nt eo © = © = a = ee ae) 4 ~ 8 = Y > A ~ wm — ~ 5 Lee Q, iS) = d 5. a Y 2) ~— Le) S = S ~ =) So Lead S SS AD) ~ ~ S ~ ~ ~ iS) ~ ~ = ~ ~ = phytogeographical and ecological publication www.phytologia.org Vol. 89, No. 3, pp 239 - 394 December, 2007 PHYTOLOGIA (ISSN 00319430) Phytologia, a journal for rapid publication in plant systematics, phytogeography and vegetation ecology, is published three times a year. Managing Editor Executive Editor micro-Molecular, Phytochemical — General systematics and Multivariate Systematics and evolution Robert P. Adams Billie L. Turner Baylor University University of Texas at Austin Baylor-Gruver Lab Austin, TX 78705 Gruver, TX 79040 billie@uts.cc.utexas.edu Robert_Adams@baylor.edu Associate Editors Nomenclature General Systematics Guy Nesom A. Michael Powell Botanical Research Dept. of Biology Institute of Texas (BRIT) Sul Ross State University Ft. Worth, TX, 76102 Alpine, TX, 79832 gnesom @brit.org ampowell @sulross.edu Macro-Molecular and Ethnobotany Phylogenetic Systematics Martin Terry Andrea E. Schwarzbach Dept. of Biology Univ. of Texas at Brownsville Sul Ross State University Dept. of Biol. Sciences Alpine, TX, 79832 Brownsville, TX, 78520 mterry @sulross.edu andrea.schwarzbach@ utb.edu Secretary-Treasurer - Subscriptions Robert P. Adams Phytologia, Box 727, Gruver, TX 79040 Robert_Adams@baylor.edu Copyright 2007 Phytologia. Texensis Publishing, Gruver, TX LuESTHER T. MERTZ LIBRARY Phytologia (December 2007) 89(3) 239 UEL WR ZUU/ Phytologia NEW YORK BOTANICAL GARDEN Contents D. S. Seigler, J. E. Ebinger and A. Kerber. Characterization of thorn- scrub woodland communities at the Chaparral Wildlife Management area in the South Texas plains, Dimmit and La Salle counties, Texas.... B. L. Turner. Melampodium moctezumum (Asteraceae: Heliantheae), a new species from Sonora, MERIC iii Ma ie sedivc esi wnvese vacee 258 R. P. Adams. Juniperus maritima, the seaside juniper, a new species Mem rupet Sound, North America... :..)....4046 3 s0steceteteseenecoanae ewes 263 M. H. MacRoberts and B. R. MacRoberts. Eryngium heterophyllum (Apiaceae) and Eriocaulon lineare (Eriocaulaceae) deleted from the MERE NOT OE LRP ae 8 rt te oe 282d eae ce cade whe vc wa dle des eee Noma ee 284 J. L. Reveal. A new variety of Eriogonum nudum (Polygonaceae) from SRNUTORNE NC DCU AIM seas Oh sdicceee ale acted ca ceateu ta dere Pdessan ten Opeeuaee ss 287 B. L. Turner. Biological status and distribution of Thelesperma jlavodiscum (Asteraceae: Coreopsideae)... 2.2... .0..22.00...deeeevetes sees. 290 D. B. Ward. Keys to the flora of Florida 17, Ruellia (Acanthaceae)..293 D. B. Ward. What in the world did Thomas Walter mean by Xxxxxx eyyyy? Part Two: The quite doublfulls: .. cde. cs..1 oe sien iis ccs 300 B. L. Turner. Heterosperma xanti, transferred to the genus Bidens Rm SIR ESE He item SNSAE). Po5: 055s cdacobackank vet ade s eat boheme vues 315 T. C. Philipps, S. B. Walker, B. R. MacRoberts and M. H. MacRoberts. Vascular Flora of a longleaf pine upland in Sabine County, Contents continued on next page 240 Phytologia (December 2007) 89(3) Contents (continued) M. Athar, A. H. Chaudhary, Z. Yousaf and S. M. Sabbir. Taxonomic reflections on the parasitic angiosperms of Pakistan..................... 339 B. L. Turner. Xanthisma spinulosum var. austrotexanum (Asteraceae: Astereae), An endemic of southernmost Texas..........2.--..-.s00cs000 349 B. L. Turner. A new species of Viguiera (Asteraceae: Heliantheae) » from Oaxaca: MexiGes. cock scant cc ano eeee aah ee a ee es 353 J. L. Panero and V. A. Funk. New infrafamilial taxa in Asteraceae...356 R. P. Adams, A. E. Schwarzbach, J. A. Morris and M. Socorro Gonzales Elizondo. Juniperus compacta (Cupressaceae),a new species fr@ina MIEKICG bse crs ae ieee re ran Rit ne ll ey 361 G. L. Nesom and B. L. Turner. Taxonomic review of the Xanthisma Spinulosum complexx Asteraceae: Asiereae).. 17 1..es00se ce ene 371 Index to article topics and scientific names in volume 89.................. 390 Phytologia (December 2007) 89(3) 241 CHARACTERIZATION OF THORN-SCRUB WOODLAND COMMUNITIES AT THE CHAPARRAL WILDLIFE MANAGEMENT AREA IN THE SOUTH TEXAS PLAINS, DIMMIT AND LA SALLE COUNTIES, TEXAS David S. Seigler Department of Plant Biology, University of Illinois, Urbana, Illinois 61801 seigler@life.uiuc.edu John E. Ebinger Emeritus Professor of Botany, Eastern Illinois University, Charleston, Illinois, 61920 jeebinger@eiu.edu Angela Kerber Department of Natural Resources and Environmental Sciences, University of Illinois, Urbana, Illinois 61801 akerber@v3co.com (Present address: V3 Companies, Ltd., 7325 James Ave, Woodridge, Illinois, 60517). ABSTRACT The composition of semiarid grasslands in southwestern North America has changed during the past 150 years with woody species increasing in density and cover. This brush encroachment involves mostly native species, many of which are woody members of the Fabaceae (legume family) that have increased in importance because of changes in local biotic and/or abiotic factors. At the Chaparral Wildlife Management Area, Dimmit and La Salle counties, Texas, thorn-scrub woodland communities are common. In some, Prosopis glandulosa (honey mesquite) is the most important species, however, other thorny woody legume species are prominent, and sometimes dominate the community. Common taxa are Senegalia berlandieri [=Acacia berlandieri (guajillo, fern acacia)], S. greggii [=Acacia greggii (catclaw acacia)], S. roemeriana [=Acacia roemeriana (Roemer’s acacia)], Vachellia bravoensis [=Acacia schaffneri var. bravoensis (twisted acacia)], V. farnesiana [=Acacia farnesiana (huisache)], and V. rigidula [=Acacia rigidula (blackbrush)]. In the communities studied, thorny woody legume species had importance values between 87 and 157 242 Phytologia (December 2007) 89(3) (possible 200). Between 11 and 20 woody and succulent species were present in these communities with a total of 28 species recorded. KEY WORDS: Acacia s./., Prosopis glandulosa, South Texas Plains, thorn-scrub woodlands. The composition and structure of semi-arid grasslands in southwestern North America has changed during the past 150 years (Van Auken 2000). Although some changes are the result of the invasion of exotic species, most changes involve dramatic increases of native woody taxa that were historically present in low densities (Johnston 1963, Archer et al. 1988, Archer 1989). The flat, deep soils of much of the South Texas Plains once supported an open savanna with a ground layer of short grasses and forbs in which Prosopis glandulosa (honey mesquite), along with lesser numbers of other shrubs and trees, which were clustered or scattered. This region also contained a mosaic of rocky, broken uplands that were dominated by relatively dense brushy vegetation. This open savanna has changed to brushy thorn-scrub woodland within the last 150 years apparently due to anthropogenic forces (Correll and Johnston 1970, Van Auken 2000, Ruthven 2001). The change to thorn-scrub woodland was primarily the result of overgrazing by domestic livestock and fire suppression (Archer et al. 1988, Ruthven et al. 2000, Ruthven 2001). Honey mesquite was the pioneer woody species involved in this transition to thorn-scrub woodland, and is currently the common dominant throughout the southwestern United States and adjacent Mexico (Ruthven 2001). Species representing two genera of thorny legumes (Senegalia and Vachellia) were also major components of these thorn-scrub woodlands. The genera Senegalia and Vachellia are segregates of the genus Acacia (sensu lato). Based on morphological and genetic evidence, it is evident that the genus Acacia s./. is polyphyletic. Also, there has been an accumulation of data, derived from molecular studies, that has lead to a better understanding of probable relationships within the genus Acacia s./., as well as the position of the genus within the Mimosoideae. These studies confirmed that the genus Acacia s.l. should be separated into as many as five genera, including Senegalia Phytologia (December 2007) 89(3) 243 (Seigler et al. 2006) and Vachellia (Seigler and Ebinger 2005). The common species of these two genera in Texas include Senegalia berlandieri [=Acacia berlandieri (guajillo, fern acacia)], S. greggii [=Acacia greggii (catclaw acacia)], S. roemeriana [=Acacia roemeriana (Roemer’s acacia)], Vachellia bravoensis [=Acacia schaffneri var. bravoensis (twisted acacia)], V. farnesiana [=Acacia farnesiana (huisache)], and V. rigidula [=Acacia rigidula (blackbrush)]. These well-armed species are common throughout the arid and semi-arid environments of the South Texas Plains (Isely 1998), and along with Prosopis glandulosa are important sources of animal fodder, fuel, and timber (Fagg and Stewart 1994). Thorn-scrub woodlands are common at the Chaparral Wildlife Management Area (CWMA), located in the northern part of the South Texas Plains ecological region. The importance and distribution of honey mesquite and other thorny legumes is determined by various biotic and abiotic factors, such as climate, moisture, edaphic conditions, present and past grazing pressures, and fire. The objective of this study was to examine the structure and composition of thorn-scrub woodland communities to understand better the importance, distribution, and habitat preferences of thorny legume species. STUDY AREA The study area was on the Chaparral Wildlife Management Area (28°20'N, 99°25'W) in the northern half of the South Texas Plains (Ruthven et al. 2000, Ruthven 2001). Located in Dimmit and La Salle counties, 12 km west of Artesia Wells, CWMA is deer-proof fenced and about 6,150 ha in size. Purchased in 1969 by the Texas Parks and Wildlife Department, it serves as a research and demonstration area. The area around CWMaA is rangeland, most holdings being large cattle ranches. Hot summers and mild winters characterize the climate of CWMaA. The average daily minimum winter (January) temperature is 5°C, the average daily maximum summer (July) temperature is 37°C, the growing season is 249 to 365 days, and the average annual precipitation (1951 to 1978) is 55 cm (Stevens and Arriaga 1985). The precipitation patterns are bimodal with peaks occurring in late spring (May and June), and early fall (September and October). Short-term periods of drought are common and rainfall can be highly variable 244 Phytologia (December 2007) 89(3) between locations (Norwine and Bingham 1985). An all-time low record of 7.16 cm fell during 1917 in Cotulla, a small town 25 km northeast of CWMA (Correll and Johnston 1970). Soils of CWMA are dominated by Duval very fine, sandy loams, gently sloping and Duval loamy fine sands, 0 to 5% slope (Gabriel et al. 1994, Stevens and Arriaga 1985). The soil surface layer is reddish brown, slightly acid, very friable, and 0 to 40 cm thick. Also present are shallow limestone ridges (calcareous rises) where soils are mildly to moderately alkaline and have a caliche layer near the surface. Topography is level to gently rolling with an average elevation of 175 m above mean sea level. Domestic livestock have grazed the CWMA since the 18th century (Lehmann 1969). Sheep production dominated from about 1750 to 1870 when cattle became the major livestock. Before 1969, grazing was continuous on the entire area. From 1969 to 1984 a four- pasture rest-rotation system was employed. Cattle were absent from the study area from 1984 to 1989. Grazing resumed in 1990, and, until 2002, CWMA utilized a high intensity, low frequency rotational grazing system. Stocking rates averaged one Animal Unit per 12 ha (Ruthven 2001). A prescribed burn program was initiated at the CWMA in 1997, but none of the sites examined in the present study has been burned (Ruthven, personal communication). Most of the CWMA was chained in 1948 (Ruthven, personal communication). Chaining involves the use of two large tractors with a very heavy linked chain connected at each end to one of the tractors. The chain is pulled across the site, disrupting and pulling out much of the vegetation (Lehmann 1984). METHODS During the summer of 2001, five thorn-scrub woodland communities were studied at the CWMA. These sites were selected based on the recommendations of CWMA site personnel who located sites where the vegetation was mature and least disturbed. All sites were upland, nearly level areas, where minimal disturbance, other than grazing, was observed. At four of the sites (1, 2, 3, and 4), a single line transect was randomly established near the center of the long axis of each community. At 30 m intervals along the length of the transect, circular plots 0.03 ha in size were located (a minimum of 10 plots) and Phytologia (December 2007) 89(3) 245 all woody plants and succulents greater than 0.4 m tall were identified and their height and average crown diameter determined to the nearest dm. As Senegalia roemeriana was found only in one small area, the entire community was examined (Site 5). This site (S50 m x 50 m in size) was divided into subplots to facilitate sampling, and all woody plants greater than 0.4 m tall were measured and identified as described above. Data from the plots were used to determine density, average cover, relative density, relative cover, and importance value (IV) for each species at each site. The IV is calculated as the sum of the relative density and relative cover. Sorensen’s Index of Similarity (ISs) was used to determine the degree of similarity between the study sites: ISs = 2C/(A+B) x 100 (Mueller-Dombois and Ellenberg 1974). At each study site, soil samples (n = 16) were taken, both under shrub and tree canopies and in open areas between shrub clusters. All samples were analyzed by the Texas Agricultural Extension Service, Soil Testing Laboratory, College Station, Texas for pH, salinity, and macro-nutrients (ppm in available form). A random sample from each site was analyzed for soil texture. Significant differences between sites for pH and the various macro-nutrients was tested with procGLM (p < 0.05) using SAS (1986). RESULTS The number of woody and succulent species recorded for the five thorn-scrub woodland sites ranged from 11 to 20 with 28 different species recorded, six being thorny species of legumes (Tables 1 thru 5). Of the 28 species encountered, seven occurred on each of the five sites: Celtis pallida (spiny hackberry), Condalia hookeri (brazil), Diospyros texana (Texas persimmon), Forestiera angustifolia (narrowleaf forestiera), Opuntia engelmannii (prickly pear), O. Jleptocaulis (tasajillo), and Prosopis glandulosa. Opuntia engelmannii, O. leptocaulis, and Prosopis glandulosa were common species in most of the communities and had high densities, covers, and IV’s. Prosopis glandulosa was the dominant species with the highest IV on two sites (2 and 5) and was second at site 3, whereas one of the two species of Opuntia ranked third or higher on all sites. The remaining species listed above were recorded for all sites; though they were sometimes common, they never dominated the community, always ranking fourth or lower in IV. For woody vegetation of all five sites, the Sorensen 246 Phytologia (December 2007) 89(3) Index of Similarity ranged from 51.6 to 84.8 (Table 6), but was usually greater than 66 indicating that sites were very similar (Mueller- Dombois and Ellenberg 1974). Though one or two species of the genus Senegalia and/or Vachellia were among the top three species at each study site, they commonly shared dominance with Prosopis glandulosa, and/or a species of Opuntia. Vachellia rigidula was found on three sites, ranking first in IV on site 4 and third on site 1 (Tables 1, 4, and 5). Vachellia bravoensis occurred on four sites, ranking third or lower in IV on all sites (Tables 1, 2, 3, and 5). Other thorny legume species were restricted to one or two sites. Senegalia roemeriana was second in IV on site 5 (Table 5), S. greggii ranked first on site 1 (Table 1), and S. berlandieri occurred on two sites, ranking second on site 4, and was uncommon on site | (Tables 1, 4). On all study sites, the soil texture was relatively uniform, being sandy loams with 61 to 75% sand, 12 to 20% silt, and 11 to 19% clay, and none were saline (Table 7). Soils of sites 1, 2, 3, and 5 were mildly to strongly acidic, mostly low to very low in available nitrates, phosphorus, and sodium, but with moderate to high amounts of available potassium, magnesium, sulfur, and calcium (Table 7). Soils of site 4, in contrast, were from a calcareous ridge (cuestas). Soil pH here was mildly to moderately alkaline with high available phosphorous and magnesium, all being significantly different (p < 0.0001) from other sites; whereas the level of available sodium was significantly lower (p < 0.0001). Although all sites had relatively high levels of available calcium, site 4, was significantly higher (P < 0.0001). DISCUSSION The thorn-scrub vegetation of CWMA and surrounding area was representative of that associated with the South Texas Plains (South Texas Brush Country or Tamaulipan Brushlands). In much of this rangeland, Prosopis glandulosa was the dominant species, with about 10 to 15 other woody or succulent, mostly thorny species, varying in abundance and composition. At the CWMA, honey mesquite was usually the dominant woody species, but, on some sites, other woody legumes were dominant or co-dominant (Johnston 1963, Correll and Johnston 1970). Phytologia (December 2007) 89(3) 247 Honey mesquite-dominated woodland with various species of Senegalia and Vachellia as co-dominants was the most common plant community on the CWMA._ This woodland community, where dominant trees were more than 3 m tall and formed a 26-60 percent canopy, would be equivalent to the Deciduous Woodland, Mesquite- Huisache Series (Prosopis glandulosa-Vachellia farnesiana) of Diamond et al. (1987) with other thorny legume species replacing huisache. At three study sites, mesquite was dominant with Vachellia bravoensis (Tables 2, 3), or Senegalia roemeriana (Table 5) as co- dominants. Vachellia bravoensis, a large shrub only rarely exceeding 3 m in height, was common at the CWMA. Except for the Vachellia rigidula/Senegalia berlandieri community on the calcareous crestas, V. bravoensis ranked third to sixth in IV on all sites. Senegalia roemeriana, in contrast, was rare at CWMA. We found this species at just one site, where, along with Prosopis glandulosa, it dominated the community in a small area less than 75 m across (Table 5). At this site, soil pH was nearly neutral (6.5), and available calcium was relatively high (1101 ppm) (Table 7). Isely (1998) did not report Senegalia roemeriana for the South Texas Plains. Correll and Johnston (1970) list the distribution of this species as farther north and west in Texas, being frequent in the Trans Pecos, and infrequent on caliche cuestas in the southern part of the Edwards Plateau. Vachellia rigidula and Senegalia berlandieri dominated limestone ridges and caliche cuestas of the CWMA. This community, in which the dominants were shrubs or small trees 0.5 to 3 m tall, and formed 26 percent of more of the total canopy, would be equivalent to the Deciduous Shrubland, Blackbrush Series (Vachellia rigidula) of Diamond et al. (1987). Vachellia rigidula appeared to be fairly site specific at the CWMA, ranking first in IV on the calcareous ridge (Site 4) and third in IV on the dry ridge that supported the Senegalia gregegii/Opuntia/Vachellia rigidula community (Site 1). This species appears to be well adapted to dry sites with high levels of available calcium. Senegalia berlandieri, in contrast, was a component of disturbed habitats at CWMA, often along roadsides, in arroyos, and other disturbed areas, but was an important stand component on the limestone ridge (Site 4). This species is common throughout southern and western Texas and is exceedingly abundant on limestone ridges and caliche cuestas (Correll and Johnston 1970, Isely 1998). At this site, 248 Phytologia (December 2007) 89(3) the alkaline soils (pH of 8.0) and extremely high concentration of available calcium appear beneficial to this species. Senegalia berlandieri and Vachellia rigidula, act as nurse trees, and facilitate the recruitment of woody and succulent species (Jurena and Van Auken 1998). Senegalia greggii had a restricted distribution at CWMA (Site 1) being common only on a dry sandy ridge. At this location catclaw acacia was the dominant member of the community, accounting for one-third of the total IV. This community, which is probably maintained by fire, grazing, and sandy soil, should be classified as the Catclaw Acacia Series, Deciduous Scrubland (Senegalia greggii). The woody vegetation at this site was short with only a few individuals being more than 2 m tall (Table 1). Canopy cover was estimated at 25 to 30 percent, and the scattered woody shrubs were mostly small and compact. No individuals of S. greggii were more than 2 m tall, and most were less than | m tall. Many were in clumps 1-2 m in diameter; the numerous, upright, bushy stems being connected by underground roots or stems. Of the thorny species of woody legume species found at the CWMA, Vachellia farnesiana (huisache) was not common. This species was rare along roadsides and near arroyos, mostly in heavily disturbed habitats. Vachellia farnesiana is common throughout the South Texas Plains, but usually in more mesic habitats (Correll and Johnston 1970), where it is a co-dominant of the Deciduous Woodland, Mesquite-Huisache Series of Diamond et al (1987). Though some of the acacia species at CWMA have distinct habitat preferences, the reasons for their continued importance, and the continued prevalence of thorn-scrub woodland communities they dominate is not entirely clear. Most information suggests that overgrazing and fire suppression were the primary causes of this encroachment (Van Auken 2000). When much of the South Texas Plains was covered with open savanna containing a dense groundcover of grasses and forbs, wildfires were frequent and of sufficient intensity to prevent encroachment by native woody species. However, overgrazing by livestock reduced the fuel load. At the same time, fire suppression allowed for a significant decrease in fire frequency creating ideal conditions for the rapid explosion of native invaders. Phytologia (December 2007) 89(3) 249 ACKNOWLEDGEMENTS We wish to acknowledge support of this work by a grant from the International Arid Lands Consortium (O1RO005), a grant from the National Science Foundation (NSF DEB 04-15803), and to thank The Texas Parks and Wildlife Department, and in particular D. C. Ruthven, J. F. Gallagher, and D. R. Synatzske of the Chaparral Wildlife Management Area for their assistance with the project. We also appreciate the critical review of the manuscript by Dr. Sean Jenkins, Ecologist, Western Illinois University, Macomb, Illinois; William McClain, Adjunct Research Associate in Botany, Illinois State Museum, Springfield, Illinois; and Dr. Chip Ruthven, Project Leader, Matador Wildlife Management Area, Paducah, Texas. LITERATURE CITED Archer, S. 1989. Have southern Texas savannas been converted to woodlands in recent history? American Naturalist 134: 545- 561. Archer, S., C. Scifres, C. R. Bassham, and R. Maggio. 1988. Autogeneic succession in a subtropical savanna: Conversion of grassland to thorn woodland. Ecological Monographs 58: 111-127. Correll, D. S. and M. C. Johnston. 1970. Manual of the Vascular Plants of Texas. Texas Research Foundation, Renner, Texas. xv + 1881 pp. Diamond, D. D., D. H. Riskind, and S. L. Orzell. 1987. A framework for plant community classification and conservation in Texas. Texas Journal of Science 39: 203-221. Fagg, C. W. and J. L. Stewart. 1994. 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New combinations in the genus Vachellia (Fabaceae: Mimosoideae) from the New World. Phytologia 87: 139-179. Seigler, D. S., J. E. Ebinger, and J. T. Miller. 2006. The genus Senegalia (Fabaceae: Mimosoideae) from the New World. Phytologia 88: 38-93. Stevens, J. W. and D. Arriaga. 1985. Soil survey of Dimmit and Zavala counties, Texas. United States Department of Agriculture, Washington, D.C. 161 pp. Van Auken, O. W. 2000. Shrub invasion of North American semiarid grasslands. Annual Review of Ecology and Systematics 31: 197-215. 251 gia (December 2007) 89(3) 7 Phytolo 0007 0001 0001 6¢°CS 6sot = Si St 609 000! 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Similarity Index of the five study sites where thorny woody legume species were common, Chaparral Wildlife Management Area, Dimmit and La Salle counties, Texas. Site 3 Site 4 Phytologia (December 2007) 89(3) Table 7. 257 Soil texture (% sand, silt, clay), salinity, pH, and macro- nutrients (ppm) of the soils on the study sites where thorny woody legume species were common, Chaparral Wildlife Management Area, Dimmit and La Salle counties, Texas. For pH and macro-nutrients, the range is given with the average value given beneath in parentheses. Different letters indicates significant difference between sites. Texture Sand Silt Clay Salinity pH Nitrate-N Phosphate Potassium Calcium Magnesium Sodium Sulfur Site | n=15 75% 12% 13% none 5.1-6.9 (6.1)a 4-6 (5.0)a 1-5 (2.7)a 151-320 (230.3)a 660-1323 (925.5)a 91-192 (134.6)ab 99-271 (198,5)a 8-26 (18.7)ac Site 2 n=14 71% 12% 17% none 4.6-6.2 (5.3)b 5-23 (8.6)b 1-10 (5.6)a 226-462 (323.9)b 469-1497 (721.9)a 84-225 (144.5)a 98-255 (214.9)a 6-29 22.2)abc Site 3 n=13 71% 16% 13% none 4.5-6.6 (5.2)b 5-9 (6.2)ab 3-8 (5.3)a 205-364 (266.5)a 495-1428 (784.8)a 76-145 (107.2)b 106-258 (189.5)a 8-30 18.8)ac Site 4 n=18 61% 20% 19% none 7.7-8.3 (8.0)c 4-9 (5.6)a 5-24 (16.8)b 198-361 (287.7)ab 3515-23291 (13886.8)b 146-413 (242.4)c 32-235 (82.9)b 14-47 26.8)be Site 5 n=2 77% 8% 15% none 6.3-6.6 (6.5)a 5 (5.0)a 2-3 (2.5)a 301-358 (329.5)ab 981-1221 (1101.0)a 141-178 (159.5)a 217-250 (233.5)a 22-24 23.0)c 258 Phytologia (December 2007) 89(3) MELAMPODIUM MOCTEZUMUM (ASTERACEAE: HELIANTHEAE), A NEW SPECIES FROM SONORA, MEXICO B. L. Turner Plant Resources Center The University of Texas at Austin Austin, Texas 78712 billie@uts.cc.utexas.edu ABSTRACT A new species, Melampodium moctezumum, is described from Mpio. de Moctezuma, Sonora, Mexico and Cochise County, southeastern Arizona, U.S.A. It is closely related to the more southwestern M. cupulatum but can be distinguished from that taxon by several features, most notably by its broader, coarser, more venose, markedly lobed leaves. KEY WORDS: Asteraceae, Heliantheae, Melampodium, Mexico, Sonora, Arizona. Routine identifications from Sonora, Mexico and _ the southwestern U.S.A. have revealed the following novelty: Melampodium moctezumum B. L. Turner, sp. nov. Melampodio cupulatum A. Gray similis sed foliis laminis venosis valde lobatis (vs. integris) in petiolos tenues gradatim descrescentibus (vs. subsessilibus vel brevipetiolatis) et flosculis radii plerumque 2-6 mm longis (vs 5-10 mm). Annual herbs 15-45 cm high. Stems reddish, erect, 1-4 mm diameter, sparsely pubescent to glabrous. Larger mid-stem leaves 4-6 cm long, 1.0-1.8 cm wide; petioles 0.3-2.0 cm long, the blades markedly venose with lobed or irregularly serrate margins, appressed-pilose and punctate-glandular on both surfaces. Peduncles 2-7 cm long. Heads 6-9 mm high, 8-20 mm wide (with rays expanded). Outer involucral bracts 5, broadly ovate, ca 5 mm high, connate for 2/3-4/5 their length, Phytologia (December 2007) 89(3) 259 their margins scarious. Receptacle conical, ca 6 mm high, 2 mm wide. Pales oblanceolate, 3-4 mm long with flabellate, fimbriate, yellow- orange apices. Ray florets12-15; ligules “yellow-orange,” 2(3)-6(7) mm long, 1-2 mm wide. Disk florets numerous; corollas yellow- orange. Achenes 3-4 mm long, somewhat falcate laterally, markedly tuberculate, epappose, hoodless. TYPE: MEXICO. SONORA: Mpio. de Moctezuma, 13.7 km NNW of Tepache on road to Moctezuma (Son 117), sparse scrub on basalt cobble plain, 730 m, 17 Aug 2003, 4. L. Reina G. 2003-943 (with T. R. Van Devender and Z. Liu). (Holotype: TEX; isotype: ARIZ). ADDITIONAL COLLECTIONS EXAMINED (TEX): MEXICO. SONORA: Mpio. de Moctezuma, 18.1 km SSE of Moctezuma, “locally common annual,” 14 Aug 2006, Reina 2006-486; 21.4 km SSE of Moctezuma, 697 m, 19 Oct 2003, Van Devender 2003-1228 (with A. L. Reina); 19.6 km SSE of Moctezuma, on road to Tepache, 19 Oct 2003, Van Devender 2003-1230; 18.9 km SSE of Junction with Moctezuma- Huasabas Hwy on road to Tepache, 635 m, 14 Sep 2006, Van Devender 2006-802. UNITED STATES: ARIZONA. Cochise Co.: W side of Peloncillo Mts., across road from Cottonwood Creek Cemetery, 12 Sep 1987, Kluever s. n. (ARIZ); W side of Peloncillo Mts., across road from Cottonwood Cemetery, 7.5 mi E of Guadalupe Canyon turnoff, 4550 ft, 9 Aug 1990. Warren 90-16 [with Kluever] (ARIZ, ASU, TEX)). Van Devender (pers. comm.) has provided the following comments regarding its habitat at the type locality and surroundings: The new species is found in the basalt lava plains along Sonora Highway 117 between Moctezuma and Tepache in the Municipio de Moctezuma. Lava plain starts at 17.3 km SSE of Moctezuma and extends 12.6 km before dropping into the Rio Tepache about 7 km N of Tepache. The lava plains are a very unusual habitat with medium to small black basalt rocks in a matrix of dark brown, clay rich soil. The vegetation is foothills thornscrub, although the plants are often widely spaced and of smaller stature compared to the same vegetation on rocky slopes. Dominants include various 260 Phytologia (December 2007) 89(3) legumes (Acacia cochliacantha, A farnesiana, A. occidentalis, Haematoxylon brasiletto, Parkinsonia praecox, and Prosopis velutina), tree ocootillo (Fouquieria macdougalii), guayacan (Guaiacum coulteri), organpipe cactus (Stenocereus thurberi), and a prickly pear (Opuntia sp.). In the summer rainy season, the combination of rocky substrate and rich soil yields a profusion of annual and perennial herbs. The basalt lava flowed south and west from Cerro Blanco about 500,000 years ago into the late Pleistocene (Mead et al. 2006), providing a maximum time for the isolation of the new species. Melampodium moctezumum is seemingly closely related to both M. appendiculatum and M. cupulatum but differs in having leaves petiolate with markedly venose blades, their margins lobate to irregularly serrate (vs sessile, weakly venose and margins entire). The characters called to the fore and its relative geographical isolation suggest specific status for the taxon; at least no intermediates between the several taxa were detected among the numerous specimens of M. appendiculatum and/or M. cupulatum on file at LL, TEX. Distribution of these taxa is shown in Figs. 2 and 3. Collections of M. moctezumum from the U.S.A. have somewhat larger heads with longer rays but otherwise appear very similar to the Mexican collections. According to its collectors, the florets of MZ moctezumum are “orange-yellow” and the plants are said to be locally “abundant.” The species is named for the Municipio of Moctezuma, where first collected. ACKNOWLEDGEMENTS I am grateful to ARIZ and ASU for the loan of specimens; to my colleague, Guy Nesom, for the Latin diagnosis and reviewing the paper; and to my colleague Tom Van Devender for his insistence that the taxon had some sort of biological reality. Phytologia (December 2007) 89(3) 261 LITERATURE CITED Mead, J.I. et al. 2006. Tropical marsh and savanna of the late Pleistocene in northeastern Sonora. Southwestern Naturalist 51: 226-239. Taree | Fig. 1. Holotype of Melampodium moctezumum. 262 Phytologia (December 2007) 89(3) MELAMPODIUM cupulatum MELAMPODIUM ® appendiculatum © moctezumum Fig. 3. Distributions of M. appendiculatum (closed circles) and M. moctezumum (open circles). Phytologia (December 2007) 89(3) 263 JUNIPERUS MARITIMA, THE SEASIDE JUNIPER, A NEW SPECIES FROM PUGET SOUND, NORTH AMERICA Robert P. Adams Biology Department, Baylor University, Box 727, Gruver, TX 79040, USA Robert_Adams@baylor.edu ABSTRACT Based on analyses of terpenoids, nrDNA and trnC-D SNPs as well as morphology and ecology, a new cryptic species, Juniperus maritima, from the Puget Sound region is recognized. The species, previously included in J. scopulorum, is characterized by having seed cones that mature in one year (14-16 months), seeds usually exserted from the cone, obtuse scale leaf tips, usually reniform seed cones, scale leaves overlap less than 1/5 the length, and branchlets smooth and reddish-brown. Called the seaside juniper, it grows on rocky areas (rarely sand dunes) near the sea, in Puget Sound. KEY WORDS: Juniperus maritima, Puget Sound, J. scopulorum, J. virginiana, cryptic species, terpenoids, nrDNA, trnC-trnD, SNPs. The smooth leaf margined (40X) junipers in the western hemisphere are very widespread and are composed of the Caribbean Juniperus: J. barbadensis L., J. bermudiana L., J. gracilior Pilg. , J. g. var. ekmanii (Florin) R. P. Adams, J. g. var. urbaniana (Pilg. & Ekman) R. P. Adams, J. /ucayana Britt., and J. saxicola Britt. & P. Wilson; the Mexican junipers: J. blancoi Mart. var. blancoi, J. b. var. huehuentensis R. P. Adams, S. Gonzales & M. G. Elizondo, and J. mucronata R. P. Adams and the Canada/ United States junipers: J. horizontalis Moench, J. scopulorum Sarg., J. virginiana L. and J. v. var. silicicola (Small) E. Murray (Adams, 2004). Juniperus scopulorum and J. virginiana are weedy junipers that occupy millions of acres in the United States and Canada. Adams (1983) analyzed the leaf terpenoids of populations of J. scopulorum 264 Phytologia (December 2007) 89(3) ' Differiential Systematics | . 6 Canonical Axes | Terpenoids 1 1 ee Figure 1. Contoured differentiation based on the first 6 canonical axes using leaf terpenoid data (from Adams, 1983). Areas with close contour lines are areas of high differentiation. Phytologia (December 2007) 89(3) 265 from throughout its range and found that much of the variation within putative J. scopulorum was due to differentiation in populations from Puget Sound from the balance of the range of J. scopulorum (Fig. 1). The differentiation of the two populations sampled in the Puget Sound (VB, Vancouver Isl., B.C.; PW, Whidbey Isl., WA) accounted for 50.2% of the variance among all 17 populations (Adams, 1983). It was hypothesized that the Puget Sound populations have been genetically isolated from the main, Rocky Mountain populations since the Pleistocene (or earlier) (Fig. 2). Notice (Fig. 2, A) that the Puget Sound Figure 2. A. Maximal Wisconsin ice cover showing the extinction of local populations of J. scopulorum. BB. Proposed refugia and recolonization following the Wisconsin (adapted from Adams, 1983). populations were thought to have retreated to a refugium south of the their present distribution and that no common refugia are indicated for the Puget Sound populations and J. scopulorum from the Rocky Mountains (Fig. 2 B). 266 Phytologia (December 2007) 89(3) Recently, Schwarzbach et al. (2008), using combined ITS and trmC-D sequence data in their study of the phylogeny of Juniperus, found that an individual from Puget Sound came out in the clade with J/. virginiana, not in the clade with J. scopulorum. This prompted the author to reexamine the terpenoid data (Adams, 1983). Figure 3 shows a PCO of the terpenoids. Four distinct entities are resolved: J. horizontalis, J. scopulorum, J. virginiana, and the Puget Sound populations. It should be noted that each stick represents the mean of 15 individuals (a total of 441 individuals analyzed for over 100 terpenoids, with the 30 terpenoids with the highest F ratios utilized for PCO). These data are robust and must be given significant weight in assigning the taxonomic position of the Puget Sound populations. 2 (15%) PCO Terpenoids J. horizontalis tne f BE ™, \ J. scopulorum é 3(8%) 1 (22%) Figure 3. Principal coordinate ordination (PCO) utilizing terpenoid data from Adams (1983). Each of the sticks represents population mean of 15 individuals, except for the 2 Puget Sound populations that contained 8 and 13 samples. Phytologia (December 2007) 89(3) 267 Because the ITS and trnC-D sequence data (Schwarzbach et al. 2008) fails to support a conspecific status of the Puget Sound population and J. scopulorum, it seemed prudent to make additional collections and analyze additional samples using several DNA methods. The purpose of this paper is to compare ITS and trnC-D SNPs (single nucleotide polymorphisms) analyses of junipers from Puget Sound with J. scopulorum and J. virginiana with previous terpenoid, morphological and ecological data to determine the taxonomic status of the Puget Sound (seaside) juniper. MATERIALS AND METHODS Specimens used in this study are shown in table 1. Voucher specimens are deposited at BAYLU herbarium Baylor University. One gram (fresh weight) of the foliage was placed in 20 g of activated silica gel and transported to the lab, thence stored at -20° C until the DNA was extracted. DNA was extracted using the Qiagen DNeasy mini kit (Qiagen Inc., Valencia CA). SNPs obtained from DNA sequencing ITS and trnC-trnD amplifications were performed in 50 ul reactions using 10 ng of genomic DNA, 3 units Qiagen Taq polymerase, 5 ul 10x buffer (final concentration: 50 mM KCl, 10 mM Tris-HCl (pH 9), 0.01% gelatin and 0.1% Triton X-100), 1.75 mM MgCl,, 20 ul Q solution (2X final), 400 uM each dNTP, 1.8 uM each primer and 4%(by vol.) DMSO. Primers (5'-3'): ITS: ITSA = GGA AGG AGA AGT CGT AAC AAG G; ITSB =CTIT TIC CTC CGC TIA TIGATA TG. ITSA and ITSB primers from Blattner (1999). trnC-trnD: CDFor: CCA GTT CAA ATC TGG GTG TC CDRev: GGG ATT GTA GTT CAA TTG GT CDFor, CDRev primers from Demesure et al. (1995). CDI0F: AAA GAG AGG GAT TCG TAT GGA CD3R: AAC GAA GCG AAA ATC AAT CA CD10F and CD3R primers from Andrea Schwarzbach (per. comm.) 268 Phytologia (December 2007) 89(3) The following PCR conditions were used: MJ Research Programmable Thermal Cycler, 45 cycles, 94°C (1 min.), 50°C (1 min.), 72°C (1 min.), with a final step of 72°C (5 min.). The PCR reaction was subjected to purification by agarose gel electrophoresis (1.5% agarose, 45 min.). The nrDNA primers (ITSA, ITSB) produced a band of approx. 1120 bp. The internal trnC-trnD primers, CD10F-CD3R produced a band of approx. 850 bp. In each case the band was excised and purified by use of a Qiagen QIAquick gel extraction kit. The gel purified DNA band with the appropriate primer was sent to McLab Inc. for sequencing. Sequences for both strands were edited and a consensus sequence was produced using Chromas, version 2.31 (Technelysium Pty Ltd.). Alignments were done using Clustal W and then manually corrected. Indels were coded with a "-" for the first nucleotide and "I" for succeeding nucleotides such that an indel was treated as a single mutation event. Sequences were deposited in GenBank (table 1). SNPs analyses Aligned data sets (nrDNA and tmC-tmD) were analyzed by CLEANDNA (Fortran, R. P. Adams) to remove invariant data and nucleotides that only varied by a single polymorphism among individuals. Mutational differences were computed by comparing all SNPs, divided by the number of comparisons over all taxa (= Gower metric, Gower, 1971; Adams, 1975). Principal coordinate analysis was performed by factoring the associational matrix using the formulation of Gower (1966) and Veldman (1967). A minimum spanning network was constructed by selecting the nearest neighbor for each taxon from the pair-wise similarity matrix, then connecting those nearest neighbors as nodes in the network (Adams, et al. 2003). Phytologia (December 2007) 89(3) 269 Table 1. Specimens collected, locations and GenBank accession numbers. All specimens deposited at BAYLU. Taxon/collection# Location GenBank acc. J. scopulorum, Adams 10895 Kamas, UT ITS: EF608963-65 -10897 tmCD EF608988-90 J. virginiana Adams 6753-6755 Hewitt, TX ITS: EF608980-82 trnCD: EF609002-04 Adams 10230 Knoxville, TN ITS: EF608973-75 -10232 trnCD: EF608996-98 J. v. var. silicicola Adams 9186-88 Ft. DeSoto Park, ITS: EF608977-79 Mullet Key, FL trnCD: EF609009-11 J. maritima Adams 11056-58 Brentwood Bay (BB) tmCD: EF608985-87 Vancouver Isl., BC Adams 11061-63 Cowichan Bay(CB) ITS: EF608968-70 Vancouver Isl., BC tmCD: EF608992, EF609007, EF608993 Adams 11064 Yellow Point (YP) ITS: EF608984 Vancouver Isl., BC tmmCD: EF608991 Adams 11065-66 — Lesqueti Isl. (LS) ITS: EF608967 BC tmCD: EF609000-01 Adams 11067-68 Friday Harbor (FH) ITS: EF608971 San Juan Isl., WA trnCD: EF608994-95 Adams 11075 Whidbey Isl. (WI) ITS: EF608983 Cranberry L., WA trnCD: EF609005 Adams 11076 Fidalgo Isl. (FI) ITS: EF608972 State Park, WA tmCD: EF609006 Adams 11077-78 Skagit Isl. (SK), WA ITS: EF608966, EF608976 trnCD: EF609008, EF608999 270 Phytologia (December 2007) 89(3) RESULTS AND DISCUSSION Analysis of the nrDNA (ITS) sequences revealed little variation among these essentially sibling species. One exception was individual 11076 from Fidalgo Island, WA that had a 67 bp deletion at position 399. The tree appeared to be morphologically similar to other trees in the area and it is assumed that this indel represents a single mutational event. A few single nucleotide mutations were found among individuals and removed from the data. This resulted in 18 SNPs among J. scopulorum, J. virginiana, J. v. var. silicicola and the Puget Sound (seaside) junipers. Factoring the associational matrix resulting in eigenroots that accounted for 55.4%, 24.8%, 6.1%, and 4.2% before they began to asymptote. Notice that two degrees of freedom (axes 1,2) accounted for 80.2% of the variance! This implies that there are only 3 groups (n-1 = 2). Ordination of the individuals (Fig. 4) revealed three groups: J. scopulorum, J. virginiana (including var. silicicola) and the Puget Sound junipers. The minimum spanning network shows (Fig. 4) that the Puget Sound junipers are nearly equidistant between J. scopulorum (5 bp) and J. virginiana (4 bp). The ITS SNPs, although not plentiful, are fully congruent with the terpenoid and morphological data. Analysis of the trnC-trnD cpDNA sequences proved to be difficult. Numerous indels and single mutational events were present. Figure 5 shows the variation encountered in the sequence length (1580 bp). This includes both nucleotide substitutions and single indels. NCBI blast of the region from CDIOF to CD3R did not yield information on the nature of the conserved regions where these primers reside. Each of the J. v. var. silicicola samples (3 indvs.) had a 254 bp deletion in the CD10F - CD3R region not found in any other samples. Juniperus v. var. silicicola is a coastal juniper from the sand foredunes of se United States. Analyses including J/. v. var. silicicola samples in the data set showed it to be quite differentiated in its trnC-trnD sequence, so these were removed from further consideration for the trnC-trnD data. Phytologia (December 2007) 89(3) 271 2 (25%) PCO 18 ITS SNPs J. scopulorum i J. virginiana Puget Sound 3 (6%) Figure 4. PCO ordination of based on 18 SNPs of ITS sequence data. Juniperus scopulorum (3 indvs.) each had a 4 bp (TATA) insert at position 986, not shared with either J. virginiana or the Puget Sound junipers. Juniperus virginiana (6 indvs.) had an insert of 4 bp (TTTT) at position 262 not found in any other samples. Four trees in the study had a 4 bp indel at position 712. These trees were from Friday Harbor (TATT, TATT) , Fidalgo Island (TAAT) and Whidbey Island (TAAT). The population from Fidalgo Island is only about 10 km north of the Whidbey Island population. However, the Skagit Island population, only 5 km east of the Whidbey Island population, did not have the indel. 272 Phytologia (December 2007) 89(3) Number of variable sites vs. position in trnCD based on J. scop(3), J. virg(6), and Puget Sound(19) tree samples : 720 960 1200 1580 +“ +— CD 3R CD Rev Figure 5. Frequency distribution of variable sites in the trnC-trnD region. Principal Coordinates analysis of the association measures using 78 polymorphic SNPs from the tmC-trnD sequences produced three eigenroots before the eigenroots began to asymptote. These three eigenroots accounted for 25.8%, 14.6% and 11.6% of the variation among individuals. Three eigenroots implies that 4 groups are present in the data. However, ordination (Fig. 6) shows two principal groups: J. scopulorum and J. virginiana / Puget Sound individuals. These two groups (axis 1) accounted for 26% of the variation among the individuals. There is a partial separation of the J. virginiana individuals (V, fig. 6), but it is incomplete. Considerable variation exists among the Puget Sound individuals, but a detailed examination failed to correlate their ordination with geography. The trnC-trnD data seem similar to the trnL-trnF cp data from J. occidentalis Hook. var. australis (Vasek) A. & N. Holmgr. Phytologia (December 2007) 89(3) 213 3 (12%) PCO trnC-trnD 78 SNPs V = J. Virginiana @ = Puget Sound J. scopulorum 1 (26%) Figure 6. PCO ordination of J. scopulorum, J. virginiana and Puget Sound individuals based on 78 SNPs. (now J. grandis R. P. Adams) and J. osteosperma (Torr.) Little from Terry et al. (2000). The latter workers found that a cp haplotype, a mutation at position 436 (at the 3' position of the 7ru 9/ restriction site), was invariant within J. o. var. australis (J. grandis), but varied clinally (with some notable exceptions) from the area of sympatry (w. Nevada) to Utah. However, several populations in UT, CO and WY, the farthest removed from J. 0. var. australis, had high frequencies of the cp haplotypes. They considered three explanations: inheritance of ancestral polymorphism, intraspecific polymorphism, and hybridization between J. occidentalis var. australis and J. osteosperma. Of course, Vasek (1966) has already made a strong case for hybridization between these taxa based on morphological data. Terry et al. (2000) opted for the hybridization (and introgression) as the explanation with gene flow (via pollen) from J. 0. var. australis to typical J. osteosperma. This 274 Phytologia (December 2007) 89(3) would be in agreement with the transfer of cpDNA via pollen from J. o. var. australis, but not the reverse flow. However, one can not rule out the persistence of ancestral cpDNA as another explanation. In any case, analysis of the trnL-trnF sequences gave a picture of incomplete separation between these morphologically well defined Juniperus species. This appears to be the case for trnC-trnD cp data for J. virginiana and the Puget Sound junipers. The trnC-trnD PCO (Fig. 6) stands in contrast to the terpenoid data (Fig. 3) and ITS data (Fig. 4). A striking aspect of the Puget Sound, seaside junipers is their habitat. They all grow at the seaside (or lakeside) on granite or sand (Fig. 7). This is a very different kind of habitat than that found in J/. scopulorum and J. virginiana. Juniperus scopulorum grows on dry, rocky mountainous soils. Juniperus virginiana is more cosmopolitan, growing in limestone areas as well as deep soils. Both J. scopulorum and J. virginiana are weedy junipers that invade old fields and disturbed roadsides. In contrast, the seaside juniper is not weedy and usually appears as if it is relictual (i.e., older trees, with few or no seedlings). The Puget Sound juniper's habitat seems to be very restricted and has only been collected in a few locations (Fig. 7). The Puget Sound climate is very different than the Rocky Mountain or the eastern US climates, having a mild, wet regime. In short, the Puget Sound juniper has evolved physiological genes to facilitate its growth in such an environment. Is the Puget Sound, juniper a distinct species? Ownbey (1950) has provided us with a very practical species definition. He emphasizes that species are natural groups, characterized by: 1. a combination of distinctive morphological features (and/or chemical/ DNA features, my addition); 2. The taxa are reproducing under natural conditions; and 3. There is not free gene exchange between the taxa concerned. How can we apply the 'Ownbey species concept' to the present taxonomic problem? 1. The taxa are natural groups, characterized by a combination of distinctive morphological features (and/or chemical/ DNA features, my addition). Phytologia (December 2007) 89(3) 275 Recently, Issakainen (1999) wrote "We easily forget that different parts of a single organism's genome may have a different evolutionary history.". We might modify his statement to read "different parts of the genome may be under differential selection pressure." We, as taxonomists, have relied on morphology as the deciding data for the recognition of species, varieties, and indeed most of our nomenclatural taxa. This is only natural, as the morphology is "what you see." The morphology is a product of the plant's genes plus the environment. The genes are composed of DNA and in tomato the genome size is about 700,000,000 base pairs (bp) versus 4,000,000 bp in E. coli and 230,000,000,000 bp in man (Brown, 1986) and these appear to represent 20,000 to 30,000 genes (Somerville and Somerville, 1999). The amount of the genome that we see in the morphology is not known precisely. But, in an interesting study of two species of goldenrod (Solidago), Charles and Goodwin (1953) made the following estimates for the minimum number of genes for several key taxonomic characters: Character Minimum number of genes leaf margins: entire vs. serrate leaf surface: glabrous to pubescent leaf thickness basal leaves: length leaf cuticle: degree of sculpturing stomatal apparatus: length WwmMaonnn Thus, for these 6 key characters separating S. sempervirens and S. rugosa, they estimated that the species differed by a minimum of 35 genes. How many DNA base pairs this represents is unknown. Irving and Adams (1973) applied these methods to estimate the minimum number of genes controlling monoterpenes in Hedeoma. They found that 20 monterpenoids were inherited by from | to 7 genes, with an average of 1.95 genes per compound. Thus, these 20 monoterpenoids appeared to be inherited by a minimum of 39 genes. Again a small sample of the total genome. 276 Phytologia (December 2007) 89(3) If Solidago and Hedeoma have 20,000 to 30,000 genes as commonly expected in plants (Somerville and Somerville, 1999), then the Solidago morphology and Hedeoma monoterpenes are small samples of these genomes. Somerville and Somerville (1999) show that, in Arabidopsis, 54% of the genes can be assigned a known function. Although they did not show morphology per se, they did show that of the genes with known function, approximately 5% control cell structure and 6% code for secondary metabolism in Arabidopsis. For the case of the seaside (Puget Sound) juniper, the taxon is distinct from both J. scopulorum and J. virginiana in its terpenoids and ITS sequences. It is also differentiated in its physiology, enabling it to grow in a habitat foreign to both J. scopulorum and J. virginiana. Clearly the Puget Sound juniper (seaside juniper) is characterized by a combination of terpenoid, ITS DNA and physiological traits, these independent of those relating to morphology. 2. The taxa are reproducing themselves under natural conditions. Of immediate concern upon examining the Puget Sound juniper, was that it might be an escaped cultivar of J. virginiana. Juniperus virginiana was (and continues to be) commonly cultivated by settlers moving westward in the United States. It is a very common ornamental tree found at homesteads, cemeteries and parks in the central and western United States. Several groups of early immigrants came to the Pacific Northwest. Likely, the earliest were the Spanish and Portuguese sailors and explorers. It is extremely unlikely that these explorers, who apparently did not build permanent settlements in the Pacific Northwest would have brought J. virginiana for cultivation. The most likely group of settlers were the Anglos from the eastern United States who used the Oregon Trail to migrate to the Pacific Northwest between 1841 and 1869. Apparently, Hudson Bay trappers and Russians visited Puget Sound as early as 1830 (Steve Erickson, pers. comm.). So any junipers older than 176 years old (in 2006) would have pre-dated the earliest known Anglo settlers. Although juniper growth rings are not reliable in desert regions due to lack of rings in dry years, the precipitation of Puget Sound is very consistent with a wet season each year. Therefore, the Phytologia (December 2007) 89(3) 277 growth rings should be a very good measure of the age of junipers in the area. In 2006, the author cored several very large junipers in Puget Sound. Table 2 shows the growth rings varied from 86 to 210 rings. A linear exploitation gives values over 400 yr. Most of the cores had uniform ring spacing for the region scored, except for 11070, Lesqueti Table 2. Estimated ages and sizes of junipers in the Puget Sound area. Tree and trunk # rings % radius approx. Location radius counted __ counted __age 11065, Yellow 22:3em 128 100% 128 yr. Point, BC in 22.8 cm 11061, Cowichan 35.5 cm 167 58.6% > 167yr. Bay, BC in 20.8 cm ca. 285 yr. 11065, Lesqueti 35 cm 163 B29% ~— > 16370. Island, BC in 29 cm ca. 196 yr. 11070, Lesqueti 64 cm 210 2 20 yr Island, BC in llem ca. 400- 500 11067, Friday 40 cm 86 60% > 86 yr. Harbor, San Juan Isl. in 24 cm ca. 140 yr. 11072, English 106.7 92 I3.0e «= 92 yr. Camp, San Juan Isl. in 18 cm ca. 273 yr. 11077, Skagit 118.6cm 140 33.7% >140 yr. Island, WA in 20 cm ca. 415 yr. 278 Phytologia (December 2007) 89(3) Island, that had very compressed rings in the 11 cm that was scoreable. It is clear that the seaside juniper predates Anglo settlement and the taxon is naturally occurring. In addition, high genetic variation between the seaside junipers, argues against the introduction by settlers. Recent introduction would have produced a genetic bottleneck effect that is not present in these populations. Although there is almost universal damage to the seed cones by insects, resulting in exserted seeds, the seaside juniper is reproducing itself under natural conditions. 3. There is not free gene exchange between the taxa. The nearest population of J. scopulorum is about 140 km east of Puget Sound at Ross Lake, BC. The nearest population of J. virginiana is in central Nebraska, several thousand km to the east. It seems unlikely that gene flow is currently occurring between the seaside juniper and either J. scopulorum or J. virginiana. In summary, the seaside juniper of Puget Sound is an entity that is genetically defined (primarily by its chemistry and DNA sequences), reproducing itself under natural conditions and is not interbreeding with other juniper species. Because of this, I recognize it as a new species as follows: Juniperus maritima R. P. Adams sp. nov. Type: Canada, BC, Vancouver Island, Brentwood Bay, Lat 48° 34.794' N; Long 123° 20.211' W, elev. 5 m., 29 May 2006, R. P. Adams 11056 (HOLOTYPE: BAYLUWSOTYPE: YY). A J. scopulorum similis sed differt strobilis seminiferis in 14-16 menses maturescentibus, seminibus plerumque ex strobilo exsertis, et apicibus foliorum squamiformium obtusis. Differt a J. virginiana strobilis seminiferis majoribus (6-8 mm) saepe reniformibus, seminibus plerumque ex strobilo exsertis, foliis squamiformibus minus quam 1/5 longitudinis imbricatis, et ramulis laevibus porphyreis. This species is similar to J. scopulorum but differs in that the seed cones mature in | year (14-16 months), seeds are usually exserted from the cone, and the scale leaf tips are obtuse (Table 3). It differs Phytologia (December 2007) 89(3) 279 from J. virginiana in having larger seed cones (6-8 mm) that are often reniform, seeds usually exserted from the cone, scale leaves overlap less than 1/5 the length, and branchlets are smooth and reddish-brown. Table 3. Morphological comparison of J. maritima, J. scopulorum and J. virginiana. J. maritima J. scopulorum _J. virginiana seed cones mature 1 yr (14-16 mos.) 2 years 1 year seed cone diam. 6-8 mm 6-9 mm 3-6(7) mm seed cone shape globose to globose to ovoid reniform reniform seeds per cone (1) 2 (1) 2 (3) 1-2 (3) exserted seeds ubiquitous rare rare scale leaf overlap < 1/5 length <1I/Slength > 1/4 length scale leaf tips obtuse acute to obtuse acute branchlets (6-15mm, smooth, smooth, brown with diam.) reddish-brown bright reddish- persistent brown old leaves Junipers maritima is known only from the Puget Sound area (Fig. 7). It is usually found in rocky areas, often within meters of the water. However, a population exists on coastal sand dunes near Cranberry Lake, Whidbey Island, WA. No other population has been found on sand, so that site 1s likely atypical. Population Status The Lesqueti Island population (LS, Fig. 7) is in a nature reserve and consists of hundreds of trees. It appears to be a robust population and not threatened. The Yellow Point population (YP, Fig. 7) at Yellow Point Resort, private land, has tens of trees that appear to be reproducing, but development and human impact at the resort threatens it. 280 Phytologia (December 2007) 89(3) Lesqueti Island o) BC Vancouver Island wi Skagit Island Juan Whid WA Island Island Figure 7. Distribution of Juniperus maritima based on Adams field collections (acronyms) and herbarium specimens (stars) from V, WS, and WTU. Phytologia (December 2007) 89(3) 281 The Cowichan Bay population (CB, Fig. 7) is on private land. Approximately 10 trees were seen. No seedlings or saplings were observed. The Brentwood Bay population (BB, Fig. 7) consists of 6 mature trees on seaside granite. It is at the north end of the Tsartlit Reserve and is protected from development. The Friday Harbor plants are found chiefly on rocks at the Univ. of Washington Marine Station (8-10 trees) and at the NPS, English Camp (6 old, mature trees) on the opposite side of San Juan Island. These sites are protected from development. The Fidalgo Island, Washington State Park, Anacortes, WA was the most robust population examined with hundreds of trees of various ages. It is in a protected park and its future looks secure. On Whidbey Island, a natural population was found on coastal sand dunes in Deception Pass Park (near Cranberry Lake). There are 10-20 trees, all very stunted from constant ocean winds and salt spray. Some age differences were observed. The site is in a park and protected from cutting. However, beach use and a large storm could threaten this population. Several other seaside junipers appear to have been planted at houses in the interior of Whidbey Island and are growing well in deep soil. About 10 individuals were seen on Skagit Island, ranging from very old to young saplings. Skagit Island is a protected area so, aside from fires, this little population appears stable. ACKNOWLEDGEMENTS Thanks to B. L. Turner and A. E. Schwarzbach for reviewing the manuscript and Guy Nesom for providing the Latin diagnosis. Field collections would not have been possible without excellent assistance by numerous collaborators: Steve Erickson, Whidbey and Skagit Island, WA; Eugene Kozloff, San Juan Island; Peter Dederich, NPS, English Camp, San Juan Island, WA; Ted Smith, Deception Pass State 282 Phytologia (December 2007) 89(3) Park, Whidbey Island, WA; Richard Hebda, Adolf Ceska, and John Pinder-Moss, Royal British Columbia Museum (V); Lance Goldy, Yellow Point Resort, BC; and Drew Chapman and Wade Calder, Lesqueti Island Ecological Reserve, BC. Thanks to Andrea Schwarzbach for trnC-trnD primers CDIOF and CD3R_ sequence information and helpful advice. Thanks to Tonya Yanke for lab assistance. This research supported in part by funds from Baylor University. LITERATURE CITED Adams, R. P. 1975. Statistical character weighting and similarity stability. Brittonia 27: 305-316. Adams, R. P. 1983. Infraspecific terpenoid variation in Juniperus scopulorum: Evidence for Pleistocene refugia and recolonization in western North America. Taxon 32: 30 - 46. Adams, R. P. 2004. Junipers of the World: The genus Juniperus. Trafford Publ., Vancouver, B. C. Adams, R. P., A. E. Schwarzbach, and R. N. Pandey. 2003. The Concordance of Terpenoid, ISSR and RAPD markers, and ITS sequence data sets among genotypes: An example from Juniperus. Biochem. Syst. Ecol. 31: 375-387. Blattner, F. R. 1999. Direct amplification of the entire ITS region from poorly preserved plant material using recombinant PCR. BioTechniques 27: 1180-1186. Charles, D. R. and R. H. Goodwin. 1953. An estimate of the minimum number of genes differentiating two species of golden-rod with respect to their morphological characters. Amer. Natl. 77: 53-69. Demesure, B., N. Sodzi and R. J. Petit. 1995. A set of universal primers for amplification of polymorphic non-coding regions of mito- chondrial and chloroplast DNA in plants. Mol. Ecol. 4:129-131. Phytologia (December 2007) 89(3) 283 Gower, J. C. 1966. Some distance properties of latent root and vector methods used in multivariate analysis. Biometrika 53: 326-338. Gower, J. C. 1971. A general coefficient of similarity and some of its properties. Biometrics 27: 857-874. Irving, R. S. and R. P. Adams. 1973. Genetic and biosynthetic relationships of monoterpenes. pp. 187-214. In Terpenoids: Structure, biogenesis, and distribution. Vol. 6. Recent Advances in Phytochemisty Genetics Series. V. C. Runeckles and T. J. Mabry, eds., Academic Press, NY. Issakainen, J. 1999. Dear Mr. Code ... Taxon 47: 341- 348. Ownbey, M. (1950) Natural hybridization and amphiploidy in the genus Tragopogon. Amer. J. Bot. 37: 487-499. Schwarzbach, A. E., R. P. Adams and J. A. Morris. 2008. Phylogeny of Juniperus based on nrDNA and trnC-trnD sequences. (in prep). Terry, R. G., R. S. Novak and R. J. Tausch. 2000. Genetic variability in chloroplast and nuclear ribosomal DNA in Utah juniper (Juniperus osteosperma, Cupressaceae): Evidence for interspecific gene flow. Amer. J. Bot. 87: 250-258. Somerville, C. and S. Somerville. 1999. Plant Functional Genomics. Science 285: 380 - 383. Vasek, F. C. 1966. The distribution and taxonomy of three western junipers. Brittonia 18: 350-372. Veldman D. J., 1967. Fortran programming for the behavioral sciences. Holt, Rinehart and Winston Publ., NY. 284 Phytologia (December 2007) 89(3) ERYNGIUM HETEROPHYLLUM (APIACEAE) AND ERIOCAULON LINEARE (ERIOCAULACEAE) DELETED FROM THE LOUISIANA FLORA M. H. MacRoberts and B. R. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana, 71104 and Herbarium, Museum of Life Sciences, Louisiana State University in Shreveport, Shreveport, Louisiana 71115 mmacrobe@Isus.edu As part of a project to develop a checklist of the Louisiana flora, we are examining species reported to be present in Louisiana and checking them against voucher specimens (MacRoberts and MacRoberts 2005, 2006). In this report, we eliminate two more species from previous Louisiana lists. Eryngium heterophyllum. Thomas and Allen (1996), Kartesz and Meacham (1999), USDA (2007), and NatureServe (2007) include Eryngium heterophyllum Engelm. (syn = E. wrightii A. Gray) in Louisiana on the basis of a specimen collected by D.S. and H.B. Correll in St. Martin Parish in 1938 (Correll & Correll 9448 DUKE; Correll and Correll 1941). We examined this specimen and found it to be E. hookeri Walp. The specimen had been previously annotated as E. hookeri by Charles Allen in 2000. Eryngium heterophyllum is a western species occurring no closer to Louisiana than west Texas (Turner et al. 2003, Kartesz and Meacham 2005). Kartesz and Meacham (2005) removed the species from the Louisiana flora but without explanation. Eriocaulon lineare. MacRoberts (1984, 1989), Thomas and Allen (1993), Kartesz and Meacham (1999, 2005), USDA (2007), and NatureServe (2007) include Eriocaulon lineare Small in Louisiana on the basis of two specimens (Correll & Correll 9960 DUKE; Correll and Correll 1941) and (Pruski & Urbatsch 2639 NO). We examined these specimens and found them to be E. decangulare L. and Lachnocaulon anceps (Walt.) Morong, respectively. The Correll and Correll specimen had already been annotated to E. decangulare by Harold Moldenke in 1945 and by Robert Kral in 1992. Eriocaulon lineare is found no closer to Louisiana than Alabama (Kral 2000). Phytologia (December 2007) 89(3) 285 Until voucher specimens to support their inclusion are found, these two species should be excluded from the Louisiana flora. ACKNOWLEDGEMENTS Anne Bradburn, NO, and Michael Windham, DUKE, supplied the plant specimens. Chris Reid, Louisiana Natural Heritage Program, and Anne Bradburn, NO, reviewed the paper. LITERATURE CITED Correll, D.S. and H.B. Correll. 1941. A collection of plants from Louisiana. American Midl. Naturalist 26:30-64. Kartesz, J.A. and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. Kartesz, J.A. and C.A. Meacham. 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical Garden. Chapel Hill. Kral, R. 2000. Eriocaulaceae.Pp. 198-210. In Flora of North America. Vol. 22. Flora of North America Editorial Committee (eds.) Oxford Univ. Press, New York. MacRoberts, B.R. and M.H. MacRoberts 2005. Agrimonia gryposepala (Rosaceae) deleted from the Louisiana flora. Phytologia 87:129- 131. MacRoberts, B.R. and M.H. MacRoberts 2006. Carex stipata (Cyperaceae), Chamaesyce geyeri (Euphorbiaceae), Eurytaenia texana (Apiaceae), Pediomelum esculentum (Fabaceae), and Talinum calycinum (Portulaceae) deleted from the Louisiana flora. Sida 22:1221-1223. 286 Phytologia (December 2007) 89(3) MacRoberts, D.T. 1984. The vascular plants of Louisiana: an annotated checklist and bibliography of the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. Louisiana State University in Shreveport 6:1-165. MacRoberts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci. Louisiana State University in Shreveport 8-9:1-756. NatureServe 2007. www.natureserve.org Thomas, R.D. and C.M. Allen. 1993-1996. Atlas of the vascular flora of Louisiana. Dept. Wildlife and Fisheries, Baton Rouge. Turner, B.L., N. Nichols, G. Denny, and O.Doran. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24:1-888. U.S.D.A. 2007. http://plants.usda.gov Phytologia (December 2007) 89(3) 287 A NEW VARIETY OF ERIOGONUM NUDUM (POLYGONACEAE) FROM CALIFORNIA James L. Reveal Department of Plant Biology, 228 Plant Science, « Cornell University, Ithaca, NY 14853-4301; jlr326@cornell.edu Professor Emeritus, University of Maryland Honorary Curator, The New York Botanical Garden ABSTRACT Eriogonum nudum var. psychicola is established for a population confined to the Antioch Dunes area of Contra Costa Co., California, where it is the primary host for the endangered Lange's metalmark butterfly (Apodemia mormo langei). Allied to var. auriculatum of western California, it may be recognized by its densely pubescent flowers. KEY WORDS: Eriogonum, Polygonaceae, Antioch Dunes, Lange's metalmark butterfly, Apodemia mormo langei. The Antioch Dunes wild buckwheat was mentioned in passing in my Flora of North America treatment of Eriogonum (Reveal 2005: 313). My failure to christen the taxon earlier was because I misplaced the specimens that now serves as the type after it was sent to Maryland. I am grateful to Dr. Charles Delwich for relocating the collection. Eriogonum nudum Douglas ex Benth. var. psychicola Reveal, var. nov. TYPE: UNITED STATES. California, Contra Costa Co., U.S. Fish and Wildlife Service’s Antioch Dunes National Wildlife Refuge (Stamm Unit) east of Antioch, north of railroad tracks in a former vineyard on fine pale brown sand associated with Bromus, Centaurea, Lactuca, Vicia, and Clarkia, ca. N38°00'00", W121°47'50", T2N, R2E, sec. 18, 18 Aug 1997, B. Ertter 15766 (Holotype: UC; isotypes: BH, BM, BRY, CAS, GH, MO, NY, OSC, RSA, TEX, US, UTC, WTU. A var. auriculato floribus dense pubescentibus differt. 288 Phytologia (December 2007) 89(3) Plants erect to spreading perennial herbs, (5) 8-15 (20) dm high, glabrous; /eaves sheathing and occasionally at lower nodes, the leaf-blades 3—7 (9) cm long, 1.54.5 cm wide, densely white tomentose abaxially, floccose or glabrous adaxially, the margins undulate-crisped; flowering stems 2-5 (10) dm long, often stout but not inflated; inflorescences cymose, 3—10 (15) dm long, 1-8 dm wide; involucres solitary or 2—3 in a cluster, (3) 4-5 mm long, glabrous; flowers white to pinkish, 2.5—3.5 mm long, densely pubescent. Sand dunes in coastal grassland communities known presently only from the Antioch sand dunes area east of Antioch and just south of the San Joaquin River in northern Contra Costa Co., California; 3-15 (20) m elev. Flowering Jul-Oct. Other specimens seen: UNITED STATES. California, Contra Costa Co.: Antioch sand dunes, 7 Sep 1965, W. Knight 1175 (CAS); Antioch sand dunes, 17 Aug 1935, E. Lee & A. Carter 1630 (JEPS), 1631 (JEPS); sand dunes E of Antioch, 8 Oct 1947, P. A. Munz 12204 (RSA); 2 mi E of Antioch, 19 Aug 1962, J. Powell 308a (CAS, MIN, UC); sand dunes E of Antioch, 7 Oct 1951, P. Rubtzoff 825 (CAS), 826 (CAS); sand dunes E of Antioch, 14 Sep 1954, P. Rubtzoff 1745 (CAS, UTC), 1746 (CAS); Antioch, 20 Aug 1958, R. W. Thorp 20 (UC). Eriogonum nudum var. psychicola (from the Greek psyche, butterfly, and —cola, dweller) is confined to the Antioch sand dunes area in west central California where it is the primary host for the federally endangered Lange's metalmark butterfly (A4podemia mormo langei J. A. Comstock). It grows in association with an atypical form of Lupinus albifrons Benth., Gutierrezia californica (DC.) Torr. & A. Gray, Quercus agrifolia Née, several introduced invasive species (Bromus diandrus Roth, Centaurea solstitialis L., Lactuca serriola L.), and two federally protected plants, Oenothera deltoides Torr. & Frém. var. howellii Munz (Antioch Dunes evening primrose) and Erysimum asperum (Nutt.) DC. var. angustatum (Rydb.) B. Boivin (Contra Costa wallflower). Historically, the sand dunes have undergone extensive modification due to industrialization to the point that most of the remaining dunes are now confined to the Antioch Dunes National Phytologia (December 2007) 89(3) 289 Wildlife Refuge. Even so, many of the dunes were mined prior to establishment of the Refuge for high-quality sand and requiring the U.S. Fish and Wildlife Service to bring in sand. Also, the wild buckwheat is being “cultivated” on newer dunes to provide additional plants for the butterfly (Nebhan & Buchmann 1996). Antioch Dunes wild buckwheat is allied to those varieties of Eriogonum nudum in California that have leaves with rather strongly crisped leaf margins that sheath up a glabrous flowering stem. A. Leaf-blades densely woolly abaxially and grayish tomentose adaxially; involucres 5—10 per cluster; rare, Sierra Nevada, Tulare eM set, 5 isch miut ou tna shaky cnuie pean tame: dobacagetithe). var. murinum AA. Leaf-blades tomentose abaxially, glabrous or nearly so adaxially; involucres solitary or 2—S per cluster; widespread, coastal ranges. B. Flowers densely tomentose; rare, Antioch Dunes, Contra RCI EL Whee tek n et cchownc adic wrath otieytd oattans var. psychicola BB. Flowers glabrous or (rarely) sparsely pubescent; flowering stems slender or more often slightly to strongly inflated; widespread, west central California. C. Flowering stems not strongly inflated; involucres (2) 3- 2; owes White 10 PMk oo... canes ee var. auriculatum CC. Flowering stems strongly inflated; involucres solitary; flowers pale yellow to yellow or white.................... ACKNOWLEDGEMENTS I wish to thanks Dr. Barbara Ertter and Margriet Wetherwax for reviewing the paper. Recent work was support by L. R. Heckard Funds from the Jepson Herbarium, University of California, Berkeley. LITERATURE CITED Nabham, G. P. & S.L. Buchmann. 1996. The lonesome evening primrose. Pacif. Disc. 49(3): 27-31. Reveal, J. L. 2005. 44a. Polygonaceae Jussieu subfam. Eriogonoideae Arnott, Encycl. Brittannica (ed. 7), 5: 126. 1832 * Wild buckwheat subfamily. F/. N. Amer. 5: 218-478. 290 Phytologia (December 2007) 89(3) BIOLOGICAL STATUS AND DISTRIBUTION OF THELESPERMA FLAVODISCUM (ASTERACEAE: COREOPSIDEAE) Billie L. Turner Plant Resources Center The University of Texas Austin, Texas 78712 billie@uts.cc.utexas.edu ABSTRACT The biological status of Thelesperma flavodiscum vis-a-vis T. filifolium is discussed, along with the habitat proclivities of each. It is concluded that 7. flavodiscum is a relatively uncommon, well-marked species that mostly occurs in deep sandy soils, while 7. filifolium is a species of calcareous soils, the two taxa rarely occurring in close proximity. A map showing the distribution of T. flavodiscum is provided, along with comments upon new distributional records of the taxon in Arkansas and Louisiana. KEY WORDS: Asteraceae, Thelesperma, Texas, Arkansas, Louisiana. Strother (2006), in his treatment of Thelesperma for the Flora of North America maintained the species, 7. flavodiscum (Shinners) B.L. Turmer, but with the admonition, “Differences between Thelesperma flavodiscum and T. filifolium are subtle; they may be better treated as one species.” The distinctions between the latter two taxa are scarcely subtle, as well documented by Melchert (1963), whose doctoral thesis on Thelesperma (albeit unpublished) was not cited by Strother. The latter author does, however, point out the major differences that mark the species, including that of habit (robust plants mostly 0.5-1.5 m high, vs 10-40 cm) and habitat (deep sandy soils vs clays or silty-clays). Observation of plants in the field by the present author show that the two taxa rarely, if ever, grow intermixed, although their distributions are partially sympatric, largely because of the disjunct distribution of 7. filifolium populations in clay outliers within the sandy forest lands of eastern Texas, as correctly noted by Melchert Phytologia (December 2007) 89(3) 291 (1963). At the time of Melchert’s study, relatively few collections of T. flavodiscum were available in herbaria, and his distribution map of the taxon was necessarily limited. I include here (Fig. 1) a map showing the distribution of 7. flavodiscum, this based upon the cited specimens of Melchert (1963) and plants assembled at LL, TEX since his study. It will be noted, as mapped by the present author, that T. flavodiscum is now known from the closely adjacent states of ARKANSAS (Hempstead County, Kral 65476, TEX; Miller County, Thomas et al. 151,334, TEX) and LOUISIANA (Caddo Parish, Thomas et al. 120,635, TEX). The Hempstead Co. collection from Arkansas was reportedly obtained from a “chalk outcrop,” but perhaps not. As already noted, chalk or calcareous outcrops in Texas harbor plants of 7. filifolium, these growing within the range of T. flavodiscum, presumably in close proximity of each other. It is possible that hybrids between these very different taxa occur upon occasion in such areas. Indeed, the cauline leaves of occasional plants of T. filifolium in eastern Texas (and eastern Oklahoma) resemble those of 7. flavodiscum, but the flowering material of the former are typical of T. /filifolium, possessing sulphur-yellow rays (vs yellow, the disc florets brownish to purplish-brown (vs yellow), not to mention the habital differences. Finally, it should be noted that Thelesperma flavodiscum is relatively rare in eastern Texas, and presumably becoming more so. Attempts to collect again from two previously collected populations of the species in Wilson County Texas (Melchert, in 1962; Turner, in 1965) proved futile in the spring of 2007. Indeed, attempts to re-collect from a population of the species obtained in 1988 from Medina County by Orzell & Bridges (6728 TEX) also proved profitless, this from a well documented locale (roadside park along IH 35 in Carrizo sands). Perhaps T. flavodiscum was rare at these several sites to begin with, but I suspect that continual mowing of the roadsides by the Highway Dept. of the State of Texas over the years concerned has been a factor in their disappearance. ACKNOWLEDGEMENTS I am grateful to my son Matt Turner and his partner, Paul Waller, for assisting me with field work during the spring of 2007. 292 Phytologia (December 2007) 89(3) LITERATURE CITED Melchert, T.E. 1963. Systematics of the genus Thelesperma. Ph.D. dissertation, The University of Texas, Austin Strother, J.L. 2006. Thelesperma, in Fl. N. Amer. 21: 199-203. Fig. 1. Distribution of Thelesperma flavodiscum. Phytologia (December 2007) 89(3) 293 KEYS TO THE FLORA OF FLORIDA - 17, RUELLIA (ACANTHACEAE) Daniel B. Ward Department of Botany, University of Florida Gainesville, Florida 32611, U.S.A. ABSTRACT Ruellia (Acanthaceae) is represented in Florida by 8 species. Ruellia heteromorpha and R. succulenta are endemic to the state, while R. brittoniana and R. ciliatiflora are introduced and _ naturalized. Ruellia noctiflora is rated as endangered. Ruellia brittoniana has been designated an "invasive" species and is rapidly spreading along Florida waterways. A nomenclatural justification is provided for use of R. brittoniana, rather than R. tweediana, as the correct name for the Mexican Petunia. Five species, elsewhere reported for the state, are here excluded. An amplified key is given to the Florida taxa. KEY WORDS: Rvuellia, Acanthaceae, Florida flora. The genus Ruellia (Acanthaceae) in eastern North America was well treated by M. L. Fernald (Rhodora 47: 1-38, 47-63, 69-90. 1945), and his documented record of collections, morphology, and range remains the basis for all later work. Ruellia then became the subject of intense study -- in the field, the test garden, and the laboratory -- by Robert W. Long (USF). His insightful reports have gone far to bring understanding to the Florida species: transplant studies of R. caroliniensis and related taxa in South Florida (Amer. Jour. Bot. 51: 842-852. 1964); the first Florida report of R. ciliatiflora (Rhodora 68: 432-434. 1966); the misapplication of R. humilis (Bull. Torrey Bot. Club 95: 16-27. 1968); the polymorphic R. caroliniensis (J. Arnold Arbor. 51: 257-309. 1970); and the distribution 294 Phytologia (December 2007) 89(3) and genetic relationships of the very different (and endangered) R. noctiflora (Bull. Torrey Bot. Club 98: 16-21. 1971); among others. Yet Long's untimely death in 1976 left still other issues unsettled. Perhaps the most regretted uncertainty centers on the variation observed in south peninsular Florida. Fernald (1945) believed herbarium materials justified recognition of two species endemic to southern peninsular Florida, R. heteromorpha and R. succulenta, while restricting R. caroliniensis and R. ciliosa to north Florida and the upper peninsula. Long (1964), after cultivation of plants from 25 Florida populations, interpreted the observed variation to represent only a single species, R. caroliniensis. But, following further opportunity to observe the Florida plants, Long (1970) partitioned the single species into two subspecies -- ssp. caroliniensis which he divided into var. caroliniensis and var. succulenta; and ssp. ciliosa, in turn divided into var. ciliosa and var. heteromorpha. D. C. Wasshausen (Castanea 63: 99-116. 1998), in a careful synopsis of southeastern species, accepted Long's analysis. Other Florida authors have followed somewhat different pathways. Wunderlin & Hansen (2003) recognized Ruellia caroliniensis and R. ciliosa as distinct; they also recogized R. succulenta into which they merged R. heteromorpha_ without distinction. As indicated by the following key, the present author views the four taxa as worthy of specific recognition, returning to the interpretation of Fernald. One species, Ruellia brittoniana, has become popular in garden and patio cultivation and is now known in diverse flower colors. What apparently is the original blue-violet flowered Mexican species has become extensively naturalized and invasive along Florida waterways to such an extent that effort is being made to find biological controls, a task made more difficult by the potential threat posed to a commercially valuable horticultural species. The Mexican species has been known as Ruellia brittoniana Leonard since 1945 when its convoluted nomenclatural history was Phytologia (December 2007) 89(3) 295 fully elucidated (ca. 1800 words, including quotation of all relevant original sources) by Fernald (1945). In outline: Nees (1847) briefly described Cryphiacanthus angustifolius; he cited two collections, one by Galeotti from Jalapa, Mexico, the other by Tweedie from Entre Rios, Argentina. These collections have since been consistently interpreted as representing distinct species, leading Grisebach (1879) to rename the Argentine plant Ruellia tweediana, with a brief description. (In modern parlance, Grisebach followed the spirit of I.C.B.N., Art. 9A.5, in segregating one of the elements as another taxon, and thus by implication designated the "residue" as the lectotype of C. angustifolius.) Britton (1893), addressing the plants of Paraguay, in recognition that the original epithet, "angustifolius," was a later homonym in Rvuellia (not R. angustifolia Sw., 1788) and_ thus unavailable and seemingly unaware of Grisebach's assignment of the name R. tweediana, again renamed the South American plant, as Ruellia spectabilis. Leonard (1941), noting that Nees's "angustifolius," as well as Britton's "spectabilis", had already been used in Ruellia, and needing a name for the Mexican plant, formed still another name, Ruellia brittoniana. Fernald (1945) then re-described the Mexican and Argentine plants in full (Latin) detail, and used R. brittoniana and R. tweediana as their names. A recent statement (30 words) by Wunderlin (1998: 662) that the Florida plant should be known as Ruellia tweediana is incorrect. He noted that R. tweediana Griseb. predates R. brittoniana Leonard; it does indeed, but the first is the South American species, while the second is the related but clearly distinct North American taxon. He remarked inter alia that R. brittoniana is illegitimate since it is based on the same type as R. tweediana; it is not, for the two names are based on the two different specimens, treated together by Nees but segregated by Grisebach. Though Fernald's descriptions are the first to establish a proper understanding of the morphology of these two species, the diagnoses by Nees and Grisebach, though scant, coupled with the unambiguous collections cited, are nomenclaturally sufficient to form legitimate names. The types on which the names are based have not been changed 296 Phytologia (December 2007) 89(3) (Art. 47.1), and the oft-used author citation, "Leonard ex Fernald," for R. brittoniana is inappropriate. [Though not a responsibility of Florida botanists, the present I.C.B.N. (Rec. 60C.1) indicates the Argentine plant, in honor of James Tweedie (1775-1862), is properly spelled R. tweedieana. | RUELLIA L. Wild Petunias 1. Cauline leaves linear to narrowly linear-lanceolate, to 25 cm. long and 2 cm. broad, sessile; inflorescence of elongate axillary peduncles; flowers 1-several, terminal; corollas 3-4 cm. long, blue-violet. Perennial herb, to | m. tall. Stream banks, pond margins. Rare in panhandle, frequent throughout peninsula. Spring-fall. INVASIVE. [Ruellia coerulea, misapplied; Ruellia malacosperma, misapplied; Ruellia tweediana, misapplied] MEXICAN PETUNIA. * Ruellia brittoniana Leonard 1. Cauline leaves broad, variously petiolate. 2. Leaves broadly ovate, the blades abruptly truncate at base, to 10 cm. long and 5 cm. broad, with petioles to 2 cm. long; inflorescence a terminal panicle, densely glandular- pubescent; corollas pale blue-violet, pubescent. Perennial herb, to 1 m. tall. Waste areas, margins of cultivated fields. Central and south peninsula (Hillsborough, Dade counties); rare. Spring-fall. [Ruellia lorentziana Griseb.] * Ruellia ciliatiflora Hook. 2. Leaves, if ovate, with blades not abruptly truncate at base, and petioles under | cm. long; inflorescence not a terminal 1. This paper is a continuation of a series begun in 1977. The "amplified key" format employed here is designed to present in compact form the basic morphological framework of a conventional dichotomous key, as well as data on habitat, range, and frequency. Amplified keys are being prepared for all genera of the Florida vascular flora; the present series is restricted to genera where a new combination is required or a special situation merits extended discussion. Phytologia (December 2007) 89(3) 297 panicle, not densely glandular-pubescent; corollas not pubescent. 3. Flowers terminating simple to sparingly branched elongate near-leafless axillary peduncles; corollas blue-purple, to 4 cm. long. Perennial herb, to 30 cm. tall. Bogs, seasonally wet pinelands. Central panhandle (Gadsden, Liberty, Washington counties); rare. Summer. [Ruellia pedunculata Torr. & Gray ssp. pinetorum (Fern.) R. Long] Ruellia pinetorum Fern. 3. Flowers subsessile or in short-peduncled glomerules, in axils of uppermost leaves. 4. Corollas white, 6-10 cm. long, opening at dusk. Perennial herb, to 50 cm. tall. Wet pinelands. Central panhandle (Jackson, Franklin, Wakulla counties), disjunct to northeast Florida (Clay, Duval, Nassau, St. Johns counties); rare. Summer. ENDANGERED (State listing) NIGHT-FLOWERING WILD PETUNIA. Ruellia noctiflora (Nees) Gray 4. Corollas lavender or bluish, sometimes with white lobes, 4-6 cm. long, opening in daytime. 5. Stems very short, usually under 10 cm. tall; leaves crowded, with narrowed bases and rounded tips, the veins white-villous. Perennial herb, to 10 cm. tall. Dry pinelands, sandhills. North Florida (s. in peninsula to Hernando, Lake counties); frequent in panhandle, infrequent southward. Summer-fall. Occasional plants with stems elongate, the leaves spatulate, remote, are apparent hybrids with R. caroliniensis. [Ruellia humilis, misapplied; Ruellia caroliniensis (Walt. ex Gmel.) Steud. ssp. ciliosa (Pursh) R. Long] DWARF WILD PETUNIA. Ruellia ciliosa Pursh 5. Stems normally elongate, above 20 cm. tall; leaves well- separated, ovate to elliptic with blunt to acute tips, usually with short but distinct petioles, the surfaces variously pubescent to glabrous, the veins not notably more so. 298 Phytologia (December 2007) 89(3) 6. Plant glabrous or nearly so, upright, the leaves thick, somewhat fleshy, often purple-tinged. Perennial herb, to 30 cm. tall. Moist to dry soils, occasionally in rocklands. South peninsula (Collier, Dade, Monroe counties); infrequent. All year. Endemic. [Ruellia caroliniensis (Walt. ex Gmel.) Steud. var. succulenta (Small) R. Long] Ruellia succulenta Small 6. Plant variously pubescent, upright or with long-spreading lower stems, the leaves thin, full green. 7. Stems upright, either simple or with short lateral branches, sparingly pubescent. Perennial herb, to 80 cm. tall. Mesic hammocks, brushy margins. Throughout; common north and central, rare south. Spring-fall. [Ruellia parviflora (Nees) Britt.] COMMON WILD PETUNIA. Ruellia caroliniensis (Walt. ex Gmel.) Steud. 7. Stems in early season upright, in mid to late season forming long trailing horizontal branches from lower nodes, hirsute to villous. Perennial herb, to 40 cm. tall. Dry sandy soils, pinelands, occasional weed in plantings. South peninsula (n. to Lee, Brevard counties); frequent. All year. Endemic. [Rwellia caroliniensis (Walt. ex Gmel.) Steud. var. heteromorpha (Fern.) R. Long; Ruellia hybrida, misapplied] Ruellia heteromorpha Fer. Excluded names: Ruellia humilis Nutt. Northern. Reported for Florida by Small (1933), who had this plant confused with the coastal plain R. ciliosa (Long, 1970). Ruellia malacosperma Greenm. Reported for Florida by Small (1933), apparently based on specimens of R. brittoniana. Cultivated in the state, but not known to escape. Phytologia (December 2007) 89(3) 299 Ruellia nudiflora (Gray) Urban Western. Reported to occur in "pinelands," Dade County, by Lakela & Craighead (1965); omitted without comment by Long & Lakela (1971). Ruellia strepens L. Northern and western. Reported for Florida by Small (1933). No documenting specimens are known. Ruellia tuberosa L. An occasional weed in shadehouses, Dade County, but yet unknown outside. 300 Phytologia (December 2007) 89(3) WHAT IN THE WORLD DID THOMAS WALTER MEAN BY Xxxxx yyyyy? PART TWO: THE QUITE DOUBTFULS Daniel B. Ward Department of Botany, University of Florida Gainesville, Florida 32611, U.S.A. ABSTRACT Flora Caroliniana, published by Thomas Walter in 1788, was the first treatment of American plants to employ the binomial nomenclature and sexual classification system of Linnaeus. Many of its species were new to science, and their names remain important in documentation of the southeastern American flora. But Walter kept no herbarium, and certain of his plants have not been confidently matched with modern names. Assistance of select readers is requested to determine to the extent possible what plants Walter must have had in hand when he named and described 33 of his insecurely identified new species. REQUEST FOR TAXONOMIC ASSISTANCE Thomas Walter published over 400 new names in his pioneer Flora Caroliniana (1788). Many of these names were accompanied by clearly written descriptions, and represent familiar species in the Southeastern flora. But the descriptions of others were so brief or cryptic that later generations of botanists have remained uncertain of what Walter had intended. A small number have remained unidentified even to genus (Ward, 2007a). A larger number, addressed here, strongly indicate the genus to which they belong but are unclear as to species. An annotated index of all of Walter's new species is now in preparation. The majority of Walter's names have been adequately identified. A few will inevitably remain intransigent and will be termed Phytologia (December 2007) 89(3) 301 irretrievably ambiguous and unusable for most nomenclatural purposes. But others, though not immediately suggesting the plant Walter intended, can be salvaged at least to the level of being assigned appropriate synonymic status. The following tabulation of 33 species described by Walter includes (a) Walter's original name, (b) Walter's Latin description, (c) known species, of the same or related genera (Walter's name is in quotes if different from the modern name), (d) the modern name, as best can be ascertained, and (e) comments and information that may aid in identification. "Spm." references are to specimens in the Fraser/Walter herbarium, London (Ward, 2006a). As in the preceding tabulation of especially intractable names (Ward, 2007a), the request is made that persons familiar with the Southeastern flora review this listing of these somewhat less refractory names and report ANY DESCRIPTIONS THAT CAN BE IDENTIFIED. Your speculations and suggestions will be appreciated. [E-mail: DBW, c/o kperkins@flmnh.ufl.edu] LIST OF SPECIES Walter's name: Actaea pentagyna Walter (p. 151) Walter's description: floribus solitariis, pedunculis e sinu foliorum ortis; corollis petalis septem obovato-oblongis, albis; pericarpio lanceolato monospermo; foliis biternatis, foliolis obtusis tridentatis. Related species also listed: Cimicifuga racemosa ("Actaea monogyna"), Aruncus dioicus ("Actaea dioica") Modern name: Perhaps Cimicifuga americana Michx. Identified as Cimicifuga americana by L.K. (1893), but not accepted by American authors. The uncertain identification makes a neotype undesirable; were one selected, Walter's name might displace that of Michaux (1803). Cimicifuga americana is frequent in western NC mountains, unknown in SC. 302 Phytologia (December 2007) 89(3) Walter's name: Aira aegilopsoides Walter (p. 78) Walter's description: spicis secundis; corollarum glumae valvula altera ovata, acuminata, altera columnari, obtusa. Related species also listed: Triplasis purpurea ("Aira purpurea"), Melica mutica, Cenchrus incertus ("Cenchrus carolinianus") Modern name: Perhaps Chloris petraea Sw. No type of Aira aegilopsoides Walt. is known. Spm. 112-A was labeled "Aegilops an Aira ?" by Walter; it was tentatively identified as Chloris petraea by Hitchcock (1905: 47). Chloris petraea is occasional along the SC coast. Walter's name: Anonymos ciliat[a] Walter (p. 197); nom. illegit. Walter's description: caule laevi 3-pedali, foliis linearibus ciliatis, floribus pedunculis brevibus subimbricatis purpureis spicatim positis, calycibus multifloris squamis minus obtusis minus conniventibus, caulibus simplicibus. Related species also listed: ?Liatris sp. ("Anonymos pilosa"), ?Liatris sp. ("Anonymos ramosa"), Carphephorus paniculatus ("Anonymos paniculata"), Carphephorus tomentosus ("Anonymos uniflora"), Carphephorus odoratissimus ("Anonymos odoratissima") Modern name: Probably Liatris sp. No type of Anonymos ciliata Walt. is known. No neotype should be selected. Walter's name: Anonymos erect[a] Walter (p. 170); nom. illegit. Walter's description: foliis linearibus pedunculo brevioribus, floribus axillaribus purpureis. Related species also listed: Agalinis purpurea ("Anonymos purpurea"), Agalinis setacea ("Anonymos setacea"’) Modern name: Possibly Agalinis obtusifolia Raf. No type of Anonymos erecta Walt. is known. No neotype should be selected. Pennell (1935) concluded he was unable to identify Walter's Anonymos erecta [= Agalinis erecta Walt. ex Pennell]. He replaced it with Agalinis obtusifolia Raf. (a name of equally dubious antecedents), and other authors have followed. Agalinis obtusifolia is Phytologia (December 2007) 89(3) 303 infrequent in eastern SC. Walter's name: Anonymos pilos[a] Walter (p. 197); nom. illegit. Walter's description: caule piloso 3 ad 5-pedali, foliis lineari-acerosis utrinque pilosis, floribus sessilibus subimbricatis spicatim positis purpureis. Calycibus multifloris squamis adpressis, caulibus simplicibus. Related species also listed: ?Liatris sp. ("Anonymos ciliata"), ?Liatris sp. ("Anonymos ramosa") Modern name: Probably Liatris sp. No type of Anonymos pilosa Walt. is known. No neotype should be selected. Walter's name: Anonymos ramos{[a] Walter (p. 198); nom. illegit. Walter's description: caule subramoso 4-pedali, pubescente, foliis lanceolatis, floribus subsessilibus purpureis spicatim positis, calycibus multifloris squamis obtusis subconniventibus. Related species also listed: ?Liatris sp. ("Anonymos ciliata"), ?Liatris sp. ("Anonymos pilosa") Modern name: Possibly Liatris sp. No type of Anonymos ramosa Walt. is known. No neotype should be selected. Walter's name: Cineraria caroliniensis Walter (p. 207) Walter's description: floribus paniculatis, foliis petiolatis oblongis denticulatis laevibus, caule herbaceo. Related species also listed: Conyza canadensis ("Cineraria canadensis") Modern name: Perhaps Conyza parva Cronq. [= Conyza canadensis (L.) Crong. var. parva Cronq.; Erigeron canadensis L. var. pusilla (Nutt.) Ahles] No type of Cineraria caroliniensis Walt. is known. Identification here is partially based on Walter's separate recognition of Conyza canadensis (as Cineraria canadensis). Both it and Conyza parva are common throughout. Conyza parva and C. canadensis are similar and often occur together, but seem not to intergrade; they merit specific status. Walter's name is prior to that of Cronquist (1943), 304 Phytologia (December 2007) 89(3) though not to that of Linnaeus (1753). The only possible specimen (spm. 13-B) appears to be C. canadensis. Conyza parva is common throughout. Walter's name: Collinsonia serotina Walter (p. 65) Walter's description: fol. magnis oppositis ovatis, petiolis longis, supremo pari unice sessili, cordato; panicula terminali ramosissima. Related species also listed: Collinsonia canadensis ("Collinsonia praecox") Modern name: Collinsonia sp. No type of Collinsonia serotina Walt. is known. The name was rejected as ambiguous by Peirson et al. (2006). Walter's name: Cucubalus polypetalus Walter (p. 141) Walter's description: foliis oppositis, ovato-lanceolatis; floribus polypetalis. Related species also listed: Silene antirrhina, Silene catesbaei Modern name: Perhaps Silene cucubalus Wibel A single crumpled flower in the Fraser/Walter herbarium was identified as "Walter's type" of Cucubalus polypetalus Walt., by Fernald & Schubert (1948: 198; plate 1105). Corrected to lectotype, by Ward (2007b). But Walter's description of C. polypetalus suggests another species, perhaps Silene cucubalus A. W. Wibel (1799). See discussion elsewhere (Ward, 2006b). Silene cucubalus is frequent in the mountains of western NC, but is unknown in SC. Walter's name: Dianthera ensiformis Walter (p. 63) Walter's description: spicis subcapitatis, pedunculo longo solitario, flor. rubris, fol. ensiformibus. Related species also listed: Justicia ovata ("Dianthera ovata") Modern name: Possibly Justicia americana (L.) Vahl [= Dianthera americana L.] No type of Dianthera ensiformis Walt. is known. Elliott (1816: ) renamed it Justicia ensiformis (Walt.) Ell. LK. (1893) listed Walter's name as synonym of Dianthera americana. Yet the flowers of Phytologia (December 2007) 89(3) 305 D. americana are white and pale lavender, not red. Justicia americana is absent on the SC coastal plain, rare inland. Walter's name: Dianthus carolinianus Walter (p. 140) Walter's description: floribus aggregatis pedunculis longis, squamis tubo dimidio minoribus. Related species also listed: Arenaria caroliniana Modern name: Perhaps Dianthus armeria L. No type of Dianthus carolinianus Walt. is known. Dianthus armeria is unknown in eastern SC, but frequent to west. Walter's name: Eupatorium Marrubium Walter (p. 199) Walter's description: foliis ovato-deltoidibus obtuse serratis pubescentibus glabris sessilibus. Related species also listed: Eupatorium fistulosum ("Eupatorium trifoliatum"), Eupatorium purpureum ("Eupatorium fusco- rubrum"), Eupatorium sessilifolium, Eupatorium album, Eupatorium hyssopifolium ("Eupatorium linearifolium"), Eupatorium pilosum, Eupatorium rotundifolium, Eupatorium capillifolium ("Eupatorium Foeniculoides"), Eupatorium compositifolium, Eupatorium aromaticum ("Eupatorium cordatum"), Eupatorium perfoliatum, Eupatorium ?rugosum ("Eupatorium odoratum"), Conoclinum coelestinum ("Eupatorium coelestinum"), Fleischmannia incarnata ("Eupatorium incarnatum") Modern name: Perhaps Eupatorium rotundifolium L. No type of Eupatorium Marrubium Walt. is known. Elliott (1822: 300) said of Eupatorium rotundifolium, "I have always suspected this plant to be the E. Marrubium of Walter." Eupatorium rotundifolium is common in eastern SC. Walter's name: Festuca multiflora Walter (p. 81) Walter's description: repens, paniculis erectis ovatis, spiculis 8 ad 40- floris, acutis, foliis angustis, acutis, fauce subplumosis. Related species also listed: Festuca ?sciurea ("Festuca quadriflora"), Festuca octoflora, Bromus sp. ("Bromus ciliatus") Modern name: Possibly Distichlis spicata (L.) Greene 306 Phytologia (December 2007) 89(3) No type of Festuca multiflora Walt. is known. Hitchcock (1905: 52) suggested Walter's description was of Leptochloa fascicularis (Lam.) Gray -- unlikely since that species is unknown in SC. He later (1951: 856) considered it a tentative synonym of Distichlis spicata (L.) Greene, common along the SC coast. Walter's name: Gratiola inaequalis Walter (p. 61) Walter's description: fol. oblongis obtusis subdentatis. Cor. labio superiore breviore suberecto; flor. pedicellatis, subcaeruleis. Related species also listed: Mecardonia acuminata ("Gratiola acuminata"), Gratiola ramosa, Gratiola virginiana, ?Gratiola sp. ("Gratiola Peruviana?"), Bacopa monnieri ("Gratiola Monnieria") Modern name: Perhaps Lindernia anagallidea (Michx.) Pennell No type of Gratiola inaequalis Walt. is known. The name has not been identified with confidence. Pennell initially (1920: 246), on the basis of its distribution in the Carolinas, considered Lindernia anagallidea to be represented by Walter's name; he noted Elliott had so interpreted the name. Later, Pennell (1935) analyzed Walter's description and concluded, "I think that we may leave the identification of Walter's plant as permanently doubtful." Lindernia anagallidea is frequent in eastern SC. Walter's name: Hamamelis monoica Walter (p. 255) Walter's description: floribus monoicis. Related species also listed: Hamamelis virginiana ("Hamamelis dioica," "Hamamelis androgyna") Modern name: Perhaps Fothergilla gardenii Linnaeus in Murray No type of Hamamelis dioica Walt. is known. The leaves of Fothergilla are similar to Hamamelis, and Walter's epithet, monoica, may be his indication of the usually bisexual flowers. Fothergilla gardenii is frequent in eastern SC. Walter's name: Ilex ----- #2 (unnamed) Walter (p. 241) Walter's description: baccis flavis. Phytologia (December 2007) 89(3) 307 Related species also listed: //ex opaca ("Ilex Aquifolium"), Ilex cassine ("Ilex Dahoon"), Ilex myrtifolia, Ilex decidua, Ilex vomitoria ("Ilex Cassine") Modern name: Ilex sp. No type of this //ex is known. No neotype should be selected. Though no other description was given, this may be a yellow-fruited form of //ex opaca, which it immediately follows. Walter's name: Lechea juncifolia Walter (p. 83) Walter's description: foliis radicalibus teretibus, calyce nullo. Related species also listed: Lechea minor, Lechea ?villosa ("Lechea major") Modern name: Possibly Lechea tenuifolia Michx. No type of Lechea juncifolia Walt. is known. Lechea tenuifolia is absent from SC coastal plain, common on the piedmont. Elliott (1816: 185) tentatively equated these two names, and was followed by Hodgdon (1938: 90). Torrey & Gray (1838: 1: 154), however, called it "wholly unknown." Walter's name: Ludwigia ramosissima Walter (p. 89) Walter's description: caule erecto, ramosissimo, rubro; fol. alternis, lineari-lanceolatis, longis; flor. axillaribus, pedicellatis; capsulis turbinato-cubicis angulis menbranaceo-alatis. Related species also listed: Ludwigia glandulosa, Ludwigia palustris ("Ludwigia apetala"), Ludwigia alternifolia, Ludwigia linearis, Ludwigia decurrens, Ludwigia pilosa, Ludwigia arcuata, Ludwigia ?sphaerocarpa ("Ludwigia rudis"), Ludwigia suffruticosa Modern name: Perhaps Ludwigia alternifolia L. No type of Ludwigia ramosissima Walt. is known. Munz (1944: 158) suggested it was a second description of Ludwigia alternifolia L. (which Walter described under that name immediately preceding), though no argument was put forward to exclude other Carolina Ludwigia not recognized by Walter. Ludwigia alternifolia is common throughout. 308 Phytologia (December 2007) 89(3) Walter's name: Melanthium spicatum Walter (p. 125) Walter's description: spica nutante, flor. hermaph. radice fibrosa, fol. caulinis subovatis. Related species also listed: Zigadenus glaberrimus ("Melanthium virginicum"), Melanthium hybridum, Veratrum virginicum ("Melanthium monoicum"), Zigadenus muscaetoxicum ("Melanthium Muscaetoxicum"), Chamaelirium luteum ("Melanthium dioicum"), Tofieldia racemosa ("Melanthium racemosum?") Modern name: Possibly Xerophyllum asphodeloides (L.) Nutt. [= Xerophyllum setifolium Michx.] No type of Melanthium spicatum Walt. is known. Suggested by LK. (1894) to be Xerophyllum setifolium (now X. asphodeloides). But Walter stated cauline leaves to be "subovatis," while X. asphodeloides leaves are linear, almost acicular. Xerophyllum asphodeloides is very rare in western NC and SC. Walter's name: Menispermum carolinianum Walter (p. 248) Walter's description: caule fruticoso volubili, foliis lobato-cordatis, racemis axillaribus. Related species also listed: Menispermum canadense ("Menispermum virginicum") Modern name: Perhaps Calycocarpum lyonii (Pursh) Gray No type of Menispermum carolinianum Walt. is known. Walter synonymized his plant with Cissamp[elos] smilacina L. But Walter's description is of Calycocarpum lyonii. That species, though it ranges both north and south, is unknown in the Carolinas. Perhaps a Fraser discovery, but from where? Walter's name is original (even though not italicized); he was not describing nor intending Menispermum carolinum L. Walter's name: Ophrys trifolia Walter (p. 221) Walter's description: bulbis fasciculatis, foliis radicalibus ovatis submagnis planis, scapo trifolio, floribus albo-viridibus adscendentibus, nectarii labio integro lato subtriangulari. Phytologia (December 2007) 89(3) 309 Related species also listed: Platanthera cristata/ciliaris ("Ophrys fimbriata"), Calopogon barbatus/pulchellus ("Ophrys barbata") Modern name: Perhaps Liparis liliifolia (L.) Richard No type of Ophrys trifolia Walt. is known. Liparis liliifolia is frequent in NC and western SC, unknown in eastern SC. Walter's name: Orchis lata Walter (p. 220) Walter's description: alba, bulbis individis, nectarii labio integro lato revoluto, cornu sublongo tenui. Related species also listed: Habenaria ciliaris/blephariglottis ("Orchis ciliaris"), Pogonia ophioglossoides ("Orchis ophioglossoides"), Platanthera lacera ("Orchis habenaria?") Modern name: Perhaps Habenaria nivea (Nutt.) Spreng. No type of Orchis calcarata Walt. is known. The white flowers suggest Habenaria nivea, which is infrequent in eastern SC. Walter's name: Origanum clinopodioides Walter (p. 165) Walter's description: capitulis rotundatis verticillatis terminalibusque, floribus sessilibus, foliis cordato-ovatis subpetiolatis glabris. Related species also listed: Pycnanthemum incanum ("Origanum incanum"), Pycnanthemum flexuosum ("Origanum flexuosum") Modern name: Pycnanthemum sp. No type of Origanum clinopodioides Walt. is known. Perhaps Pycnanthemum aristatum Michx., as suggested by I.K. (1894), though not reported by that name in Carolina floras. Apparently not Pycnanthemum clinopodioides Torr. & Gray, which does not reach the Carolinas. Walter's name: Panicum speciosum Walter (p. 73) Walter's description: panicula longa erecta geniculata, ramiis 4, 6, s.8 verticillatis simplicibus brevibus, e singulis nodis radiatis; flor. solitarii subsessilibus, suscis, longitudine eorum remotis. Related species also listed: Panicum virgatum ("Panicum coloratum"), Agrostis hyemalis ("Cornucopiae hyemalis") Modern name: Possibly Sporobolus junceus (Beauv.) Kunth 310 Phytologia (December 2007) 89(3) No type of Panicum speciosum Walt. is known. Hitchcock (1905: 38) considered it possible that Walter was addressing Sporobolus junceus, which is frequent in SC. A neotype is not to be desired, since that action would displace Heleochloa juncea Beauv. (1812), the basionym of the familiar later name. Sporobolus junceus is rare on SC coastal plain, common inland. Walter's name: Plantago caroliniana Walter (p. 85) Walter's description: spica floribus confertis. Related species also listed: Plantago virginica Modern name: Perhaps Plantago lanceolata L. No type of Plantago caroliniana Walt. is known. Walter's "flowering spike crowded" well matches Plantago lanceolata. That species is introduced, but was likely familiar in pioneer days, now common throughout. Walter's name: Prasium coccineum Walter (p. 166) Walter's description: foliis subovatis, floribus coccineis. Related species also listed: Physostegia virginiana ("Prasium incarnatum"), Physostegia purpurea ("Prasium purpureum") Modern name: Perhaps Calamintha coccinea (Hook.) Benth. in DC. No type of Prasium coccineum Walt. is known. Identification is uncertain, but scarlet flowers ("floribus coccineis") limit the possibilities. Calamintha coccinea is not known north of GA. Walter's name: Ranunculus nitidus Walter (p. 159) Walter's description: foliis caulinis nitidis trifidis, lobis obtusis, floribus luteis. Related species also listed: Ranunculus recurvatus ("Ranunculus abortivus") Modern name: Perhaps Ranunculus abortivus L. No type of Ranunculus nitidus Walt. is known. Elliott (1821) called this Ranunculus abortivus L. and I.K. (1895) tentatively agreed, although Walter had used that name for another species. Elliott may have believed this description fitted Linnaeus' R. abortivus better than Walter's R. abortivus which was perhaps R. recurvatus. Ranunculus abortivus is unknown on the SC coastal plain, common inland. Phytologia (December 2007) 89(3) 311 Walter's name: Schoenus umbellatus Walter (p. 70) Walter's description: cu/mo subtriquetro folioso, floribus in umbellis compositis. Related species also listed: Rhynchospora glomerata ("Schoenus glomeratus"), Rhynchospora corniculata ("Schoenus umbellatus") Modern name: Perhaps Rhynchospora corniculata (Lam.) Gray No type of Schoenus umbellatus Walt. is known. Elliott (1816: 59) tentatively attributed Walter's name to Rhynchospora longirostris Ell., a synonym of R. corniculata. Rhynchospora corniculata is common throughout. Walter's name: Senecio Tussilaginoides Walter (p. 208) Walter's description: corollis flosculosis, foliis crenatis, infimis cordatis petiolatis superioribus pinnatifidis lyratis. Related species also listed: Erechtites hieracifolia ("Senecio hieracifolius?") Modern name: Possibly Senecio smallii Britt. No type of Senecio tussilaginoides Walt. is known. I.K. (1895) interpreted this species to be Senecio aureus L., which the description fits quite well (lower leaves cordate and petiolate, upper leaves pinnatifid and lyrate). But Walter's description is not original; it is copied directly from that of Linnaeus for Senecio aureus. Yet that species is unknown in eastern SC, while S. smallii is common throughout. Walter's name: Silphium scabrum Walter (p. 217) Walter's description: foliis alternis lato-lanceolatis serratis scabris ciliatis subsessilibus, caule bipedali glabro, pedunculis laevibus. Related species also listed: Silphium compositum ("Silphium laciniatum") Modern name: Possibly Silphium dentatum Ell. No type of Si/phium scabrum Walt. is known. Spm. 98-C was labeled "Si/phium" by Walter and "Scabrum" by Fraser. Silphium asteriscus L. is the more scabrous plant, but is essentially absent from SC. Silphium dentatum is common throughout. 312 Phytologia (December 2007) 89(3) Walter's name: Tragia innocua Walter (p. 229) Walter's description: foliis lanceolatis subdentatis pilosis, spica terminali. Related species also listed: Tragia urens Modern name: Perhaps Tragia urticifolia Michx. No type of Tragia innocua Walt. is known. Tragia urticifolia is frequent in eastern SC. Walter's name: Urtica filiformis Walter (p. 230) Walter's description: foliis oppositis ovatis serratis trinerviis, amentis filiformibus sere longitudine foliorum. Related species also listed: Laportea canadensis ("Urtica canadensis"), Boehmeria cylindrica, Pilea pumila ("Urtica pumila") Modern name: Perhaps Parietaria praetermissa Hinton [= Parietaria floridana Nutt.] No type of Urtica filiformis Walt. is known. Parietaria praetermissa is rare along SC coast. Walter's name: Veronica caroliniana Walter (p. 61) Walter's description: flor. solitariis, pedunculis brevibus; fol. radicalibus obovato-oblongis subincisis, caulinis oblongis subserratis obtusis oppositis; caule suberecto; flore albo. Related species also listed: Veronica serpyllifolia, Veronica arvensis Modern name: Perhaps Veronica officinalis L. No type of Veronica caroliniana Walt. is known. Walter's sectional description, "planta tota tomentosa," suggests Veronica officinalis, a species unknown in coastal plain SC but common in the western, higher part of NC. But that species has racemes of blue flowers, while Walter's plant had solitary white flowers ("flor. solitariis...albo"). Perhaps not a Veronica. LITERATURE CITED Cronquist, A. 1943. The separation of Erigeron from Conyza. Bull. Torrey Bot. Club 70:629-632. Phytologia (December 2007) 89(3) 313 Elliott, S. 1816-1824. A Sketch of the Botany of South Carolina and Georgia. Charleston, South Carolina. Fernald, M. L. & B. G. Schubert. 1948. Studies of American types in British herbaria. Part 1V: Some species of Thomas Walter. Rhodora 50: 190-208, 217-229. Hitchcock, A. S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rep't 16:31-56. Hitchcock, A. S. & A. Chase. 1951. Manual of the Grasses of the United States, 2nd ed. U.S. Dept. Agric. misc. publ. 200. 1051 pp. Hodgdon, A. R. 1938. A taxonomic study of Lechea. Rhodora 40: 30- 69, 87-131. Jackson, B. D. 1893, 1894, 1895. Index Kewensis. Cambridge. Munz, P. A. 1944. Studies in Onagraceae -- XIII. The American species of Ludwigia. Bull. Torrey Bot. Club 71: 152-165. Peirson, J. A., P. D. Cantino & H. E. Ballard. 2006. A taxonomic revision of Collinsonia (Lamiaceae) based on phenetic analyses of morphological variation. Syst. Bot. 31: 398-409. Pennell, F. W. 1920 ["1919"]. Scrophulariaceae of the southeastern United States. Proc. Acad. Nat. Sci. Phil. 71: 224-291. Pennell, F. W. 1935. The Scrophulariaceae of Eastern Temperate North America. Acad. Nat. Sci. Phil. monogr. 1. 650 pp. Torrey, J. & A. Gray. 1838. A Flora of North America. New York. Walter, T. 1788. Flora Caroliniana. London. 263 pp. 314 Phytologia (December 2007) 89(3) Ward, D. B. 2006a. Thomas Walter Typification Project, I: Observations on the John Fraser folio. Sida 22: 1111-1118. Ward, D. B. 2006b. Silene catesbaei, rather than S. polypetala, the correct name of the endangered Fringed Catchfly. Castanea 71:324-329., Ward, D. B. 2007a. What in the world did Thomas Walter mean by Xxxxx yyyyy? Part one: The complete unknowns. Phytologia 89: 228-235. Ward, D. B. 2007b. Thomas Walter Typification Project, Il: The known Walter types. J. Bot. Res. Inst. Texas 1:407-423. Phytologia (December 2007) 89(3) bf HETEROSPERMA XANTI TRANSFERRED TO THE GENUS BIDENS (ASTERACEAE: COREOPSIDEAE) B. L. Turner Plant Resources Center The University of Texas at Austin Austin, Texas 78712 billie@uts.cc.utexas.edu Bidens xanti (A. Gray) B.L. Turner, comb. nov. Based upon Heterosperma xanti A. Gray, Proc. Amer. Acad. Arts 5: 162. 1861. Bidens coreocarpoides Sherff Bidens xantiana Rose ex Vasey & Rose Heterosperma brandegeei Sherff Heterosperma coreocarpoides (Sherff) Sherff Heterosperma microglossum Sherff As indicated in the above synonymy, this species was included in the genus Heterosperma by several authors. Gray, in his original description, stated, “The disk-achenia, and indeed the whole structure, except the fertile achenia, accord with Bidens.” Vasey and Rose (1890) in their description of Bidens xantiana, and Sherff (1935) in his description of B. coreocarpoides, also retained the taxon in Bidens, albeit with mistaken identifications. Sherff (1955) subsequently placed Heterosperma xanti in the genus Heterosperma where it was retained by most workers until Clement (by annotation, TEX) and Melchert (also by annotation, TEX) again placed it in the genus Bidens, this in agreement with Melchert’s forthcoming treatment of Bidens for Mexico (in prep.). Clement never published the new combination concerned, nor did Melchert. Heterosperma (and the genus Coreocarpus) is distinguished from Bidens largely by its heteromorphic achenes, those of the disc florets differing from those of the ray florets, as noted by Panero (2007), in his key to genera of the tribe Coreopsideae. Regardless, I have included Heterosperma xanti in Bidens because it will not fit comfortably, all characters considered, within the former genus nor in 316 Phytologia (December 2007) 89(3) Coreocarpus, in spite of the fact that the plants concerned possess heteromorphic achenes. Indeed, Melchert and Turner (1990) transferred two species of the genus Coreocarpus (C. gracillima and C. hintonii), as conceived by Smith (1989), into Bidens, and Melchert intended to transfer Heterosperma xanti into the latter genus, as judged by his annotations on herbarium sheets at TEX. In short, the presence of dimorphic achenes in this or that species of Bidens is to be expected. This is also implicit in the work of Kimball and Crawford (2003) and Kimball et al. (2003) whose DNA studies confirm the position of Coreocarpus hintonii within Bidens, and that of Coreopsis cyclocarpa (having heteromorphic achenes) in the genus Heterosperma. Bidens xanti does, however, possess radial achenes similar to those of Heterosperma, if not Coreocarpus; the former, however, has floral traits like those of Bidens, hence the above transfer. LITERATURE CITED Kimball, R.T. and D.J. Crawford. 2004. Phylogeny of Coreopsideae (Asteraceae) using ITS sequences suggests lability in reproductive characters. Molecular Phylogenetics Evol. 33: 127-130. Kimball, R.T., D.J Crawford and E.B. Smith. 2003. Evolutionary processes in the genus Coreocarpus: insights from molecular biology, Evolution 57: 52-61. Melchert, T. and B.L. Turner. 1990. New species names and combinations in Mexican Bidens (Asteraceae: Coreopsideae). Phytologia 69: 20-23. Panero, J.L. 2007. Compositae: Tribe Coreopsideae. /n, Kadereit, J.W. Jeffrey,C. [Eds.], Families and Genera of Vascular Plants, Vol. 8. Smith, E.B. 1989. A biosystematic study and revision of the genus Coreocarpus. (Compositae). Syst. Bot. 14: 448-472. Phytologia (December 2007) 89(3) ak? VASCULAR FLORA OF A LONGLEAF PINE UPLAND IN SABINE COUNTY, TEXAS Thomas C. Philipps U.S.D.A. Forest Service, National Forests and Grasslands in Texas 415 S. Ist. Street, Lufkin, Texas 75901 TPhillipps@fs.fed.US Suzanne Birmingham Walker Azimuth Forestry Services, Inc., 14671 State Highway 87 South, Shelbyville, Texas 75973 Barbara R. MacRoberts and Michael H. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104 and Herbarium, Museum of Life Sciences, Louisiana State University in Shreveport, Shreveport, Louisiana 71115 ABSTRACT We describe the vascular flora of select plots within longleaf pine uplands at Fox Hunter's Hill in the Sabine National Forest in eastern Texas. The eight established 0.1 ha plots contained a total of 196 species and averaged 87.25 species (range 71 to 112) per plot; sixteen 0.001 ha plots averaged 28.75 species (range 17 to 46); and sixteen 0.0001 ha plots averaged 12.44 species (range 5 to 25). A comparison between longleaf pine uplands in central Louisiana and Fox Hunter's Hill shows that they have similar floristic composition. KEY WORDS: longleaf pine, Pinus palustris, longleaf pine uplands, Sabine National Forest, Sabine County, Texas. Longleaf pine uplands are among the most extensively studied and best known ecosystems in the southeastern United States (Marks and Harcombe 1981, Platt et al. 1988, Frost 1993, Peet and Allard 1993, Ware et al. 1993, Streng et al. 1993, Glitzenstein et al. 1995, Noel et al. 1998, Platt 1999, Christensen 2000, Conner et al. 2001). Surprisingly, considering the amount of attention given to this ecosystem and its eponym, relatively little is known about the 318 Phytologia (December 2007) 89(3) herbaceous layer. Either little or no information has been collected or only partial descriptions are available. This is especially true of longleaf pine communities in the West Gulf Coastal Plain (Streng and Harcombe 1982, Bridges and Orzell 1989, Orzell 1990, Harcombe et al. 1993, MacRoberts and MacRoberts 1998, Turner et al. 1999, Haywood et al. 1998, 2001, Haywood and Harris 1999, Van Kley 1999a, 1999b, 2006, MacRoberts et al. 2004a, Lester et al. 2005, Diggs et al. 2006), where far less research has been done than in the Atlantic and East Gulf Coastal Plain (Peet and Allard 1993, Platt 1999, Christensen 2000). In our search of the literature, we were able to find only one detailed study of the floristic composition of longleaf pine uplands in the West Gulf Coastal Plain (MacRoberts et al. 2004a). If management of longleaf pine communities is to be undertaken effectively, more than just eliminating offsite woody vegetation and reintroducing fire may be needed. At a minimum, the herbaceous layer must be known, for historical evidence indicates that many currently rare species were more common prior to recent anthropogenically influenced declines, and if current trends continue, today’s common species may become rare in the near future (Glitzenstein et al. 2001). In order to reconstruct any plant community, whether by adding rare species to intact communities or by restoring badly degraded sites, one must know what was there initially and, while we cannot go back to pre-settlement vegetation, we can at least begin by studying or by documenting today’s best managed sites. Gathering information on the herbaceous layer of longleaf pine uplands is not always easy, since virtually all West Gulf Coastal Plain longleaf pine was cut during the last two centuries (Noss 1988, Frost 1993, Outcalt 1997, Platt 1999, Diggs et al. 2006). At best, second growth exists but even where there is second growth, there is seldom much, if any, herbaceous layer because of shading by shrub growth resulting from fire suppression (Platt et al. 1988, Streng et al. 1993, Olson and Platt 1995, Brewer 1998, Frost 1998, Platt 1999, Haywood et al. 1998, 2001, Drewa et al. 2002). In pre-European North America, longleaf pine extended from Virginia to Texas (Schwarz 1907, Ware et al. 1993, Platt 1999, Conner et al. 2001). In the West Gulf Coastal Plain, it occurred in Louisiana Phytologia (December 2007) 89(3) 319 and Texas. In central and southwestern Louisiana and southeastern Texas there were large tracts of longleaf pine (Eldredge 1934, Smith 1991, Evans 1997, Outcalt 1997), which were cut in the late 19th and early 20th centuries. Over the total original range of longleaf pine, less than 3 percent remains in a semi-natural condition, and most of this is on public land (Frost 1993, Peet and Allard 1993, Bezanson 2000, Van Kley 2006). Information about longleaf pine uplands before the arrival of Europeans can be gleaned from historical descriptions, lumber company records, and from the few acres that have miraculously survived logging, for example, the Wade Tract in Georgia (Evans 1997, Platt 1999). Early travelers write of monospecific longleaf pine uplands in central Louisiana and eastern Texas (MacRoberts et al. 2004a, Diggs et al. 2006). They depict a landscape with widely spaced uneven aged pines, an open canopy with frequent gaps, and a rich herbaceous layer of grasses, composites, and other forbs. There was little or no midstory and little or no woody vegetation. Every one to three years low intensity fires moved through these pinelands, usually in the spring and summer. Since documentation of floristic composition can be found only for a small portion of this community --- notably lacking is documentation for the herbaceous layer --- it was the purpose of this study to locate a longleaf pine upland where the understory appeared to be intact and to obtain a floristic list. While the aim was to gather baseline data, the question of the quality of longleaf pine uplands in the West Gulf Coastal Plain is also briefly addressed (see Conner et al. 2001 for detailed discussion). STUDY SITE Previous surveys of the Texas National Forests and Grasslands in Texas, notably the Sabine National Forest and Angelina National Forest, have pinpointed several high quality longleaf pine uplands (Orzell 1990). One of these is Fox Hunter’s Hill in southern Sabine County, Texas. 320 Phytologia (December 2007) 89(3) Fox Hunter’s Hill is situated in the Mayflower Uplands Landtype Association (LTA). This LTA is associated with the Catahoula formation overlain with sandstones, sandy clays, and volcanic tuffs. Clay outcrops are present as are deep sands and loams. The topography is generally a rolling hill landscape with some steep hills. The LTA is noted for the longleaf-little bluestem herbaceous community, Catahoula barrens (glades), and hillside seeps/bogs (Figure 1). However, Fox Hunter's Hill, like the remainder of longleaf pine uplands in the West Gulf Coastal Plain, is not pristine. Pine stands are generally young, over-stocked, and even-aged; the canopy is dense, with insufficient gaps, and there is often too much shrub and mid-story woody vegetation. Forest Service records indicate that prescribed fire has been introduced mainly in the non-growing season (however, recent Ribure i Shingle Branch Bog occurs ; within Fox Hunter’ S Hill burns have been applied as late as May) and often with long intervals (2-4 years) between ignitions. In spite of these problems, Fox Hunter's Hill (Figure 2) has a diverse ground layer in many places. Phytologia (December 2007) 89(3) 321 Community types at Fox Hunter's Hill include extensive areas of arenic dry uplands, loamy dry mesic uplands, and small patches of xeric sandylands and glades. Along creeks are herbaceous seeps, particularly bogs and baygalls (Orzell 1990, Diggs et al. 2006, Van Kley 2006). High-quality longleaf pine upland is habitat for such al : PF, ’ e > > 406 Fig. 1. Distribution of Xanthisma spinulosum var. austrotexanum, vat. chihuahuanum and var. spinulosum in Texas. Phytologia (December 2007) 89(3) 353 A NEW SPECIES OF VIGUIERA (ASTERACEAE: HELIANTHEAE) FROM OAXACA, MEXICO B. L. Turner Plant Resources Center The University of Texas at Austin Austin, Texas 78712 billie@uts.cc.utexas.edu Routine identification of Mexican Asteraceae has revealed the following novelty: Viguiera paneroi B.L. Turner, sp. nov. Viguierae sylvaticae Klatt similis sed differt capitulis majoribus (10-12 mm altis vs 5-7 mm), captulescentiis capitulis paucioribus (1-2 vs numerosis), et pappo magis prominenti (aristae laterales ca 5 mm longa vs ca 2 mm). Perennial herb or shrublet to 1 m (?) high. Stems sparsely pilose with slender, multiseptate, appressed to spreading, hairs 3-5 mm long, beneath these a shorter array of recurved hairs ca 0.5 mm high. Leaves alternate along the upper stems, 10-14 cm long; petioles 2-4 cm long, grading into the blades; blades ovate, pubescent above and below with recurved hairs, their margins coarsely serrate, the under surfaces atomiferous-glandular with golden globules. Capitulescence of only | or 2 terminal heads on elongate peduncles. Heads ca 6 cm wide across the extended rays. Involucres 10-12 mm high, the outer bracts subequal, linear-oblanceolate with somewhat ovate apices. Receptacle convex, ca 4 mm wide, paleate with stiff lanceolate bracts ca 14 mm long. Ray florets 13, neuter; tube sparsely pubescent, ca 1.5 mm long; ligules yellow, ca 3 cm long, 0.5-0.7 cm wide. Disk florets 40-60; corollas glabrous, the tube ca 0.5 mm long, the throat 4.5-5.5 mm long, markedly 5-nerved, their lobes ca 1 mm long. Achenes (immature) ca 5 mm long, densely appressed silky-white pubescent; pappus of two lateral, rigid awns, 5-6 mm long, between these 4-8 membranous scales 1-2 mm high. 354 Phytologia (December 2007) 89(3) TYPE: MEXICO. OAXACA: Distrito Tehuantepec; Mpio. San Pedro Huamelula, "noroestw [sic] de la laguna del Rosario Manglar, con Conocarpus erectus...Manglar. orilla de laguna." altitud ca 10 m, 15 Oct 2003, Alfredo Saynes V. 3951 ( con Silvia Salas). Holotype: TEX . ADDITIONAL COLLECTIONS EXAMINED: MEXICO. OAXACA: Distrito Tehuantepec: Mpio. San Pedro Huamelula, 25 m, 2 Oct 2003, M. Elorsa C. 7364 (TEX); 30 m, 29 Oct 2003, M. Elorsa C. 7478 (Holotype: TEX). To my knowledge this novelty has no close relatives in Viguiera (sensu Blake 1918), having a very distinctive 2-seriate involucre, the outer bracts longer than the inner and possessed of broad, oblanceolate apices. Vegetatively, it can be compared to V. sylvatica Klatt of the section Diplostichis, which has broad, ovate, markedly petiolate blades, the under surfaces of which contain golden-colored glandular punctations, similar to those found in V. paneroi. Interestingly, the pales of V. paneroi resemble those found in species of Simsia, as do the peculiar long epidermal hairs found on its stems, these readily matched by the long epidermal hairs found on the stems of Simsia foetida. The achenes, however, both in shape and pappus, place the species in Viguiera, but not neatly into any of the series propounded by Blake (1918). The species is named for my colleague, Prof. Jose L. Panero, long-time student of the genus Viguiera and exceptional teacher at The University of Texas, Austin. ACKNOWLEDGEMENTS I am grateful to Guy Nesom for the Latin diagnosis and for reviewing the paper. Jose L. Panero also reviewed the paper and kindly provided helpful suggestions. LITERATURE CITED Blake, S.F. 1918. A revision of the genus Viguiera. Contr. Gray Herb., n. s. 54: 1-205. Phytologia (December 2007) 89(3) Fig. 1. Viguiera paneroi (Holotype: TEX) 353 356 Phytologia (December 2007) 89(3) NEW INFRAFAMILIAL TAXA IN ASTERACEAE Jose L. Panero Section of Integrative Biology, 1 University Station, A6700, The University of Texas, Austin, TX 78712, USA panero@mail.utexas.edu V. A. Funk US National Herbarium, Department of Botany, P.O. Box 37012, Smithsonian Institution MRC 166, Washington, DC 20013-7012, USA ABSTRACT Phylogenetic studies based on chloroplast DNA have recently identified several lineages that we recognize at the tribal and subfamily levels. Subfamilies Stifftioideae and Wunderlichioideae and tribes Hyalideae, Onoserideae, and Wunderlichieae are described. KEY WORDS: Compositae, Stifftioideae, Wunderlichioideae, Mutisioideae, Hyalideae, Onoserideae, Wunderlichieae Molecular studies using 10 loci of the chloroplast DNA and reported elsewhere (Panero & Funk submitted) reveal several clades that require naming to maintain a classification that recognizes only monophyletic groups. We describe the following new taxa formerly placed in Mutisioideae: Mutisieae. These names complement those already published in Panero & Funk (2002), based on the same molecular phylogenetic studies. Stifftioideae (D. Don) Panero subfam. & comb. nov.; basionym: tribe Stifftieae D. Don, Trans. Linn. Soc. London, 16: 291, 1830. Type: Stifftia J. C. Mikan Asteraceae subfamilia ad Mutisioideae similis sed differt (charactera in combinatione) foliis crasse coriaceis rare tenuibus in extremitatibus ramulorum fasciculatis, corollis grandibus _ tubis antherarum exsertis, capitulis phyllariis multiseriatis, stylis brachiis Phytologia (December 2007) 89(3) 357 glabris, cypselis plerumque 10-nervatis, et pappo aliquando vivide colorato. Small trees, shrubs, vines. Leaves alternate, rarely opposite, petiolate, membranaceous to coriaceous, with margins entire, glabrous to pubescent. Capitula terminal, rarely axillary, solitary to large paniculiform cymes, rarely tightly grouped in glomerule-like cymes, homogamous, rarely heterogamous. Involucres narrowly cylindric to hemispheric; phyllaries in 3-several series, imbricate, gradate. Receptacles epaleaceous. Florets hermaphrodite; corollas ligulate, bilabiate or actinomorphic, the lobes sometimes strongly coiled, white, yellow, pink, purple, orange or red; anthers 5, caudate, calcarate; anther appendages apiculate, rarely tapered; styles glabrous, style arms glabrous, rarely papillate or bullate on abaxial surface. Cypselae cylindric; pappi of multiple capillary bristles, sometimes subplumose, mostly white or stramineous, rarely brightly colored, yellow-orange or pink. The subfamily contains 10 genera: Achnopogon Maguire, Steyermark & Wurdack, Dinoseris Griseb., Duidaea S. F. Blake, Eurydochus Maguire & Wurdack, Glossarion Maguire, Gongylolepis R. H. Schomb., Hyaloseris Griseb., Neblinaea Maguire & Wurdack, Quelchia N. E. Br., and Stifftia J. C. Mikan. Wunderlichioideae Panero & V. A. Funk, subfam. nov., Wunderlichieae Panero & V. A. Funk, tribus nov. Type: Wunderlichia Riedel ex Benth. & Hook. f. Asteraceae subfamilia ad Mutisioideae similis sed differt (charactera in combinatione) foliis crasse coriaceis aut deciduis, paginis abaxialibus foliorum aliquando dense pubescentibus albis in facie, corollis grandibus tubis antherarum exsertis, corollis plerumque actinomorphis rare bilabiatis aut ligulatis, capitulis phyllariis multiseriatis, antheris plerumque apiculatis, stylis brachiis papillatis aut laevibus, et cypselis plerumque cylindricis 10-nervatis. Perennial herbs or shrubs. Leaves alternate, petiolate or sessile, coriaceous; blades linear, ovate to pandurate, obovate, with margins entire, undulate to tightly inrolled and then leaves (tubular, 358 Phytologia (December 2007) 89(3) cylindrical), sometimes densely ferrugineous pubescent on abaxial surface. Capitula terminal, solitary, simple dichasia to paniculiform cymes, homogamous or heterogamous. Involucres cylindric to hemispheric; phyllaries in 3-several series, imbricate, gradate. Receptacles paleaceous or epaleaceous. Florets hermaphroditic; corollas bilabiate, actinomorphic, the lobes erect or rightly coiled, pink, white, purple, magenta, lavender, white or cream-colored; anthers 5, caudate, calcarate; anther appendages apiculate or tapered; styles glabrous, the arms glabrous, papillose or bullate abaxially. Cypselae cylindric, 10-ribbed; pappi in 3-4 series, sometimes plumose. The subfamily contains 8 genera: Chimantaea Maguire, Steyerm. & Wurdack, Janthopappus Roque & D. J. N. Hind, Hyalis D. Don ex Hook. & Arm., Leucomeris D. Don, Nouelia Franch., Stenopadus S. F. Blake, Stomatochaeta (S. F. Blake) Maguire & Wurdack, Wunderlichia Riedel ex Benth. & Hook. f. Hyalideae Panero, tribus nov. Type: Hyalis D. Don ex Hook. & Arn. Tribus subfamiliae Wunderlichioideae (in combinatione) distinctus appendicibus antherarum apiculatis, brachiis stylorum laevibus, et pappo et corollis conspicue exsertis supra involucra in speciebus plurimis. Perennial herbs, shrubs, or small trees. Leaves alternate; leaf blades entire, linear to broadly obovate, sericeous to pannose white on abaxial surfaces, margins entire to slightly serrate. Capitula terminal, solitary or in small to compact paniculiform cymes, rarely corymbiform cymes, discoid or radiate. Involucres turbinate to campanulate; phyllaries in 3-multiple series, imbricate, gradate. Receptacles epaleaceous. Florets hermaphroditic; corollas ligulate, bilabiate with adaxial lobes strongly coiled, or actinomorphic, white, burgundy or pink; anthers 5, caudate, calcarate; anther appendages apiculate; styles glabrous; style arms glabrous. Cypselae cylindric to obovoid; pappi of multiple capillary bristles. This tribe is placed in subfamily Wunderlichioideae and contains 4 genera: Janthopappus Roque & D. J. N. Hind, Hyalis D. Don ex Hook & Arm., Leucomeris D. Don, and Nouwelia Franch. Phytologia (December 2007) 89(3) 359 Onoserideae (Bentham) Panero & V. A. Funk, tribus & comb. nov. basionym: subtribe Onoseridinae Benth. & Hook. f., Gen. Pl. 2: 168, 215, 1873. Type: Onoseris Willd. Tribus subfamiliae Mutisioideae (in combinatione) distinctus corollis in morphologia similis et setis paleaceis dimorphis. Annual or perennial herbs, shrubs, sometimes dioecious. Leaves alternate; blades entire, linear to ovate, rarely deltate, suborbicular or obovate. Capitula solitary, of a few dichasia or rarely forming large capitulescences with hundreds of capitula, radiate, rarely discoid. Involucres campanulate to hemispheric, with several series of imbricate phyllaries. Receptacles epaleate, rarely alveolate or fimbrillate. Ray florets female; corollas bilabiate with a 3-toothed outer lobe and 1-2-toothed inner lobe, rarely absentred, orange, purple, pink, white or bicolored white-purple. Disc florets hermaphrodite, fertile or functionally staminate; corollas 5-lobed, the lobes short to long, straight or recurved, equal or unequal in length, sometimes with one lobe enlarged, red, yellow, greenish-yellow, purple, violet, white or pink; anthers 5, caudate, calcarate; styles glabrous, rarely papillose on abaxial surface of style arms. Cypselae cylindric to turbinate, glabrous to pubescent; pappi 2-4-seriate, mostly heteromorphic. Chromosome number, x = 9. This tribe is placed in subfamily Mutisioideae and contains 6 genera including Aphylloclados Wedd., Gypothamnium Phil., Lycoseris Cass., Plazia Ruiz & Pav., Onoseris Willd., and Urmenetea Phil. ACKNOWLEDGEMENTS We thank B. L. Turner and Jim Henrickson for reading the manuscript and providing helpful suggestions. We thank Guy Nesom for providing the Latin diagnoses. Molecular studies and fieldwork were supported by NSF grant 0344116 (to JLP) and Mellon Foundation and Smithsonian Scholarly Studies grants (to VAF). 360 Phytologia (December 2007) 89(3) LITERATURE CITED Panero, J. L. and V. A. Funk. 2002. Toward a phylogenetic subfamilial classification for the Compositae. Proc. Biol. Soc. Wash. 115: 909-922. Panero, J. L. and V. A. Funk. (submitted). The value of sampling anomalous taxa in phylogenetic studies: major clades of the Asteraceae revealed. Mol. Phylogenet. Evol. (submitted) Phytologia (December 2007) 89(3) 361 JUNIPERUS COMPACTA (CUPRESSACEAE) A NEW SPECIES FROM MEXICO Robert P. Adams Biology Department, Baylor University, Waco, TX 76798, USA Robert_Adams@baylor.edu Andrea E. Schwarzbach and Julie A. Morris Department of Biological Sciences, University of Texas at Brownsville Brownsville, TX 78520, USA. M. Socorro Gonzalez Elizondo CIIDIR Unidad Durango, Instituto Politecnico Nacional, Apdo. Postal 738, Durango, Dgo., 34000 Mexico ABSTRACT Recent nrDNA and tmC-tmD sequence data revealed that J. monticola and J. m. f. compacta are not monophyletic, and this prompted additional research using Single Nucleotide Polymorphisms (SNPs). The SNPs data clearly show that J. monticola f. compacta is not conspecific with J. monticola f. monticola. Juniperus monticola f. compacta Mart. is raised to the specific level as: Juniperus compacta (Mart.) R. P. Adams, comb. et. stat. nov. KEY WORDS: Juniperus jaliscana, Juniperus monticola, J. compacta, J. saltillensis, nrDNA, trn C-trnD, SNPs, Cupressaceae Adams (2004), in his monograph of Juniperus, followed traditional classifications in recognizing J. monticola Mart. f. monticola, J. m. f. compacta Mart. and J. m. f. orizabensis Mart. However, DNA sequencing of nrDNA and tmC-tmD for Juniperus (Schwarzbach, et al., 2008) has shed new light on the relationships within this group. One surprising finding was that J. m. f. monticola formed a clade with J. jaliscana whereas J. m. f. compacta formed a clade with J. saltillensis M. T. Hall. These clades were well separated. 362 Phytologia (December 2007) 89(3) To further investigate this problem, additional sequencings of nrDNA and the trnC-tmD cp DNA region were performed to obtain SNPs to reexamine the relationship of J. monticola to J. m. f. compacta. MATERIALS AND METHODS Specimens collected: J. jaliscana, Adams 6846-6848, 12/12/1991, 940 m, 19 km E of Mex. 200 on the road to Cuale, Jalisco, Mexico; J. monticola f. compacta, Adams 6898-6902, 12/21/1991, 3490 m, Cerro Potosi, Nuevo Leon, Mexico; putative J. m. f. compacta, S. Gonzalez et al. 7169a,b 6/17/2006, (=~Adams 11221, 11222), 4000 m, Nevado de Colima, Jalisco, Mexico; J. monticola f. monticola, Adams 6874-6878, 12/20/1991, 2750 m, El Chico National Park, Hidalgo, Mexico; J. saltillensis, Adams 6886-6890, 12/21/1991, 2090m, on Mex. 60, 14 km E. of San Roberto Junction, Nuevo Leon, Mexico. Voucher specimens are deposited at BAYLU. One gram (fresh weight) of the foliage was placed in 20 g of activated silica gel and transported to the lab, thence stored at -20° C until the DNA was extracted. DNA was extracted using the Qiagen DNeasy mini kit (Qiagen Inc., Valencia CA). SNPs obtained from DNA sequencing ITS (nrDNA) and tmC-tmD amplifications were performed in 50 ul reactions using 10 ng of genomic DNA, 3 units Qiagen Taq polymerase, 5 ul 10x buffer (final concentration: 50 mM KCl, 10 mM Tris-HCl (pH 9), 0.01% gelatin and 0.1% Triton X-100), 1.75 mM MgCh, 20 ul Q solution (2X final), 400 uM each dNTP, 1.8 uM each primer and 4%(by vol.) DMSO. Primers (5'-3'): ITS: ITSA = GGA AGG AGA AGT CGT AAC AAG G; ITSB = CIT TTC CTC CGC TFA TIG ATA-TG, ITSA and ITSB primers from Blattner (1999). tmC-tmD: CDFor: CCA GTT CAA ATC TGG GTG TC CDRev: GGG ATT GTA GTT CAA TTG GT CDFor, CDRev primers from Demesure et al. (1995). Phytologia (December 2007) 89(3) 363 CD10F: AAA GAG AGG GAT TCG TAT GGA CD3R: AAC GAA GCG AAA ATC AAT CA CD10F and CD3R primers from Andrea Schwarzbach (per. comm.). The following PCR conditions were used: MJ Research Programmable Thermal Cycler, 45 cycles, 94°C (1 min.), 50°C (1 min.), 72°C (1 min.), with a final step of 72°C (5 min.). The PCR reaction was subjected to purification by agarose gel electrophoresis (1.5% agarose, 70 v, 55 min.). The nrDNA primers (ITSA, ITSB) produced a band of approx. 1120 bp. The internal trnC-trnD primers, CD10F-CD3R produced a band of approx. 850 bp. In each case the band was excised and purified by use of a Qiagen QIAquick gel extraction kit. The gel purified DNA band with the appropriate primer was sent to McLab Inc. for sequencing. Sequences for both strands were edited and a consensus sequence was produced using Chromas, version 2.31 (Technelysium Pty Ltd.). Alignments were made using Clustal W and then manually corrected. Indels were coded with a "-" for the first nucleotide and "I" for succeeding nucleotides such that an indel was treated as a single mutation event. Overall sequences have been deposited in GenBank (Schwarzbach et al., 2008). SNPs analyses Aligned data sets (nrDNA and trmC-tmD) were analyzed by CLEANDNA (Fortran, R. P. Adams) to remove invariant data. Mutational differences were computed by comparing all SNPs, divided by the number of comparisons over all taxa (= Gower metric, Gower, 1971; Adams, 1975). Principal coordinate analysis was performed by factoring the associational matrix using the formulation of Gower (1966) and Veldman (1967). A minimum spanning network was constructed by selecting the nearest neighbor for each taxon from the pair-wise similarity matrix, then connecting those nearest neighbors as nodes in the network (Adams et al., 2003). RESULTS AND DISCUSSION Analyses of the nrDNA sequences revealed 13 SNPs among the taxa. PCO of the SNPs resulted in 3 eigenroots that accounted for 42, 22 and 19 % of the variation among the OTUs. Ordination (Fig. 1) shows 4 364 Phytologia (December 2007) 89(3) groups as J. jaliscana, J. m. f. monticola, J. m. f. compacta and J. saltillensis. Notice that the two alpine plants (NCI, NC2) from Nevado de Colima (4000 m) appear as somewhat intermediate between taxa. PCO 13 SNPs, nrDNA J. saltillensis J. monticola 1 (42%) J. jaliscana 3 (19%) Figure 1. PCO ordination based on 13 SNPs from nrDNA. Dashed lines are the minimum spanning network with the number of nucleotide differences noted on the dashed line. Clearly, J. m. f. compacta (Cerro Potosi) is quite different from J. m. f. monticola. No variation was found among the 3 individuals of J. m. f. compacta (Cerro Potosi), or among the 3 individuals of J. jaliscana. (a single stick is used in Fig. 1 to represent 3 individuals for these taxa). Analyses of a portion of trmC-trnD revealed several indels, with a total of 15 SNPs. PCO ordination extracted 3 eigenroots that accounted for 66, 25 and 5% of the variation, implying that 4 groups were present Phytologia (December 2007) 89(3) 365 (Fig. 2). These four groups are the same groups as with the nrDNA: J. jaliscana, J. m. f. compacta, J. m. f. monticola, and J. saltillensis. However, the two alpine plants from Nevado de Colima (NC1, NC2) had no differences from J. m. f. monticola (- NC1) or from J. jaliscana (- NC2). It is possible that NC1 is of hybrid origin with pollen and cp DNA from J. m. f. monticola and that NC2 is of hybrid origin with pollen and cp DNA from J. jaliscana. 2 (25%) PCO 15 SNPs, trnC-D f. compacta J. monticola +NC1 1 (66%) J. jaliscana +NC2 Figure 2. PCO ordination based on 15 SNPs from trnC-trnD. Dashed lines are the minimum spanning network with the number of nucleotide differences noted on the dashed line. No variation was found within J. jaliscana, J. monticola, or J. m. f. compacta. However, the 3 individuals of J. saltillensis differed among themselves by a single nucleotide. This differs a little from the nrDNA where more variation within taxa was detected. It appears that in 366 Phytologia (December 2007) 89(3) this instance, the cp DNA has not accumulated mutations as quickly as nr DNA. Zanoni and Adams (1976) analyzed leaf volatile oils from several locations of J. monticola, J. m. f. compacta and J. m. f. orizabensis. They reported that the oils from these taxa were rather uniform, except for the oil of J. m. f. compacta from Cerro Potosi. Adams et al. (1980) compared the leaf terpenoids of J. monticola (El Chico), J. m. f. compacta (Nevada de Toluca) and J. m. f. orizabensis (Pico Orizaba). Table 1 shows an abbreviated summary of their results. Several compounds appear to discriminate between the three formas. These compounds include tricyclene, a-pinene, sabinene, a-terpinene, 4-terpineol, borny] acetate, y-terpinene, the eudesmols and 8- a-acetoyxelemol. It should be noted that the sample (average of 5 plants) of J. m. f. compacta was from Nevada de Toluca not Cerro Potosi (as used for the SNPs in this paper). Zanoni and Adams (1976) reported that the leaf oil from Cerro Potosi was quite different from J. m. f. compacta from Nevada de Toluca and Popocatepetl. Table 1. Comparison of volatile leaf oils of J. monticola (El Chico), J. m. f. compacta (Nevada de Toluca) and J. m. f. orizabensis (Pico Orizaba). Several compounds that appear to separate the taxa are indicated in boldface. t = trace (<0.05%). Compound mont. comp. Oriz. tricyclene 0.6 t 0.9 a-pinene 25.8 8.8 6.0 camphene 0.8 t 1.2 verbenene 0.5 - - sabinene t 26.9 t B-pinene 0.8 t t myrcene 2A | 2.8 4-carene 3.3 0.9 oa a-phellandrene t t t 3-carene - - t a-terpinene - 1.8 t Phytologia (December 2007) 89(3) 367 p-cymene t 0.5 t camphene hydrate 0.5 t I. borneol 4.0 2S LF 4-terpineol t 10.1 0.7 a-terpineol t t t piperitone 0.9 t t bornyl acetate 25.6 12.8 48.6 a-terpiny! acetate t t t thymol t t t (E)-caryophyllene - - t germacrene D - t - B-phellandrene os2 0.6 te limonene 12.4 8.0 L32 y-terpinene t 3.3 0.6 p-menth-1(7),3-diene - 0.5 - terpinolene t - 0.5 linalool t t de) cis-sabinene hydrate t 0.6 1.4 camphor 33 1.0 4.2 trans-sabinene hydrate t t 0.7 elemol 25 2.3 1.4 y-eudesmol 1.0 0.6 t p-eudesmol 33 1.4 t a-eudesmol 1.6 0.5 t 8-a-acetoxyelemol 1.4 0.8 t manoy] oxide t 3.0 t manool - 0.6 - It is clear from SNPs of both nrDNA and tmC-trnD cp DNA that Juniperus monticola f. compacta is not allied with J. monticola. In fact, it is as different from J. monticola as several other recognized species (J. jaliscana, J. saltillensis, Figs. 1, 2). It is also different in its volatile leaf oils (Table 1) and its morphology (Adams, 2004; Zanoni and Adams, 1976, 1979), having tightly compacted foliage and being prostrate shrubs. Silba (2006) recognized it as a subspecies (J. m. subsp. compacta (Mart) Silba) but did not discern its affinity to J. saltillensis (due to cryptic variation in the morphology). 368 Phytologia (December 2007) 89(3) Based on the data presented in this paper, it is appropriate to recognize Juniperus monticola f. compacta as a distinct species: Juniperus compacta (Mart.) R. P. Adams, comb. et. stat. nov. Basionym: Juniperus monticola Martinez f. compacta Martinez, Bol. Soc. Bot. Mexico 7: 19 (1948). Compact mountain juniper. Type: Mexico, Volcan Popocatepetl, Martinez 7003 (HOLOTYPE: MEXU!). Distribution: 3000-4300 (-4500) m Sierra Mojada, Coahuila; Cerro Pelado and Ajusco, Distrito Federal; Nevado de Colima, Jalisco; Popocatepetl, Iztaccihuatl, Tlaloc and Nevado de Toluca, Mexico; Cerro Potosi, Nuevo Leon; Malinche, Tlaxcala; and Cofre de Perote, Vera Cruz, Mexico. Synonyms: Cupressus sabinoides H.B.K., Nova Gen. et Sp. Pl. 2:3. 1817. J. mexicana Sprengel, Syst. Veg. 3: #909 (1826), nom. superfl. illeg. J. sabinoides (Kunth) Nees. Linnaea 19: 706 (1847), non Griseb., Spec. Fl. Rumel. 2: 352 (1846). J. sabinoides Humb. (erroneously attributed) in Lindley and Gordon, J. Hort. Soc. 5: 202 (1850). J. monticola Martinez var. monticola f. compacta Martinez, Bol. Soc. Bot. Mexico 7:19 (1948). J. monticola Martinez subsp. compacta (Martinez) J. Silba, J. Int. conifer Preserv. Soc. 13(1): 12 (2006). Several questions remain unanswered concerning the alpine junipers of Mexico. What is the biological status of J. monticola f. orizabensis? Might all the disjunct alpine populations be variants of J. compacta? Do the large leaf oil differences correlate with more wide based genetic differences? Additional collections and analyses are being conducted to address these questions. ACKNOWLEDGEMENTS Thanks to Saul Pandey and Crispin Borunda for lab assistance. This research was supported in part with funds from NSF grant DEB-3 16686 (A. Schwarzbach and R. P. Adams) and funds from Baylor University. Phytologia (December 2007) 89(3) 369 LITERATURE CITED Adams, R. P. 1975. Statistical character weighting and similarity stability. Brittonia 27: 305-316. Adams, R. P. 2004. Junipers of the World: The genus Juniperus. Trafford Publ., Vancouver, B. C. Adams, R. P., A. E. Schwarzbach, and R. N. Pandey. 2003. The Concordance of Terpenoid, ISSR and RAPD markers, and ITS sequence data sets among genotypes: An example from Juniperus. Biochem. Syst. Ecol. 31: 375-387. Adams, R. P., E. von Rudloff, L. Hogge and T. A. Zanoni. 1980. The volatile terpenoids of Juniperus monticola f. monticola, f. compacta and f. orizabensis. J. Nat. Prods. 43: 417 - 419. Blattner, F. R. 1999. Direct amplification of the entire ITS region from poorly preserved plant material using recombinant PCR. BioTechniques 27: 1180-1186. Demesure, B., N. Sodzi and R. J. Petit. 1995. A set of universal primers for amplification of polymorphic non-coding regions of mito- chondrial and chloroplast DNA in plants. Mol. Ecol. 4:129-131. Gower, J. C. 1966. Some distance properties of latent root and vector methods used in multivariate analysis. Biometrika 53: 326-338. Gower, J. C. 1971. A general coefficient of similarity and some of its properties. Biometrics 27: 857-874. Schwarzbach, A. E., R. P. Adams and J. A. Morris. 2008. Phylogeny of Juniperus based on nrDNA and trnC-trnD sequences. (in prep). Silba, J. 2006. An International census of the coniferae, III, a checklist of the Cupressaceae, Part one (Actinostrobus, Callitris, Diselma, Fitzroya, Libocedrus, Juniperus and Widdringtonia only. .J. Int. Conifer Preserv. Soc. 13: 1-20. 370 Phytologia (December 2007) 89(3) Zanoni, T. A. and R. P. Adams. 1976. The genus Juniperus (Cupressaceae) in Mexico and Guatemala: Numerical and chemosystematic analysis. Biochem. Syst. Ecol. 4: 147-158. Zanoni, T. A. and R. P. Adams. 1979. The genus Juniperus (Cupressaceae) in Mexico and Guatemala: Synonymy, Key, and distributions of the taxa. Bol. Soc. Bot. Mexico 39: 83-121. Veldman D. J., 1967. Fortran programming for the behavioral sciences. Holt, Rinehart and Winston Publ., NY. Phytologia (December 2007) 89(3) 371 TAXONOMIC REVIEW OF THE XANTHISMA SPINULOSUM COMPLEX (ASTERACEAE: ASTEREAE) Guy L. Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4060 gnesom@brit.org Billie L. Turner Plant Resources Center The University of Texas Austin, TX 78712 ABSTRACT A taxonomic review of the Xanthisma spinulosum complex is presented, with county-level (for the U.S.A.) dot maps documenting distribution of the taxa. Xanthisma incisifolium, X. glaberrimum, X. paradoxum (Turner & Hartman) Turner & Nesom, comb. et stat. nov., and X. scabrellum (Greene) Turner & Nesom, comb. nov., do not intergrade with other taxa of the complex and are treated at specific rank. In regions of sympatry, X. spinulosum var. spinulosum, var. gooddingii, var. chihuahuanum, and var. austrotexanum intergrade and are maintained here at varietal rank. Xanthisma spinulosum var. hartmanii Turner & Nesom, var. nov., is described from northern Coahuila, Mexico. KEY WORDS: Asteraceae, Xanthisma, taxonomy Xanthisma spinulosum (Pursh) Morgan & Hartman sensu lato is a species complex in the western United States composed of intergrading infraspecific taxa distributed within two subspecies, as treated by Turner and Hartman (1976, as Machaeranthera pinnatifida). Hartman (2006) treated the complex within the genus Xanthisma in his account for the Flora of North America, using recently modified generic circumscriptions as defined by Morgan and Hartman (2003). Nesom reviewed the complex in Mexico (1990) and in panhandle Texas (2003), in each case concluding that one of the taxa was aie Phytologia (December 2007) 89(3) genetically isolated from the others and justifiably treated at specific rank. Turner (2007) added a taxon at varietal rank from southern Texas, citing collections that appear to represent intergrades with typical X. spinulosum in their region of contact. In the present study, we examine geographic distributions in detail and further evaluate biological integrity and taxonomic rank. Xanthisma glaberrimum at specific rank. Xanthisma spinulosum var. glaberrimum (Rydb.) Morgan & Hartman was elevated to specific rank as X. glaberrimum (Rydb.) Nesom & O'Kennon (Nesom 2003). Hartman (2006), however, retained the taxon as a variety, noting that “it occurs sympatrically with variety spinulosum in the Texas panhandle, where the two taxa behave like biological species” but that “on the eastern plains of Colorado, however, hybridization and autoploidy have been documented (D.B. Hauber 1986).” Xanthisma glaberrimum (diploid, 2n = 8) occurs from the southern panhandle region of Texas northwards into southern Manitoba, Canada (Fig. 1). Over most of this region it is confined to mid-grass regions of the central U.S.A. and only rarely comes in contact with populations of typical X. spinulosum (diploid, 2n = 8), which appears to be largely confined to the more western short grass prairies. The two taxa are sympatric in westernmost Nebraska and adjacent Wyoming, southeastern Colorado, and panhandle Texas and adjacent areas of New Mexico and Oklahoma (Figs. 1, 2). Typical X. glaberrimum in southeastern New Mexico mostly grows on gypsum. In DeBaca and Chaves counties, N.M., some plants have a vestiture of barely perceptible tomentum but they contrast sharply in both vestiture and habit with many collections of typical X. spinulosum from the same area. a. Stems usually unbranched until the upper third, the heads usually distinctly clustered; leaves strictly ascending, narrowly oblong in outline, 1-pinnatifid, midportion 1—2(—2.5) mm wide, lobes oblong- lanceolate to lanceolate or triangular, sometimes shallowly toothed, glabrous or less commonly lightly tomentose, eglandular or less commonly glandular; involucres cupulate................ X. glaberrimum Phytologia (December 2007) 89(3) 373 a. Stems branched from midstem or below to the upper third, the heads diffusely arranged; leaves loosely ascending to spreading, oblong to obovate in outline, especially the basal and lower cauline, 1-2- pinnatifid, midportion 0.5—1.0(-1.5) mm _ wide, lobes linear to lanceolate, usually shallowly toothed, and glandular to varying degrees, slightly to densely tomentose, rarely without eglandular hairs; involucres shallowly hemispheric....................0.0000 var. spinulosum Hauber (1986) noted that putatively hybrid tetraploids he investigated in Otero Co., Colo., show intermediacy toward the ascending stems with greatly reduced lateral branching characteristic of X. glaberrimum (vs. the spreading habit with strong lateral branching in var. spinulosum) as well as intermediacy in their degree of light tomentum. Nesom (2003) observed that in panhandle Texas, where X. glaberrimum and X. spinulosum occur as discrete entities in close proximity, the former is typical in morphology but X. spinulosum commonly shows genetic influence of X. glaberrimum. Morphological discontinuity and apparent isolation of the two taxa at local sites were emphasized in the decision to recognize each of the taxa at specific rank. It is plausible that a significant portion of the Texas panhandle populations of X. spinulosum are tetraploid (likely of hybrid origin, as in Hauber’s study) and as such, effectively isolated from sympatric, diploid X. glaberrimum. Plants mapped as X. glaberrimum in Colfax Co., N.M. (Averett 345, TEX, Lucas 125, TEX) and in Weld Co., Colo. (Raven & Gregory 19521, TEX) also are intermediate between X. glaberrimum and X. spinulosum, but they probably indicate the close proximity of X. glaberrimum. Hybridization occurs among well-marked species in many genera (e.g., among Baptisia species in Texas, Alston & Turner 1963) and, as in the case here with Xanthisma, the occurrence of hybrids does not necessarily make the case for specific negation. Xanthisma spinulosum var. paradoxum as X. paradoxum. Xanthisma spinulosum var. paradoxum (Turner & Hartman) Morgan & Hartman is localized in distribution (Fig. 3) and restricted to the Four Corners region, occurring most abundantly in San Juan Co., Utah. It grows on low, rolling, sparsely vegetated hills formed from the Mancos or Fruitland shale formations (Cretaceous), which produce a 374 Phytologia (December 2007) 89(3) substrate of highly alkaline, gypsiferous clay. Common associates include Atriplex corrugata, A. confertifolia, Frankenia jamesii, and Opuntia polyacantha. At least three species are known to be endemic to this area: Sclerocactus mesae-verdae (Boissevain ex Hill & Salisbury) L. Benson, Proatriplex pleiantha (W. Weber) Stutz & Chu, and Abronia bolackii Atwood, Welsh, & Heil (NM Rare Plant Technical Council 1999). Study of Four Corners Asteraceae by Nesom revealed that morphology of var. paradoxum is consistent and discontinuous from that of var. spinulosum, even where their distribution slightly overlaps (see couplet immediately below). Many collections of var. paradoxum have been made from the vicinity of Hatch Trading Post along Alkali Canyon in San Juan Co., Utah; from that locality, Porter 1255 (SJNM 4260) is var. paradoxa, but Porter 1255 (SJNM 7252) is_ var. spinulosum. The two Porter plants evidently were collected in close proximity. a. Stems 6—15 cm; basal leaves persistent and dense, cauline mostly on proximal half of stems; heads on naked or bracteate peduncles 1-4 cm; involucres 15:2 Sint, 26)in. cage cut seein end ea X. paradoxum a. Stems (10—)15-—30 cm; basal leaves mostly deciduous by flowering, cauline relatively even sized upwards to near heads; heads on bracteate peduncles 0.5—2(—3) cm; involucres 8—12 mm wide.....var. spinulosum Turner and Hartman (1976, p. 314) noted that var. paradoxum "is fairly well-marked and does not seem to intergrade with its more eastern allopatriarch [var. spinulosum], ... it does appear to grade into var. gooddingii to the southwest, although not strikingly so." Var. paradoxum was compared in the original description to var. gooddingii, but as further study has shown, the two taxa are allopatric and do not have the opportunity to hybridize or intergrade. Thus, in parallel with X. glaberrimum, we observe that var. paradoxum is distinct and genetically isolated from X. spinulosum and propose (below) that it be treated at specific rank. Xanthisma spinulosum var. scabrellum as X. scabrellum. Turner and Hartman (1976) observed that Xanthisma spinulosum var. scabrellum intergrades with X. arenarium (Benth.) Phytologia (December 2007) 89(3) 375 Morgan & Hartman to some degree and that X. arenarium might with justification also be treated within X. spinulosum, although they retained it at specific rank. Both taxa are densely glandular. Xanthisma arenarium is restricted to southern Baja California Sur and sympatric there with var. scabrellum, which is more widely distributed and extends northward into the adjacent state of Baja California. In southern Baja California var. scabrellum apparently is parapatric or slightly sympatric with var. gooddingii, which usually is eglandular in the zone of contact with var. scabrellum. The two also are conspicuously different in habit. In the current study, we find that neither var. scabrellum nor X. arenarium intergrades with X. spinulosum and, as Shinners did earlier (1950). Consequently, we treat var. scabrellum at specific rank. Xanthisma incisifolium at specific rank. Plants Xanthisma spinulosum var. incisifolium from a variety of habitats on the islands of San Lorenzo (Baja California) and San Esteban and Tiburon (Sonora) are consistent in morphology and distinct from other X. spinulosum. Xanthisma spinulosum var. gooddingii is the only expression of the species that approaches the geographic range of var. incisifolium and that is sympatric with it (on San Lorenzo and Tiburon). The insular endemic was recognized in a previous study (Nesom 1990) at specific rank as X. incisifolium. Xanthisma spinulosum sensu stricto and varieties. We treat Xanthisma spinulosum as comprising five varieties. Even with recognition of these geographic variants, X. spinulosum var. spinulosum is variable, consisting of a panorama of individuals and/or local populations that have received formal taxonomic recognition. Variation within var. spinulosum may be complex even within a single county of Texas. For example, at one locality (Taylor Co., 7 mi SW of Merkel), plants of X. spinulosum may be found that are completely glabrous with once-pinnatifid leaves (Henderson 63-787, TEX), similar to those of X. glaberrimum, or that are cottony pubescent with similar leaves (Henderson 63-785, TEX). Additionally, plants referable to the "cotula" form (very glandular individuals lacking cottony pubescence) may also occur there, along with various intermediates between the latter and the previously mentioned expressions (LL,TEX). We have recognized all such individuals and/or population segregates in this area 376 Phytologia (December 2007) 89(3) as belonging to var. spinulosum. The distribution of densely glandular plants among the collections at LL,TEX is shown in Fig. 4. Xanthisma spinulosum var. spinulosum intergrades with var. chihuahuanum (Fig. 5) in Mexico, southern Texas, southwestern New Mexico, and southeastern Arizona. In southern Arizona var. spinulosum may intergrade slightly with var. gooddingii (Fig. 7), but this needs detailed investigation. Var. austrotexanum, recently recognized as an endemic of the Rio Grande Valley in southern Texas (Turner 2007), apparently intergrades with var. spinulosum (Fig. 6). Var. hartmanii, first described in the present manuscript, occurs in north-central Coahuila, surrounded on all sides by var. spinulosum (Fig. 5), with which it may intergrade. Status of subsp. spinulosum and subsp. gooddingii. Turner and Hartman (1976) treated Xanthisma spinulosum (= Machaeranthera pinnatifida) as having two allopatric subspecies: subsp. gooddingii, a western assemblage including four varieties and a more eastern subsp. spinulosum with three varieties. Morgan and Hartman (2003) informally divided the species into the two subspecies but did not provide a valid combination for subsp. gooddingii in the new generic position. Because we have elevated three of the four original taxa of subsp. gooddingii to specific rank, and because we observe that var. gooddingii (Fig. 7) may intergrade with var. spinulosum and var. chihuahuanum, it no longer seems useful to formally recognize subspecies among the varieties of X. spinulosum treated here. Key to the taxa of the Xanthisma spinulosum complex. 1. Plants eglandular to sparsely or densely glandular, stems usually at least slightly arcuate, with leaves reduced in size near the heads. (3) 1. Plants densely stipitate-glandular, stems stiffly erect, branches stiffly spreading-ascending, with leaves even-sized and evenly arranged to immediately below the heads. (2) Phytologia (December 2007) 89(3) 377 2. Densely stipitate-glandular, also sparsely to densely villous with 2. Moderately stipitate-glandular, sometimes with a few eglandular hairs; leaves narrowly obovate-oblong, 2-4 mm wide; involucres 9— eR MRRLINON MERE P80 cls cee a Gs bons Sem mv dcedseysihes iecones X. scabrellum 3. Plants glabrous or less commonly lightly tomentose, eglandular or less commonly slightly glandular; stems usually unbranched until the upper third, the heads usually distinctly clustered................... EE aera ry sa akens ervetenin jcUvse anes ctinth s dubetae tended X. glaberrimum 3. Plants usually glandular to varying degrees, slightly to densely tomentose, rarely without eglandular hairs; stems branched from mid-stem or below to the upper third, the heads more diffusely arranged. (4) 4. Involucres mostly 8—15 mm wide (12-22 in var. gooddingii); leaves mostly cauline (if basal persistent, then involucres relatively small). (6) 4. Involucres 15—25 mm wide; leaves mostly basal or basal and lower cauline. (5) 5. Leaf lobes lanceolate to oblanceolate; heads held barely above the level of the leaves, on short, bracteate peduncles; subshrubs with woody, ascending, caudex-like branches................ X. incisifolium 5. Leaf lobes linear; heads above leaves on naked or bracteate peduncles 15-40 mm long; perennial herbs without caudex een Re I Te Ms Rae 4 Made ce nn uetineke RC eek wea X. paradoxum 6. Leaves evenly arranged along entire stem, lobes linear, 4-8 mm long, and usually falcate-recurving; heads epedunculate, usually immediately subtended by leaves; stems 30-50 cm high; stems and leaves inconspicuously granular-glandular, without other vestiture. sight Settee ais et Th 8 lg Did aerate one re Ae var. hartmanii 6. Leaves usually mostly on proximal 2/3, serrate or with lobes of varying length ; heads pedunculate or pedunculate; stems 10—70 cm high; vestiture various. (7) 378 Phytologia (December 2007) 89(3) oo — NO oS) . Plants evidently coarsely or minutely stipitate-glandular, usually without other pubescence; stems with leaves reduced distally, heads pedunculate; involucres mostly 12—22 mm wide. (9) . Plants eglandular or very sparsely and inconspicuously glandular, variably villous; stems relatively uniformly leafy up to heads, heads epedunculate; involucres mostly 8—12 mm wide. (8) . Stems mostly 10-40 cm tall, ascending from the base, intricately branched at midstem; midcauline leaves deeply toothed to divided; persistently thinly floccose tomentose; widespread...var. spinulosum . Stems 30-70 cm tall, stiffly erect from the base, usually branched only near the heads; midcauline leaves shallowly serrate; glabrescent and often glabrous; Brooks, Hidalgo, Kleberg, and Jim Wells gourties: Texans. 66 dic Peed Oe ch ete ahs var. austrotexanum . Entire plants usually densely and coarsely stipitate-glandular; leaf surfaces dull-textured, lobes and teeth antrorse, not falcate; involucres mostly 12-16 mm wide................. var. chihuahuanum . Plants minutely stipitate- to granular-glandular (U.S.) to eglandular (Baja California); leaf surfaces shiny, lobes and sometimes blades (distal cauline) commonly falcate; involucres 12-22 mm wide WARE aes, SSE A i aii a Cites ob Ao ES Sivas iA var. gooddingii . XANTHISMA GLABERRIMUM (Rydb.) Nesom & O’Kennon, Sida 20:1586. 2003. Sideranthus glaberrimus Rydb. [1900]. Machaeranthera pinnatifida var. glaberrima (Rydb.) Turner & Hartman. Xanthisma spinulosum var. glaberrimum (Rydb.) Morgan & Hartman. Sideranthus laevis Woot. & Standl. [1913]; Haplopappus spinulosus subsp. laevis (Woot. & Standl.) Hall; Machaeranthera laevis (Woot. & Standl.) Shinners. . XANTHISMA INCISIFOLIUM (I. M. Johnston) Nesom, Sida 20:1585. 2003. Aplopappus arenarius var. incisifolius 1.M. Johnston; Machaeranthera pinnatifida var. incisifolia (1.M. Johnston) Turner & Hartman; Machaeranthera incisifolia (1.M. Johnston) Nesom. . XANTHISMA PARADOXUM (B. L. Turner & Hartman) Nesom & B. L. Turner, comb. et stat. nov. Based on Machaeranthera pinnatifida var. paradoxa B. L. Turner & Hartman, Wrightia 5:314. 1976. Phytologia (December 2007) 89(3) 379 Type: USA. Colorado. Montrose Co.: Paradox, 21 Jun 1912, E.P. Walker 147 (holotype: NY!, internet image!; isotypes: DS!, GH!, US! internet image!). Xanthisma spinulosum var. paradoxum (B. L. Turner & Hartman) Morgan & Hartman. Stems numerous from base, ascending, 6-15 cm high, stems and leaves thinly tomentose-puberulent, eglandular. Leaves: basal persistent and dense, cauline mostly on proximal half of stems, basal and cauline deeply dissected, lobes linear, spreading-ascending. Heads on naked or bracteate peduncles 15-40 mm long; involucres cupulate, mostly 15-25 mm _ wide; phyllaries linear-lanceolate, minutely puberulent and finely granular-glandular. Washes, clay hills, sandstone, disturbed sites, desert scrub, pinyon-juniper. Arizona, Colorado, Utah; 1400-1800 m; (Mar—)Apr-— Oct; Arizona, Colorado, Utah. 4. XANTHISMA SCABRELLUM (Greene) Nesom & B. L. Turner, comb. nov. Based on Eriocarpum scabrellum Greene, Erythea 2:108. 1894. Type: Mexico. Baja California: Los Angeles Bay, 1887, E. Palmer 539 (holotype: ND-G?; isotypes: US-2 sheets! internet images!). Machaeranthera scabrella (Greene) Shinners; Xanthisma spinulosum var. scabrellum (Greene) Morgan & Hartman; Machaeranthera pinnatifida var. scabrella (Greene) B. L. Turner & Hartman. Plants with a woody caudex; stems, leaves, and phyllaries densely stipitate-glandular. Stems usually spreading from the base, (6-) 15—35(-45) cm tall. Leaves oblong oblanceolate, 5-25 mm long, 1-3 mm wide, relatively even-sized along the stems (smaller immediately beneath heads), shallowly but coarsely toothed. Heads on peduncles 0- 5(—10) mm long; phyllaries usually spreading to recurving at apex. 2n = 8, 16. Sandy roadsides, rocky slopes, shrublands, thorn-forests, short-tree woodlands, 10-400 m; most commonly Jan—Apr but sporadically all seasons; Baja California Sur, Baja California. 380 Phytologia (December 2007) 89(3) 5. XANTHISMA SPINULOSUM (Pursh) Morgan & Hartman, Sida 20:1406. 2003. Amellus spinulosus Pursh (not Machaeranthera spinulosa Greene). 5a. Var. AUSTROTEXANUM B. L. Turner, Phytologia 89:350. 2007 [nom. nov.]. Based on Haplopappus texensis R.C. Jackson; Machaeranthera texensis (R.C. Jackson) Shinners. 5b. Var. CHIHUAHUANUM (B. L. Turner & Hartman) Morgan & Hartman, Sida 20:1408. 2003. Machaeranthera pinnatifida var. chihuahuana Turner & Hartman 2n = 8, 16. 5c. Var. HARTMANIIB. L. Turner & Nesom, var. nov. TYPE: Mexico. Coahuila. 85 mi from San Miguel on Muzquiz road, common along road beneath Acacia - with Hymenoxys, Sorghum, Verbena, Lantana, Aristida, 24 May 1972, R.L. Hartman 3333{c] (holotype: TEX!; isotypes: TEX, RM). A Xanthismo spinuloso var. spinuloso distinctus foliis profunde dissectis lobis linearibus falcato-recurvatis. Stems and leaves inconspicuously — granular-glandular, otherwise glabrous. Leaves relatively even-sized and similar in morphology from base of stem to heads, 2(—3)-pinnately parted with linear and usually falcate-recurving lobes. Heads usually immediately subtended by leaves. Additional collections examined. Mexico. Coahuila. 1 mi W of La Rosita on road to Muzquiz, common in stream bed, 24 May 1972, Hartman 3334 (LL, RM). 3 mi W of Rancho La Rosita, 43 mi NW of Muzquiz, E of Sierra de la Encantada, 22 May 1968, Powell, Patterson & Ittner 1577 (TEX - 2 sheets). These plants have been collected from only a single small area in north-central Coahuila -- they are highly distinct in a morphology not seen anywhere else in the range of the species. Three separate collections indicate the morphology is consistent. Collections of Xanthisma spinulosum from around the city of Muzquiz (Marsh 196- Phytologia (December 2007) 89(3) 381 TEX, Marsh 1196-TEX), about 55 kilometers southeast of typical var. hartmanii, have leaves more deeply dissected than normal and it is possible that these plants show genetic influence of var. hartmanii. On the other hand, their leaves are smaller and less dissected than in var. hartmanii and their vestiture is densely tomentose. Otherwise, plants surrounding var. hartmanii in all directions in Coahuila are var. spinulosum. It is plausible that further field study may show that var. hartmanii would be appropriately treated at specific rank. 5d. Var. GOODDINGH (A. Nels.) Morgan & Hartman, Sida 20:1408. 2003. Sideranthus gooddingii A. Nels.; Machaeranthera pinnatifida var. gooddingii (A. Nels.) B. L. Turner & Hartman Baja California plants are slightly tomentose but completely eglandular or nearly so, while those of Sonora northward into the U.S.A. are minutely stipitate- to granular-glandular and have consistently narrower leaves. The tall and erect stems, large heads, linear leaves with long, sharp teeth, and distinctive vestiture of var. gooddingii are distinctive. It is maintained here within Xanthisma spinulosum because of putative intergrades with var. spinulosum, but these two taxa may prove to be isolated. 2n = 8. Se. Var. SPINULOSUM Diplopappus pinnatifidus Hook.; Machaeranthera pinnatifida (Hook.) Shinners. Sideranthus cotula Small; Haplopappus spinulosus subsp. cotula (Small) H.M. Hall 2n = 8, 16. LITERATURE CITED Alston, R.E. and B.L. Turner. 1963. Natural hybridization among four species of Baptisia (Leguminosae). Amer. J. Bot. 50:159-173. Hartman, R.L. 1990. A conspectus of Machaeranthera (Asteraceae: Astereae). Phytologia 68:439-465. 382 Phytologia (December 2007) 89(3) Hartman, R.L. 2006. Xanthisma. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford University Press, New York and Oxford. Vol. 20, pp. 383-393. Hauber, D.P. 1986. Autotetraploidy in Haplopappus spinulosus hybrids: Evidence from natural and synthetic tetraploids. Amer. J. Bot. 73:1595—1608. Morgan, D.R. 2003. nrDNA external transcribed spacer (ETS) sequence data, reticulate evolution, and the systematics of Machaeranthera (Asteraceae). Syst. Bot. 28:179-190. Morgan, D.R. and R.L. Hartman. 2003. A synopsis of Machaeranthera (Asteraceae: Astereae), with recognition of segregate genera. Sida 20:1837—1416. New Mexico Rare Plant Technical Council. 1999. New Mexico Rare Plants. Albuquerque, NM: New Mexico Rare Plants Home Page. http://nmrareplants.unm.edu (Latest update: 19 April 2007). Nesom, G.L. 1990. Notes on variation in the Machaeranthera pinnatifida complex (Asteraceae) in Mexico with a new combination. Phytologia 69:108—114. Nesom, G.L. 2003. New combinations in Xanthisma (Asteraceae: Astereae). Sida 20:1585—1586. Shinners, L.H. 1950. Notes on Texas Compositae—V. Field & Lab. 18:32—-42. Turner, B.L. 2007. Xanthisma spinulosum var. austrotexanum (Asteraceae: Astereae), an endemic of southernmost Texas. Phytologia 89:349-352. Turner, B.L. and R.L. Hartman. 1976. Infraspecific categories of Machaeranthera pinnatifida (Compositae). Wrightia 5:308-315. 383 Phytologia (December 2007) 89(3) < = @ = = r A 4 >< © E a he © 2 3 D Fig. 1. Distribution of Xanthisma glaberrimum by county in the U.S.A. Phytologia (December 2007) 89(3) 384 E 3 o 2 =| £& a w u ec > E ° Fig.2. Distribution of Xanthisma spinulosum var. spinulosum by county in the U.S.A. Phytologia (December 2007) 89(3) 385 XANTHISMA spinulosum var. © spinulosum 4 var. ® paradoxum Fig.3. Distribution of Xanthisma spinulosum (open circles) and X. paradoxum (closed circles) in the Four-Corners region of the southwestern U.S.A. 386 Phytologia (December 2007) 89(3) tee Fig.4. Distribution of plant and/or population forms of Xanthisma spinulosum in the southcentral U.S.A.: var. spinulosum (open circles); "cotula" form (closed circles); intermediates (half circles). Phytologia (December 2007) 89(3) 387 ae y) 0 Hi i. isl 4 aii ey aan At LY] Fig.5. Distribution of Xanthisma spinulosum in the southwestern U.S.A. and closely adjacent Mexico: var. chihuahuanum (open circles); var. spinulosum (dotted circles). Var. hartmanii is not mapped. Phytologia (December 2007) 89(3) 388 spinulosum & var. austrotexanum E 3 E CI 3 = i] 3 = = 0 = > ° E a = 3 £ e . $ ° Fig.6. Distribution of Xanthisma spinulosum in Texas: var. austrotexanum (triangles); var. chihuahuanum (open circles); var. spinulosum (closed circles). Phytologia (December 2007) 89(3) 389 XANTHISMA spinulosus @ var. goodingii © var. spinulosus O var. chihuahuanum =25° i$? Fig. 7. Distribution of Xanthisma spinulosum var. gooddingii (closed circles). 390 Phytologia (December 2007) 89(3) INDEX TO ARTICLE TOPICS AND SCIENTIFIC NAMES IN VOLUME 89 (New scientific names and combinations printed in bold face) PRCAGTA SP sieh cis Sec ogee covet ete tae cha tees Aas eas TAR a Geaee hare 241-257 Agaloma graminea, COMD. MOV...................0cceceeeeee teens eee eees 226 Aceratina humochica; Sp. WOVs. ci. 66255-0seseaete re. ccessporseneoe tees. 193 Ageratina serbOaha, SPs DOV... (52.5050 obese 0s ovens «canine enseaebiecence 194 Alloispermum guerreroanum, Sp. NOV.................0.:e0c eee eee eees 131 Anzona plant localities: 375 e.diae tsacan Bones secre ase tee tes saies ves 178-189 ASP AIT TAURI ee sie cine « Oe vino cea diel sianase de’ dren sedesnes teas 230 Ap dae SCH DIGIMES: .cescats tenet nakst testers cowed eae sen gan aorngaaee 230 Avctotus care Wntaina,o:5 Sst rk oo ues et poe ete Roe tee dos aaa eee aweneneeteese 230 Are thisar Toliwsat.. a tnsta tyne Modess tesco Seneew oe eh She Dew ee ans vane 230 Asteraceae. iiratamilial Tax «:. i tet ib vcaidhokos oben dneucuess <2 356-360 PAStCE Ve IatSeeiirs pet ihros as wuePiren Banna nasa teh stmndc eau eae Ae Ce aameay Zon Walea Carolimbana rn oa 3 cacceaatuss he ins dew eainch seebeceeeinn maismamaere saesee er 231 Caloreziay: COU. TOV a: 453 6 cis descnistiged 2 suns omaatemnsae reese abana ete cmon 198 Calorezia;: nutans, COMD. NOV. .: 3. 4.