oe Grasses of Louisiana. The University of Southwestern Louisiana, Lafayette. 35 HITCHCOCK, A. - 1936. Manual of the Grasses of the West Indies. Misc. Publ. 243, U. S. Dept. Agric. Government Printing Office. es Di -G. LELONG, M. G. 1988. Noteworthy monocots of Mobile and Baldwin counties, Alabama. Sida 13:101— 113. OCCURRENCE OF POTAMOGETON — PERFOLIATUS _ L. (POTAMOGETONACEAE) IN LOUISIANA — Louisiana collections of Potamogeton perfoliatus are few in number and are restricted to the northern and eastern shorelines of Lake Pontchartrain and adjacent marsh areas. Ogden (1943) listed a specimen collected by Riddell from the Tchefuncte River lighthouse at Lake Pontchartrain on 16 August 1838. Haynes (1968) reported a specimen collected by Clair A. Brown (LSU 5676) from St. Tammany Parish at Mandeville in 1945. Montz (1978) observed that P. perfoliatus was abundant in 1973 near Pointe aux Herbes in Orleans Parish. However, Mayer (1986) noted the conspicuous absence of P. perfolzatus from the Lake Pontchartrain estuarine system in 1985. Lester (1988) listed the current status of the species in Louisiana as unknown and reported that recent attempts to locate populations as unsuccessful. On 21 June 1990, we found a7 X 17 m submerged bed of P. perfoliatus along the northern shoreline of Lake Pontchartrain, ca. 1500 m west of Bayou Lacombe, south of the St. Tammany State Wildlife Refuge, St. Tammany Parish, Louisiana. The St. Tammany State Wildlife Refuge ts classified as brackish marsh according to Chabreck (1972). The bed was Ca. 20 m from the shoreline in water ca. 0.6 m deep. This location is about 11 Sipa 14(4):617. 1991. 618 km northeast of the last reported site for this species in Louisiana (Pointe aux Herbes). A subsequent investigation at this location on 18 August 1990 yielded fruiting specimens, and additional smaller beds of P. perfoli- atus. Voucher specimens have been deposited in the herbaria of Louisiana State University-Baton Rouge (LSU 78177, LSU 78178; Brantley & Platt s.n.) and University of New Orleans (NOLS 5378). The exact cause for the periodic occurrence of Potamogeton perfoliatus within Lake Pontchartrain is unknown. — Christopher G. Brantley and Steven G. Platt, Wettanda Ecologi- cal Services, PO. Box 701, Madisonville, LA 70447, U.S.A REFERENCES CHABRECK, R. H. 1972. Vegetation, water and soil characteristics of the Louisiana coastal region. eon State Univ. Agric. Exp. Sta. Bull. No. 664. 72 pp HAYNES, R. R. 1968. Potamogeton in a Proc. Louisiana Acad. Sci. 31: 82 — 90. LESTER, G. 1988. Plants and animals of special concern in the Louisiana coastal zone. Louisiana Nat. a Prog. Spec. Publ. No. 2. Louisiana Dept. Wildl. Fish., Baton Rouge, 291 5 MAYER, M. S. 1986. The Teas ae vegetation of the Lake Pontchartrain estuar- ine system, ae M.S. Thesis, University of New Orleans. 100 p MONTZ, G. N. mae Ps ne vegetation of Lake ee en Louisiana. Castanea i 115 — OGDEN, E. C. 1943. ae broad-leaved oe : at ad of North America north of Mexico. Rhodora 45:57 — 105, 119— 163, RECENT COLLECTIONS OF HELIOTROPIUM EUROPAEUM (BORAGINACEAE) FROM TEXAS — Heliotropium europaeum L. is an annual native to southern and central Europe, northern Africa, and regions of the Caucasus and Iran (Johnston 1960). In the United States, H. enropaeum has become sporadically naturalized along roadsides and in waste places from New Jersey to the Carolinas, southward to Florida, and west- ward to Texas and California (Small 1903; Radford et al. 1968; Correll and Johnston 1970; Fernald 1970; Munz 1968). The single historical record of the species from Texas was a 19th century collection from Hays County (San Marcos and vicinity, Spring 1897, Stanfield s.n. (NY)). In the fall of 1989 and summer of 1990, H. ewropaenm was collected at three separate locations in and around San Marcos, Texas. Two of the collec- tion sites were gravel terraces of the Blanco River, where the species was locally abundant and growing in association with Polanisia dodecandra (L.) DC., Sesbania macrocarpa Muhl. ex Raf., Exphorbia serpens Kunth, E. nutans Lag., Phyla nodiflora (Michx.) Greene, Leucospora multifida (Michx.) Nutt.., Sipa 14(4):618. 1991. 619 and Justicia americana (L.) Vahl. The third site was a disturbed flower bed on the campus of Southwest Texas State University in San Marcos, where only a few individuals were found. Voucher specimens: TEXAS. Hays Co.: west campus of Southwest Texas State Univer- sity, behind San Saba Hall, San Marcos, 12 Oct 1989, Hutzler 23 (SWT, TEX); 4.0 mi N of San Marcos on county road 140 at the Blanco River crossing, 17 Oct 1989, Hutzler 122 (SMU, SWT); gravel terrace of the Blanco River 0.1 mi upstream from the 1-35 hwy bridge, N of the city of San Marcos, 19 Sep 1990, Lemke 3327 (NY, SMU, SWT, TEX). Heliotropium europaeum is readily distinguished from all other Texas species of Heliotropium by its nutlets, which are typically 4 in number, 15-2 mm long and distinctly tuberculate on the abaxial surface. — Patrick L. Hutzler and David E, Lemke, Herbarium, Department of Biology, Southwest Texas State University, San Marcos, TX 78666, U.S.A REFERENCES CORRELL, D. S. and M. C. eae 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. 1881 pp. FERNALD, M. L. 1970. Gray’s Bene of botany. 8th ed. D. Van Nostrand Company, New York. 1632 pp. JOHNSTON, I. M. 1960. Boraginaceae. Pp. 123 — 221 In: Flora of Texas, vol. 1, C.L. Lundell, ed. Texas Research Foundation, Renner MUNZ, P. A. 1968. Supplement to a California dota University of California Press, Berkeley. 224 pp. RADFORD, A. E., H. E. AHLES, and C. R. BELL. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. 1183 pp. SMALL, J. K. 1903. Flora of the southeastern United States. Publsihed by the author, New York. 1370 pp. Stipa 14(4):619. 199 L. EDSKO ae DYKSTERHUIS (1908 — 1991) Dr. Edsko Jerry Dyksterhuis, age 82, of Bryan, Texas since 1964 died in Bryan on August 10, 1991 Dr. Dyksterhuis had beda professor of rangeland ecology in the Depart- ment of Rangeland Ecology and Management, Texas A&M University, 1964-1970 and was named professor emeritus in July 1970. He was born on December 27, 1908 and raised on a farm near Hospers, lowa. He received the BS degree in forestry and range management from lowa State University in 1932 and the Ph.D. in Plant Ecology and Soil Science from the University of Nebraska in 1945 Before joining Texas A&M University he had retired from the federal civil service after 10 years with the U.S. Forest Service and 20 years with the U.S. Soil Conservation Service. Following work with the University he was employed as Ecologist by Diamond Head Corporation of New Jersey; and later by the U.S. Department of State as Consultant on Natural For- ages entailing two trips to Turkey and Iran. Author of many scientific and popular articles, he received the Mercer Award of the Ecological Society of America for his monograph on the “Western Cross Timbers of Texas” and the authorship award of the U.S. Department of Agriculture for his “Savanna Concept and Its Use”. He is credited in Compton's Encyclopedia for the five pages on ecology. His field experience began as Range Examiner on National Forests of Utah, Arizona, and New Mexico and final lly included administrative, staff, and research positions ranging from Forest Ranger, and Supervisor's Assis- tant in charge of grazing on the Carson National Forest out of Taos, New Mexico to Sr. Forest Service Representative on Watershed Flood Control Surveys in Texas, Oklahoma, Kansas, Arkansas, and Missouri while with the Southern Forest Experiment Station in New Orleans. He then joined the Soil Conservation Service at Fort Worth as Range Conservationist for the Western Gulf Region and later served 15 years as Head Range Conservationist for the Northern Plains Region out of Lincoln, Nebraska, receiving the USDA award for Outstanding Leader- ship. During bricf leaves he had held visiting professorships at the State Universities of Montana, South Dakota, Kansas and Colorado, and served as first Extension Range Specialist for South Dakota. He was installed as President of The Society for Range Management in Calgary, Canada in 1968 having received the society’s highest awards as Sipa 14(4):620. 1991. 621 well as those of the Texas Section of that society. Asa result of developing a quantitative ecological approach to inventory and management of range- lands making it possible to quantify range degeneration and to predict potentials, his full biography appears in “World Who's Who tn Science; from Antiquity to the Present”. He was a Fellow of the Association for the Advancement of Science. Survivors include wife Margarett A. (Cox) Dyksterhuis, son Dr. Jerry E. Dyksterhuis, and daughters, Jantina Kay Clegg and Edna Leona Marge Selee, their spouses and eight grandchildren. — FE. Smeins, Texas AGM University, College Station, TX 77843. Edsko Jerry Dyksterhuis (1908 — 1991) Sipa 14(4):621. 1991. ANNOUNCEMENT RONALD L. STUCKEY INITIATES ENDOWMENT FUND FOR THE OHIO STATE UNIVERSITY HERBARIUM Ronald L. Stuckey, Professor of Botany at The Ohio State University, presented a gift of $30,000 to the University Foundation to initiate an endowment for the support of the University Herbarium. The presentation was made as a final surprise announcement at Professor Stuckey’s retirement party celebrating 26 years of teaching at the University. The event, held at the University Ramada Hotel, Olentangy River Road, Columbus, was atren- ded by 130 colleagues, former students, relatives, and close friends. They came from the central Ohio area, elsewhere in the state, and eight other states. Designated as the Ronald L. Stuckey Herbarium Fund, the gift was accepted by Drs. Tod E Stuessy, director of the University Herbarium; Ralph E.J. Boerner, chairperson of the Plant Biology Department; and Gary L. Floyd, Dean of the College of Biological Sciences. Anne K.P. Kochman, College representative to the University Foundation and Develop- ment Fund, eta for the Foundation by reading a letrer from its Executive Director, Donald D. wer. Dr. sees stated that the endowment was a “wonderful gift” that will aid in the studies of the fora of Ohio, which are of particular concern of the donor. Director Stuessy also praised Prof. Stuckey for his dedication, thoughtfulness, and genuine care for the future development of the Herbarium. Chairperson Boerner pointed out that the donor, who had worked so long and so hard for the students, cared enough to make this kind of gift to help future students. Dean Floyd referred co the donor's qualities of being sensitive, dedicated, committed, loyal, and generous. He wished the donor “good luck” in his second career. The establishement of the endowment fund for the University Herbarium not only marks the occasion of Dr. Stuckey’s retirement from teaching, but also commemorates the LOOth anniversary of the Herbarium. The fund creates a foundation for its future as a part of the Biological Sciences’ new Museum ap noe Diver Celebrating 100 years of continued as ation, The O hi > Scare University Herbarium was founded in 1891 by the University’s first Professor of Botany, William A. Kellerman, Ph.D. Initially the Herbarium was i. in Botanical Hall (site of the present-day Faculty Club Building) and moved in 1914 to the Botany and Zoology (B&Z Building), 1735 Neil Avenue. Beginning its second century of operation, the Herbarium will be relocated in the former food facility building (1315 Kinnear Road) now being renovated to house all of the eel collections in the University. Prof. Stuckey served as curator from 1967 through 9 Ronald L. Stuckey. Sipa 14(4):622. 1991. FLORENCE MONTGOMERY GIVENS (1933 — 1990) Florence Montgomery Givens was born 19 April 1933 in County Tyrone, Northern Ireland, but grew up in the United States. She graduated from high school in Sherrill, New York. [In 1958 she earned an associate degree in science (AAS) in Ornamental Horticulture and Biological Technology from the State University of New York, Agricultural and Technological Institute, at Farmingdale, New York. To earn money for college, she worked as a senior scientific assistant for the American Cyanamid Company in Princeton, New Jersey. She then was able to attend the University of Georgia at Athens where she obtained her bachelor of science degree in botany in 1964. She continued on there, undertaking postgraduate work under Wilbur Duncan. It was also at Georgia that she met and married Ray Givens. Her master’s degree was awarded in 1971 for her thesis “Vascular Flora of Echolls Mill Granitic Outcrop.” As a result of this work, Florence had an abiding love for these granitic outcrop areas of the southeastern states. From 1969 to 1972, Florence worked as a scientific assistant at the Academy of Natural Sciences of Philadelphia, Pennsylvania, doing curato- rial work and providing identifications of local plants. She also worked with Alfred Schuyler and Wayne Ferren on a survey of aquatic plants of the Delaware River and its tributaries. For the next four years, Florence was Sas as botanist at the Henry Foundation for Botanical Research in Gladwyn, Pennsylvania, where she performed curatorial work and conduc- ted tours for visitors. In 1976, Florence and her husband, Ray, moved to Louisiana where Ray took a faculty position in geology at Nicholls State University in Thibo- daux. Four years later, Florence began her association with the Department of Botany at Louisiana State University in Baton Rouge when she was chosen for the position of assistant curator of the herbarium, working under the direction of Lowell E. Urbatsch. One of her first duties was to organize Professor Clair Brown’s various plant collections which had been in storage since his death. Florence’s dedication to her botanical work went far beyond routine herbarium work. She made many collecting excursions to all parts of Louisiana, often to botanically interesting and unusual sites, thereby augmenting the general collections of the herbarium. These trips were often made in the company of her colleagues at Louisiana State University Stipa 14(4):623. 1991. FIG. 1. Field trip co St. Tammany Parish, Louisiana. ta to right: Florence Givens, ee Stones, and Shirley Tucker. Photograph by Lowell Urbats and other institutions. She and Ray also took many trips associated with Ray's geological research. During 1983 and 1984, she went on two trips to Costa Rica with geologists from Louisiana State University. Even though Florence's primary duty on these trips was to collect specimens for pollen analysis, she found time to collect many interesting plants. Over the years, Florence collected many fine specimens; these have included not only Howering plants and ferns but also lichens and mosses. At the time of her death, her collection numbers totaled over 5000 specimens, many of which are represented in the herbarium of Louisiana State University and, through exchange, in other herbaria. Another part of Florence's botanical work will have enduring signifi- cance. In 1977, the noted botanical artist, Margaret Stones, was engaged by Louisiana State University to prepare watercolor drawings of 200 plants for the Flora of Louisiana project. Spanning more than ten years, the final number of plant drawngs came to slightly more than 200. Florence became a close friend of Margaret Stones and provided interesting and unusual Sipa 14(4):624. 1991. FIG. 2. Florence at Leeds Castle, England, 1989. Photograph by Ray Givens. plants for her to draw. Approximately one fifth of the drawings are based on collections made by Florence alone or in association with others. The Stones’ watercolor drawings have received high praise and have been ex- hibited at the Smithsonian Institution in Washington, D.C.; three British institutions exhibited them in 1991. e year before Florence died, she and Ray travelled to Northern Ireland to see her native country. During this trip they also spent time in England where Florence was able to fulfill her dream of visiting the Royal Botanic Garden at Kew. There she saw in cultivation some of the plants — living representatives of the Flora of Louisiana project — which she had donated over the years. She was pleased to find that some of the plants, such as Trillium recurvatum and Pachysandra procumbens, had done very well in cultivation. Florence died on 15 November 1990 after a long struggle with cancer. She will be remembered by members of the Department of Botany here and by her colleagues at other institutions. Sipa 14(4):625. 1991. PUBLICATIONS AND PUBLISHED ABSTRACTS Montgomery, E 1. 1967. A new habitat and physiographic province for Botrychinm lunarioi- des. Amer. Fern J. 57:6—8. and D. Blake. 1969. Talinum mengesii Wolf from a new physiographic province. Bull. Torrey Bot. Club 96:719— 720. Givens, EM. 1982. Rediscovery of Ceratopteris in Louisiana. Proc. Louisiana Acad. Sci. 45:187. Givens, C. R. and E M. Givens. 1987. Age and significance of fossil white spruce (Picea glauca), Tunica Hills, Louisiana-Mississippi. Quaternary Research 27:283 — Givens, EF M. 1989. Review of Ceratopreris in Louisiana. Proc. Louisiana Acad. Sci. 51:62. — Alan W. Lievens, Marie S. Standifer, and Shirley C. Tucker, Department of Botany, Loutstana State University, Baton Rouge, LA 70803, U.S.A. Sipa 14(4):626. 1991. Gilbert Onderdonk. The Nurseryman of Mission Valley, Pioneer Horticulturist by Evelyn Oppenheimer. 1991. University of North Texas Press, PO. Box 13856, Denton, TX 76203; Fax (817)565-4590. Distributed by University Distribution, Drawer C, College Station, TX 77843. 200 pp, 16 b&w illus. $22.50 (cloth) ISBN 0-929398-24-6; $12.95 (paper) ISBN 0-929398-23-8. Pioneer Horticulturist in Texas and Mexico Gilbert Onderdonk, born in Sharon, New York, came to Texas in 1851. He was twenty-two. Soon he wrote home “I often forget that I was sick. . . chasing deer on horseback over the prairie.” He was working on a ranch on the San Antonio River about twenty miles from the Gulf of Mexico. At first the cattlemen opposed Onderdonk’s planting trees on what they considered prime range land. Years later he told David Fairchild that the cattle barons were the curse of Mission Valley, Victoria Co. U. PB. Hedrick called him “the only [fruitgrower} by profession in Texas before the Civil War.” Prof. L. H. Bailey said that he introduced the native Golden Beauty plum in 1874, and Samuel Wood Geiser, historian of Texas horticulture, wrote that Onderdonk did “very distinguished work on peach breeding and selec- tion.” Onderdonk wrote articles for Meehan’s Gardener's Monthly, and he received a bronze medal at the Louisiana Purchase Exposition in 1904. In his The World was my Garden David Fairchild tells of his visit with the “genial Dutchman” at his nursery. He wrote that “Onderdonk was a true plant enthusiast at heart.” Miss Oppenheimer has ransacked the Onderdonk family papers; records of the Old Dutch Church at Flatbush, New York; and university collec- tions in Texas and Delware, to give us a triptych “of a man to remember.” The first panel is a brief biography; the second, Onderdonk’s Mexican ex- periences in his own words; and his third, the most interesting for the his- torian of Texas horticulture, his account of his nursery business, as quoted from his Descriptive Catalogue of Mission Valley Nurseries for 1888, and from his Pomological Possibilities of Texas, published in Austin in 1911 when he was eighty-two. Besides a portrait of Onderdonk and reproductions of title pages, there are photographs of figs, grapes, and bananas, the last two from the farm of H. G. Stillwell in San Benito near Brownsville. Altogether this is another chapter in Professor Geiser’s call for “what shall someday be written — a comprehensive historical account of horticulture in Texas.” The author's comment is not distinguished by type font from the Onder- donk text, and there is no effort to identify plants mentioned. Some of Onderdonk’s early nursery sale items are especially interesting. For ex- Sipa 14(4):627. 1991. 628 ample, he was offering the native bignoniaceous Chi/opsis, “flowering willow,” and Ziziphus jujuba, “jajube.” We would have welcomed more on “umbrella China” — the Chinaberry, Melia azedarach, and in particular the appearance of the cultivar “umbraculiformis” reputedly of Texan origin dating from 1874. (What a fun-thing for some future essayist to document the history of this valued shade tree of the Southern States from its Persian (7) or Indian origins!). “Camphor trees” (p. 55) are mentioned, “reputedly brought to Louisiana during the 1840's” and noticed in Southern Garden in 1894 (Charlotte Seidenberg, The New Orleans Garden, New Orleans, 1990, p. 100). When did it come to south Texas? Onderdonk’s notes on his travels in Mexico will interest readers beyond horticulture. His trips were made on behalf of The U. S. Department of Agriculture in a search for overlooked fruits suitable for Texas orchards and home gardens. He is known to have brought back sixteen varieties of Mexican peaches. It is presumptuous, however, to say “no other American had reported on travel in Mexico except William Cullen Bryant in 1872.” For an example, Frederick Albion Ober, “a prolific writer” and orni- thologist, published Travels in Mexico and Life among the Mexicans (Boston, 1884). His Chapter 17 on Monterey, Saltillo, and the silver mines were described later by Onderdonk. (American Travellers Abroad by Harold E Smith (1969) is a reader's Baedeker). Fortunately Onderdonk’s family kept copies of the Cuero, Texas, newspapers that serialized his Mexican travels. These and his “private jo- urnal” have been excerpted, rarely with inclusion of year dates, but they tell us, for example, how it was to ride the Mexican National Railway in 1898. His enthusiasm comes through: “Such apricot trees!” and it is evident that he regretted that he passed the “bushes new and nameless to us” (P31). Now and then we may name his plants: his “red pepper trees” (p. 35) are easily identified as Schinus molle. There are some highly readable paragraphs: “you may never before then have felt that you would hug a burro. . . Oh! How you love that burro.” You will learn about a cayuse, how Catorce was named, the “grandest” wilderness of mountains he had ever seen; and there is an index for the curious. — Joseph Ewan, Missouri Botani- cal Garden, PO. Box 299, St. Louis, MO 63166-0299, U.S.A Sipa 14(4):628. 1991 ANNOUNCEMENT OPENING OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS, INC. IN FORT WORTH FORT WORTH, TEXAS — When the Botanical Research Institute of Texas, Inc. (BRIT) opened Friday, October 4, 1991, individuals and organizations as well as the scien- tific community were provided access to one of the nation’s 25 National Resource Centers, featuring an herbarium with a collection of more than 450,000 dried and pressed plant specimens representing most of the earth's plant families, and a botanical library with more than 30, 000 volumes. is anonprofit international botanical resource center organized in 1987. It serves as and professionals and a primary interpreta- a primary plant information center for scientists tion center for people interested in learning more about the plant world. Explaining the importance of BRIT, George Sumner, chair of BRIT’s 15-member board of trustees, said, “The human race ts totally dependent upon plants for existence. We must have information that BRIT provides about the identities, characteristics and requirements of organisms to sensibly manage this vital resource we take for grantec ward P. Bass, vice chair of BRIT’s board of trustees, said, “These are stgine times for brimary resource to help us —e our earth, and more than ever the plant world serves as a understand the changes occurring and seek solutions for improving our environment. Fortunately, BRIT's facilities are available to support everyone interested in accomplishing these goals. In size, BRIT is in the top 25 of the nation’s 628 collections. Significantly, BRIT’s research facility, together with the Fort Worth Botanic Garden's display gardens and educa- tional programs, places the Metroplex among the nation’s leading botanical cente s collection of specimens, books and periodicals — valued at more a $10 million — are primarily the Lloyd H. Shinners’ Collection started in 1943 at Southern RIT. Methodist University, which the University has placed on permanent loan Periodicals published by BRIT include Szda, Contributions to Botany, and Sida, Botanical Miscellany. Scientists and other professionals from thoughout the world regularly borrow the moun- iest dating back to 1791 — for study and comparison. In- — ted specimens — the ear and pe cae ae using BRIT include teachers, students, plant enthusiasts, mers, businesses, governmental agencies, dividuals veterinarians, physicians, ranchers and farm museums, ace oe. hospitals and poison control centers. BRIT is open Monday through Friday, 9 a.m. to 5:00 p.m. Appointments are encoura- ged. As a nonprofit organization, BRIT is funded through tax-deductible contributions. Director: Wm. E Mahler Executive Director: Andrea Pistorius McFadden Curator: Barney L. Lipscomb Sipa 14(4):629. 1991 Eprror’s Notre AND REVIEWERS FOR VOLUME 14 SiIpA, CONTRIBUTIONS TO BOTANY has a new home with the Botanical Research In- stitute of Texas, Inc. (BRIT) in Fort Worth. In 1962 Lloyd H. Shinners started the publica- tion at Southern Methodist University (SMU) as a private journal. Even after Lloyd's death in 1971 Wm. E Mahler continued privately publishing Sina at SMU up through 1990. Beginning with this issue Sipa is now a publication of BRIT. This brings to close an era of — some 30 years as a privately published journal. May the next 30 years at BRIT serve the — botanical community even better. The following individuals have kindly supported Sipa through their time and efforts in reviewing manuscripts submitted and/or published in volume 14, 1990 — 1991. Without your interest and support, Siba would not be the journal that you have come to expect. I do not take reviewers for granted. Your support is vital and very much appreciated and with ene support SIDA can remain a top quality journal of systematic botany. Sipa’s subscription base continues to expand each year with subscriptions approaching 800 in 80 countries. Thanks to all authors, reviewers, subscribers, and readers for your continued interest and support. Anderson, Loran C. Johnson, Ann E Popenoe John Ayers, Tina J. Johnson, poe E. Powell, A. Michael Bates, Vernon Keener, Carl S. Pringle, James S. Beaman, John H. Kiger, Robert W. Pullen, Thomas M. Beauchamp, R. Mitchel Koch, Stephen D. Rabeler, Richard K. Beetle, Allan A Koutnik, Daryl L. Rembert, David H. Bruederle, Leo P setae Thomas G. Reznicek, Anton A. Bryson, Charles T. Lane, edith A. Rogers, George K. Burandt, Charles ane Michel G. Rowell, Jr, Chester M. Carter, Richard Lewis, Walter H. Schultz, Leila M Castaner, David Lonard, Robert Simpson, Benny J Cholewa, Anita FE Luteyn James L ith, Ale Clewell, A.E Mason, Jr., Charles T. Smith, Edwin B. Conese, Barry McDearman, Will Smith, James P Cranfill, Raymond McDonald Andrew Stuckey, les L. Crins, William J. McDonald, seer Taylor, Joh Dennis, W. Michael McKenzie, Paul Thieret, ne W. Denton, Melinda FE McKinney, eee E. Thomas, R. Dale Dickinson, Timothy A. McVaugh, - Bers Thompson, Rahmona A. Diggs, George Mickel, Jol nit Timme, Steve L Eckenwalder, = E. Mitchell, an By Tucker, Gary Ertter, Barbar Moore, Michael O. Utech, Fred H. Ewan, Joseph Montz, Glen A tee 2. a A. Fletcher, Reggie A. Morden, Clifford W. Wain Gandhi, K.N. Morgan, David Warnock, cae acy. Godfrey, R.K Naczi, Robert EC. Whi aker, Thomas W Hall, David W Nee, Michael Wilbur, vert | Hartman, Ronald esom, Windham, Michael ayden W. John Nicolson, Dan H. Worthington, Richard D. Henderson, Doug Norquist, Cary Wunderlin, Richard P Hill, L. AMchnel Peterson, Paul M. Holmes, Walter C. Pinkava, Donald J. Sipa 14(4):630. 1991. INDEX TO VOLUME 14 Names of contributing authors are in CAPITAL letters. New names and combinations are in boldface. TITLES OF ARTICLES A natural intergeneric hybrid in the x = group of the Astereae (Asteraceae) by ynald L. Hartman and Meredith A. Lane 321— 329. s of Acourtia (Asteraceac- Mutisieae) from southern Mexico by 144. Le A new species of Cheslanthes (Adiantaceae) from northeastern Mexico by John M. Mendenhall and Guy L. Nesom 551-554. A new species of Conradina (Lamiaceae) from northeastern peninsular Florida by Robert Kral and Robert B. McCar- tney 391— A new species of Ibervillea See ae from western Mex xico by Rafae a — the Carolina ae by Joa M Stucky and Milo Pyne 189 — 20 A new species of Vrola ergs = the Guadalupe Mountains, Trans- Pecos Texas by A. Michael Powell and Brent Wauer 1—6. A new status for Quercus shumardii vat. ese ace by Nick Stoynoff William J. Hess 267 — 271. A See note on Eupatorinm fistu- losum (Asteraceae) by Kancheepuram N ndhi and Paul A. Fryxell 129: 132; A note on the geographic range of Anten- naria aromatica Evert (Asteraceae: In- uleae) by R. J. Bayer 505 — 507. A note concerning the typification of two plants described from Texas by Laur- ence J. Dorr 309. A report of a grayloides and Cyperus retroflexus (Cyperaceae) new to Mis- sourt and notes on other selected Mis- sourt Cyperus by Rick se — and Charles T. Bryson 475 — Sipa 14(4):631—635. 1991. A revision of Paxistima Snes by Ann M varo and Will H. Black- well 231—249. A taxonomic comparison of Aristida ternipes and Aristida hamulosa (Gram- ineae) by Jonathan S. Trent and Kelly W. Allred 251— 261. Additional notes on the Louisiana by Kancheepuram Dale Thomas Asteraceae of Additional seconds of Cyperus entrerianus (Cyperaceae) in the Unite s | Richard eae and Stanley D. Jones 615-616 Additions to the flora = rede by R. , Edw . Smith, Eric onal Philip E. nas and Carl Amason 483 — 491. Additions to the flora of pie from El Paso. County op ichard D Worthington 135 — An explanation for the ne in the chromosome count of the redbud (Cercis canadensis, Leguminosae) by Will H. Blackwell 7-11. Annotated ania of Arizona Con- by Daniel E Austin . ae 443 7 Annoti a checklist of New Mexican Convolvulaceae by Daniel E Austin — 286 Bisedal Sida (Malvaceae) a Charles L. Burande, Jr. 405 — Brachiaria plantaginea, oe cylindrica, and Panicum maximum. three grasses osceen new to Louisiana a extension for Rei bine ae by Cl eek M. Allen, R. Dale and Michel G. Seer 615.615. Carex bale (Cyperaceae) new to Mis- with notes on Carex picta and Carex impressinervia in Mississippi by 632 — T. Bryson, Sam W. Rosso, and Robert R.C. Naczi 493 — 499 ba comosa (Cyperaceae), new to Mis- SISSIPpi ae T. Bryson and Stanley D. Jones 311— 312 Chloris inflata esas) new to ‘Teuiian : . Macgregor and Charles M. Allen 313. Circumscription of Agarista boliviensis (Ericaceae) by Walter S. Judd and Paula M. Hermann 263 — 266. Classification and systematics of eastern North American Vitis L. (Vitaceae) as of Mexico by Michael O. Moore 3) 367 > Common names for vascular plants: guidelines for use and application by John T. Kartesz and John W. Thieret 421 — 434. Correct names for _ the varieties of Carex emmonsit by J. Rettig C ratae gus Secreta (Rosaceae), a new species of hawthorn from t Edwards Plateau, Texas by J.B. Phipps lor 19, Cyperus entrerianus ir Lene an over- pa spe in temperate North Am ie Richard ¢ Carter 69 —77. Deletions pee restorations in the flora of southern Florida by Daniel B. Ward and Frank C. Craighead 287 — 304, Documented aes 1990: : iscellaneous American ee plants by Bruce D. ae Donald J. Pinkava, Debbie ickel, Dan Fillipi, Beth Eggers, and ee 1 J. Keil 305 — 308 Documented chromosome numbers hromosome numbers in Hyhanthus (Violaceae) by B.L. ae and Linda K. Escobar 501 = 90 Edsko Jerry Dykster! quis (1908 — ay by EE. Smeins 620-621. Euphorbia Gi ee (Euphorbiaceae), ec Acutae by Mark Mayfield 5 i a lathyris phorbicen n new for Texas re J 609 — *"Kennon Extension of native range of Sabal mexicana (Palmae) in Texas to include Central Coast by Landon Lockett and Robert W. Read 79— 85. Florence Montgomery Givens (1933 — 1990) by Alan W. Lievens, Marie S. Standifer and Shirley C. Tucker 623 — 626. Gentianella canosol ica ak a new ango, Mexi acd grass (Sclerochloa dura, Poaceae) in the United States by David Brandenburg, James R. ae aid John W. Thieret 369 — 3 ee ae of the plants ie and cribed in Catesby’s Natural History Carolinas, Florida and the Bahamas by Robert Wilbur 29 — 48. Koeleria eae! (Vill.) Shinners Nae new to Louisiana by Charles M. Allen Harland D. peau and C oe H. Stagg 133 — 134. Leaf venation studies in Indian Sida Malvaceae) by A.M. Saibaba and S. Raja Shanmukha Rao 215 — 222. Lectotypification of Laciniaria carinata (Aste bees : Milo Pyne and Jon M. Stucky 13, Lemma sj eens am in the Eragros- tideae (Poaceae) sus V; s Sone: L. Hatch > a 25 a Reyna and ISL 546 Lycianthes wel (Solanaceae), new to North America by Stephen P. Darwin ind Toby Feibelman 605 — 606. Montia linearis (Portulacaceae), new Mississippi by Richard K. as in Charles T. Bryson 310 —31 Morphological characters as ne of rubber content in Guayule aa argentatim — ee by M. Foster, S.E. Gabel . Ward, L i Kile: ine, PK, “MeCann, and Jaroy Moore — 33 Noteworthy ae from north Florida. V. oy Loran C. Anderson 467 — 474, Observations — on pygmaea Malvaceae) by Paul A. ve ell and Jesus Valdes R. 399 — 40 Occurrence of Potamogeton perfoliatus L. (Potamogetonaceae Louisiana by Christopher G. Brantley and Steven G. Platt 617-618. Paliurus spina-christi (Rhamnaceae) new America in Texas by Robert J. O’Kennon 606 — . at chartacea = SSp. minima yophyllaceae): a new subspecies of a rare Florida endemic by Loran C. 441. vae Mexicanae: Sedum eee (Crassulaceae) by J. An- w McDonald 315-31 ee of the genus Cinna (poieeae) by David M. Brandenburg, Wt oe and John W. Thieret Plantae alpinae no 596. Royo gee (Cyperaceae), new s by Stanley D. ae and aes D. ie 134— Seedling morphology in ae ene and its taxonomic im- piications by Frederick B. Essig D0 B00. S yphoricarpos aes (Caprifoliaceae), y Benny J. Simpson Synopsis of Carex section Lupulinae Jones and Steph Synopsis of the Mexican and Centra American representatives of Lobelia section Tylominm (Campanulaceae: aad e) by Robert L. Wilbur eae: notes on western America Gentianaceae by James S. peel Lip = IST: Taxonomy of Digitaria section Aequiglumae (Poaceae: Paniceae) by vo and Stephan L Hatch 145 — The alpine- xen flora of northeastern Mexico by J. Andrew McDonald 21— 28. The — blue-eyed-grasses ee Iridaceae) of Arkansas Pid Kathleen L. Hornberger 597 — 604 633 The Clematis virginiana (Ranunculaceae) e southeastern United States by Frederick B. Essig 49 — 68. The genus Hosta Tratt. ee in Korea by ee Gi Chung and Jong Won Kim 411—420 The herbaceous oe of three Weches formation outcrops in eastern Texas by eee George and Elray $. Nixon Ly = The oe naturalization of Péstacta chinensis ee ae in east Texas Ed McWilliams 508 i The Piiahee aeee in the flora of the southeastern ee States by James L. Luteyn 169 — The rediscovery - on aes (Cyperaceae) in Texas by Stanley D. Jones, J.K. vee and Gretchen D. Jones 511-31 Two new Vitis ae from mountai- nous Mexico by Barry L. Comeaux 459 — 4660. Two weedy species, Ammoselinum butleri (Umbelliferae) and Lepidium austrinum (Cruciferae), new to ae by Charles T. Bryson 506 — 508 Typification of Vernonia fehl Small and V. jamesit Sac & Gray (Compo- ce by vid E. Boufford yf ke Ulmus ee (Ulmaceae) naturalized in by Max E. Medley and John Ww. Thieret 613. especie de oe eee Tulipeae) del Sur de Méxi by Abisai Garcia- Meu. Xylothamia (Asteraceae: Astereae), a new nus related to Exthamia by Guy Nesom, Youngbae Suh, Dav id. R. Morgan, and Beryl B. Simpson 10L—116 634 Acacia angustissima var. hirta 467 Acourtia ovatifolia 141 Agalinis homalantha 490 Agarista boliviensis 263, eucalyptoides 263, paraguayensis 263 Amaranthus arenicola 484 Ammoselinum butleri 506 Amorpha herbacea 46 Amsonia tomentosa 306 Antennaria aromatica 505 Anthemis arvensis 467 Archiclematis 379 pre ane oe 307 Aristida ternipes 25 | r. hamulosa 260 Arnoglossum desiblun 470 Artemisia vulgaris 468 7 aN Asclepias rubra 4 Aster spinosus 514 Baccharis sarothroides 136 Berlandiera lyrata var. monocephala 306 Bothriochloa ischaemum_ var. songarica 468 Brachiaria plantaginea 613 Bulbostylis ciliatifolia 487 Callirhoe leiocarpa 309 Calochortus balsensis 525 Calystegia longipes 443, macounii 274, 444, pubescens 285, sepium 274, ssp. angulata 444 Carex albicans var. albicans 133, var. australis 133, var. emmonsii 133, baltzellii 470, 493, comosa 311, gigantea 87, S11, hirtifolia 487, im- pressinervia 493, intumescens 87, leavenworthii 468, louisianica 87, lupuliformis 87, lupulina 87, picta 493, seorsa 470 Centella asiatica 484 Cercis canadensis 7 Cerastium semidecandrum 468 Chaenactis stevioides 306 Cheilanthes hintoniorum 551 Chloris inflata 313 Cicuta maculata 468 Cinna 581, arundinacea 585, bolanderi 390, latifolia 586, poaeformis 590 Cirsium drummondii X C. wheeleri 306 Cladium mariscoides 468 Clematis 377, addisonii 380, alpina 380, aptifolia 380, aristata 380, baldwinii 380, barbellata 380, ere 380 buchaniana 380, campanuliflora 380, canadensis 64, catesbyana 49, 380, chinensis 380, chrysocoma 380, cir- rhosa 381, cordata 60, crispa 381, denticulata 381, drummondii re filamentosa 381, flammula 381, fusca 381, gentianoides 381, ieee 381, grata 381, heracleifolia 381, hexapetala 381, hirsutissima 381, holosericea 65, integrifolia 381, in- tricata 381, kirilowi lasiantha 381, leschenaultiana 381, ligustici- folia 381, macropetala 381, mandshu- microphylla napaulensis rica 381, micrantha 62, 381, miussouriensis 65, nz 381, orientalis 381, papuasica 381, patens 381, peterae 381, pierotii 381, pitcheri 381, ranunculoides 381, recta 381, rehderiana 381, reticulata 381, serratifolia 38 Sauer 381, texensis 381, viorna , vitalba 381, viticella 381, virgi- 1, tangutica 381, a 49 bee psis 379, anethifolia 381, scabi- osifolia var. kirkii 381, villosa 381 Cleome gynandra 486 Conopholis americana 470 a 391, grandiflora 393, etonia Coavelvuilus arvensis 275, 445, var. helleri 352, equitans 275, 445 Conyza bonariensis 484 Crataegus secreta 13 Cressa truxillensis 276, 445 Crotalaria angulata 488 Curcuma zedoaria 468 Cuscuta coryli 486, obtusiflora — var. glandulosa 487 Cycladenia humilus 306 Cyperus acuminatus 72, croceus 475, dis- tinctus 72, deocstaondil 71, echinatus 468, entrerianus 69, 615, eragrostis 72, grayioides 474, hystricinus 487, luzulae 69, X mesochorus 475, ochr- aceous 72, pseudovegetus 72, reflexus 72, retroflexus 474, surinamensis 73, irens 71 Decumaria barbara 490 Desmodium cuspidatum var. longifolium 488 Dichondra argenta 277, 446, brachypoda 277, 446 es 145, aequiglumis 150, con- vens 152, costaricensis 153, ie 154, ekmanii 156, erios- ta i ya 157, lanuginosa 158, leucites 59, pauciflora 161, sabulicola 162, texana 163 Dioclea multiflora 470 Drosera filiformis 47 1 Eleocharis flavescens 487, rostellata 468 Ericameria 101 Eryngium hoo Eupatorium Retican 129, glaucescens 514, verticillata 129 Euphorbia 573, acuta 577, angusta 577, johnstonii 574, lathyris 609 Euthamia 101 Evolvulus alsinoides — var. pane 277, 447, arizonicus 447, 278, nuttallianus 278, 448, sericeus 30. 448 Filago californica 136 Fryxellia pygmaea 399 Fuirena simplex var. aristulata 487 Gentiana newberry var. tiogana 186 Gentianella canosoi 227 Geranium texanum 488 Heliotropium europaeum 618 Hybanthus attenuatus 302, aurantiacus 502, bilobus 502, calycinus 502 communis 502, cymulosus 502, en- neaspermus 502, epacroides 502, floribundus 502, monopetalus 502, parviflorus 502, verticillatus 502, volubilis 502 Hedyotis nuttalliana 471, purpurea 469 Helenium arizonicum 306 Hibiscus coccineus 489 Hosta 411, capitata 416, clausa 413, jonesii 418, minor 415, taquetii 415, yingeri 4 635 Hydrochloa caroliniensis 489 Hypericum denticulatum 488 Ibervillea maxima 223 Imperata cylindrica 613 Indigofera spicata 469 Ipomoea alba 285, barbatisepala 280, 449, capillacea 449, cardiopylla 280, 449, costellata 281, 450, crisculata — 281, , dumetorium 2 hederacea 282, 450, tu vee 283, ae a 283, 451, ha , lindheimeri 283, ye see ia . plummerae 283, 452, pubescens 284, 452, purpurea 284, 453, tenuiloba 285, 453, thurberi 454 Iris hexagona 4 Isocoma veneta 321 Jacquemontia agrestis 455, pringlei 455 Juncus gymnocarpus 47 1 Keckiella antirrhinoides ssp. microphylla 307 Koeleria gerardii 133 Laciniaria carinata 209 Lepidium austrinum 506, latifolium 136 Leptochloa scabra 616 Liatris 189, carinata 209, cokeri 205, earlei 206, ne var. gram- inifolia 207, — var. malli 207, regimontis 189, 209, as 207, spicata 207, squarrosa 206, tenuifolia 207 Limonium 170, carolintanum = 171, limbatum 17 Lithospermum incisum 306 Lobelia 555, calochlamys 562 , dressleri 557, guatemalensis 564, nubicola 566, let ga 558, tatea 559, zelayensis 563 Ludwigia erecta 471, lanceolata 471 Lycianthes pee ae 605 Machaeranthera grindelioides 306 s 307 Marah gilens Marshallia a ee 484, graminifolia ssp. uifolia 515 Matelea ere 471 Matricaria discoidea 516 Mentha arvensis 488, var. villosa 307 Mirabilis jalapa 489 636 Montia linearis 310 Murdannia keisak 469 Myosotis discolor 485 9 Naravelia 37 Nephropecalum ea 309 Neptunia pubescens 4 Opuntia repens 307 Paliurus spina-christi 606 Panicum maximum 613 Paronychia cha Pp rtacea ssp. minima 436 arthenium argentatum 33 aspalum minus 469 Paxistima 231, canbyi 244, myrsinites ssp. myrsinites 240, ssp. mexicana 243 a P Penstemon eatonii 307, ssp eatonii X P. pseudospectabilis var. connatifolius 308, pseudospectabilis — var. natifolius Pennellia longifolia 306 Physostegia godfreyi 471 Pinckneya bracteata 47 | Pistacia chinensis 508 Pluchea oblongifolia 472 Plumbago 174, auriculata 178, scandens 174 Potamogeton perfoliatus 617 Prunus caroliniana 489 Psoralea mephitica 307 s acerifolia 268, shumardii 267 Ranunculus marginatus var. trachycarpus 466 Rhexia salicifolia 472 Rhynchospora capillacea 134, crinipes 472 Rhus aromatica var. pilosissiine 305 Rocttboellia cochinchinensis 6 Sabal mexicana 79, palmetto 79 Sclerochloa dura 369, 372 Les) Sedum chrysicaulum 315, parvum ssp. nanifolium 315 Sida 215, acuta 217, cordata 217, grewl- oides 217, mysorensis jatrophoides — 405, 7, palmata 405, rhombifolia var. rhombifolia 217, var. retusa 217, schimperiana 215, 217 spinosa 217 Silene virginica 469 Sisyrinchium 597, albidum 600, angusti- fohum 600, — atlanticum 601 601, langloisit 601 rosulatum 601, sagittiferum 602 Smilax ecirrhata 489 campestre Solanum pseudocapsicum 469 Solidago drummondii 485, 485, rugosa 516 Soliva mutisii 4 ludovicitana Stachys hyssopifolia var. lythroides 472 Stellaria pallida 486 Stephanomeria exigua var. exigua 306, pauciflora 306 jes Stylocline micropoides 136 Swertia 179, fastigiata ms umpquaensis 85 Symphoricarpos occidentalis 512 Thamnosma montana 307 Imus parvifolia 61 Uvularia sessilifolia 472 Vernonia altissima var. marginata 570 jamesit 569, 570, marginata 570, var. tenuifolia ee a 569, 570 Vicia pulchella Viola evadapensis nuctalit 1, valli- cola Vitis se 459, acerifolia 359, aestivalis 347, var. bicolor 347, var. lincecumiti 348, bloodworthiana 460, cinerea 351, var. baileyana 352 351, var. hellert 352, champinit 362, X doantana 363, jaegeriana 464, labrusca 356, monticola 355, ve aaa 358, X novae-angliae 364, palmata 354, riparia 360, rotun- ‘ifoli lia 344, var. munsoniana 345, rupestris 361, shuttleworthii 357, vulpina 353 Wahlenbergia marginata 486 Xanthocephalum humile 321 Xylothamia diffusa 109, johnstonii 10, palmeri 110, parrasana 111 pseudobaccharis 112, urpusii indit 113, triantha 113 3 , var. floridana A Xyris longisepala 47 Sipa Guidelines For Contributors SIDA, CONTRIBUTIONS TO BOTANY is an international journal of systematic botany containing primary research papers sensu lato, including anatomy, biogeography, chemota- xonomy, Cladistics, ecology, floristics, epee ceva alan, numerical taxonomy, paleobo- tany, and palynology. 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Latin Descriptions or Diagnoses A description or diagnosis of a new taxon should mention the taxonomic characters that distinguish the taxon from its allies. These diagnostic characters should be éta/ized in the description. Since a diagnosis of a new taxon only refers to the taxonomic characters dis- a) it from other taxa, no alization in the manuscript is necessary. The ICBN (Rec. 32B.1) recommends that the main points in which the taxon differs from it its allies should be mentioned and italizing (ms) these in the description is in accordance with that recommendation. Sipa 14(4):638. 1991. Figures A scale should be in or on the illustration or photogra reduce the scale and subject proportionately. When this returned with the galleys for the author to determine the magnification of the final reduc- tion. In half-tones, sharp glossy photographs with good contrast are necessary for good ph and reduction will automatically is is not feasible, the proofs will be reproduction. References ‘his includes all of the literature cited in the text and may include other article citations the author may deam desireable. Only the first letter of the initial word, proper nouns, anc adjectives of titles should be capitalized. For journal abbreviations, use Botanico-Periodicum- jes Huntianum. Abbreviations When using the following abbreviations, the period is omitted. Distance: mm, dm, cm, m, km, ft, mi; directions: §, N, E, W, (in caps); months: first three letters only, Jan, Feb, etc. Example: Potter Co.: 2 km W of Dot, 5 Jun 1971, Smith 118 (SMU). Notes When references are few in number, cite them in the text and omit reference section. Text citations of literature are in parentheses and should state the journal abbreviation, volume, vages and year. The citation of a book includes the last name of the author, if not mentioned previously, the full unabbreviated title of the book, and the year published in parentheses. Documentated Plant Chromosome Numbers The year and number of each is assigned by the editor. Refer to Sipa 14(3):501— 503. 91. 199 (Continued from front cover) NOTES. Lycianthes asarifolia (Solanaceae), new to North America. 605 — Paliurus spina- christi (Rhamnaceae) new for North America in Texas. 606 — Euphorbia oe (Euphorbiaceae) new for Texas. 609 — Ulmus parvifolia (Ulmaceae) naturalized 1 Saas 610 — Brachiaria hee Imperata cylindrica, and Panteum max- imune, three grasses (Poaceae) new to Louisiana and a range extension for Rotthoellia nh. 613 — Adc licionall records of Cyperus entrerianus (Cyperaceae) in the United States. 615 — A firse report of Leptochloa scabra Nees (Poaceae) from Alabama. 616 — Occurrence of Potamogeton perfoliatus L. (Potamogetonaceae) in Louisiana. 617 — Recent collections of Hedsotropium exropaenm (Boraginaceae Texas. O18 from ~~ Edsko Jerry Dyksterhuis (1908 — 1991) 620 Florence Montgomery Givens (1933 — 1990) 623 Reviewers for volume 14 630 Announcements 622, 629 Book reviews 530, 550, 568, 571, 572, 579, 596, 627 Index to volume 14 631 Guidelines for contributors 638