CONTENTS Ethnobotany serving society: A case study from the Sierra de Manantlan Biosphere _ Bruce F. Benz, Judith Cevallos E., Elizabeth Munoz M., and Francisco Santana M. Hybridization in two cisantly related Mexican black oaks Quercus conzattii and Quercus eduardii (Fagaceae: Quercus: Section Lobatae) Jeffrey R. Bacon and eae Spellenberg 17 Invasion and spread of Coincya monensis (Brassicaceae) in North America Robert F.C. Naczi and Jobn W. Thieret 43 Resegregation of Barbieria from Clitoria (Leguminosae: Phaseoleae: Clitoriinae) Paul R. Fantz 55 Es Fuertesimalva, a new genus of neotropical Maly Paul A. Fryxell 69 raceae Robinsonecio (Asteraceae: Senecioneae) a new genus from Mexico and Guatemala Theodore M. Barkley and John P. Janovec 7 A new variety of Swertia radiata (Gentianaceae) James Henrickson 83 Notes on Spigelia (Loganiaceae James Henrickson 89 A new combination in Eragrostis (Poaceae: Eragrostideae) Paul M. Peterson 105 A new combination in Bouteloua (Poaceae Joseph K. Wipff and Stanley D. Jones 109 A new subspecies of Gentianella heterosepala (Gcntianaceae) from Mexico = — José A. Villarreal Q. 111 Contributions toward a new flora of the Philippines: I. A synopsis of the genus Myrsine (Myrsinaceae) John J. Pipoly Il 115 Morphology of Pennisetum orientale (Poaceae: )iniceae) J. Ramu, S.L. Hatch, M.A. Hussey, and EC’. Bashaw 163 New speeies of Uredinales on Bignoniaceae from Brazil Joe F. Hennerand Helen M.P. Sotao 173 A new combination in“ihevetia (Apocynaceae) Justin Kirk Williams 185 ; CONTRIBUTIONS Una nueva especie de Agave subgenero Agave (Avavaceae) dé Mexico TO BOTANY José A. Villarreal Q. 191 The Mexican genera of the Apocyma@eae (sensu A. DC), with/key and additional taxonomic notes Justin Kirk Williams 197 VOLUME 17 NUMBER 1 SEPTEMBER, 1996 SIDA CONTRIBUTIONS TO BOTANY POUNDED BY LLOYD H. SHINNERS 1962 Win. F. Mahler Publisher 1971-1992 Director Emeritus SH. Sohmer Director Barney L. Lipscomb Editor Botanical Research Institute of Texas, Inc. 509 Pecan Street Fort Worth, Texas 70102-4060, USA 817 432-4441 / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/sida John W. Thieret Prof. Dr. Felix Llamas Associate Editor Contributing Spanish Editor Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia Northern Kentucky University Universidad de Leon Highland Heights, Kentucky 41076, USA E-24071 Leon, Spain Guidelines for contributors are available upon request and on the inside back cover of the last issue of each volume subscription per year: $25. Individual, $35. USA Institutions, $45. Outside USA numbers issued twice a vear “SIDA, CONTRIBUTIONS TO BOTANY, Volume 17, Number L. pages 1-303, Copyright 1996 Botanical Research Institute of Texas, Inc. Printed in the United States of America | ; a SSN OOS6- 148 5 ETHNOBOTANY SERVING SOCIETY: A CASE STUDY FROM THE SIERRA DE MANANTLAN BIOSPHERE RESERVE BRUCE F. BENZ, JUDITH CEVALLOS E., ELIZABETH MUNOZ M., and FRANCISCO SANTANA M. Instituto ‘Manantldn’ de Ecologia y Conservaciin de la Biodiversidad Centro Universitario de la Costa Sur Universidad de Guadalajara A.P. 64 Autldn, Jalisco C.P. 48900 MEXICO ABSTRACT The move to place biological diversity and traditional cultural knowledge into the main- stream global economy is based on the premise that economic subsidies will stimulate conservation. This premise is arguably counterproductive if the effects of such subsidies are not felt at the local level. Ethnobotanical research should focus on characterizing tradi- tional knowledge to establish eae with the local community to ensure that local values are translated into rational use of resources and effective conservation of biological diversity and cultural eee nv case study focussing on twenty-five of the most lo- cally valued plants from a community in the Sierra de Manantlan Biosphere Reserve pro- vides an example of recommendations that can be made based on basic ethnobotanical inventories. RESUMEN La tendencia a ubicar la diversidad biolégica y el conocimiento tradicional dentro de la corriente de la economia global, se fundamenta en la premisa de gue las subvenciones estimularian la conservacién. Esta premisa resultarfa contraproducente si los efectos de tales subvenciones no llegaran al nivel local. La investigaci6n etnobotanica deberfa enfocarse en el uso del conocimiento tradicional para establecer las prioridades conjuntamente con la comunidad local, para asegurar que los valores locales se traduzcan en el uso racional de los recursos naturales, una conservaciOn efectiva de la biodiversidad y el conocimiento cul- tural. Un estudio de un caso en una comunidad de la Reserva de la Biosfera Sierra de Manantlan, ee examina veinticinco de las especies vegetales mayor valor local, proporciona de las recomendaciones que pueden rey en ae a la informacié6n de inventarios ease ee. Ethnobotany has arrived at center stage in the world of biological con- servation and sustainable development (Alcorn 1995). The research efforts of ethnobotanists who document and analyze how people and cultures use and classify plants is now much more than an academic exercise, it has become a focal point for biological conservation (Wilson 1992) and sustainable S1pa 17(1): 1-16. 1996 2 Sipa 17(1) development (Reid et al. 1993). Chemical ecologists and natural product chemists have renewed interest in the research ethnobotanists and anthro- pologists conduct among traditional peoples from remote corners of the world. Renewed interest in ethnobotany stems from a new applied research field known as chemical or biodiversity prospecting (Eisner 1990; Eisner & Beiring 1994; Reid et al. 1993; see also Balick 1994; Cox 1994; Farnsworth 1994). Prospecting refers to the identification and use of previously un- studied genes and the primary and secondary metabolites of organisms that can remedy existing human illness or improve well-being by reducing loss due to crop pests, or by simply increasing the yield of our few cultivated crops. Entrepreneurs hope that placing biodiversity and diverse cultural knowledge into the mainstream global economy, nature’s products will fa- cilitate treatment of the many ills that befall society today and at the same time will help us to avert the human induced surge of biological extinction. The biological conservation agenda (Wilson 1992) sees the importance of biological diversity from this same vantage pointing out that the utili- tarian or economic arguement is the strongest in favor of conservation. The valuation of biological diversity will help to avoid extinction which re- duces our ability to confront existing and future social costs such as pro- ducing enough food, and curing existing and unforeseen disease. There are myriad other arguments that form part of this agenda that hinge on the indirect values of biological diversity or ethical issues such as the right of one species to extinguish another. However, the value of biological diver- sity has become a central issue among conservationists. And many devel- opment efforts today are investing in biological diversity banking on the possibility that this vast store of natural products might yield the gene sequence or new chemical compounds that will revolutionize plant indus- try or health care possibilities. Many conservationists believe that the value of such discoveries can be translated into far-reaching conservation efforts. We believe they can. However, the economic arguement has significant consequences both for echnoborany as a discipline and ethnobotany’s long- term integration with biological conservation. Placing ethnobotany and ethnobotanists in this situation may have cre- ated a gold rush mentality that is justified because extinction rates are higher than ever before and are increasing most dramatically where bio- logical diversity is greatest. We hope that is not simply an accommodation to global economy that will subject the discipline to the ultimate risk of any fad, viz. waning long-term interest. If we cannot save it, perhaps we can at least model the chemical structure of its secondary metabolites and gain insights that will improve our survival possibilities. The risks of los- ing future options makes ethnobotanical research worth the investment (Martin 1994; King & Tempesta 1994). BENZ ET AL., Ethnobotany serving society =) An equally important justification for ethnobotanical research is no less academic than chemical prospecting but is complementary and practical. Ethnobotanical research should not focus solely on the big ticket items but also on a more general knowledge of local plant use because such informa- tion is relevant in regional land use planning frameworks to ensure rational use of natural resources at the local level and provide the basis for sustain- able conservation programs. Efforts to conserve biological diversity are con- tingent upon the willingness and capacity of the local human population. Willingness is determined by the value the biota has to the local commu- nity. Capacity is determined by the local communities’ ability to translate this value into realistic economic terms. Loss of traditional knowledge about utility of the local biota hinders conservation efforts because the local popu- lation may no longer consider the biota or its utility valuable or lack the ecological capacity to translate it into relevant social and economic terms. Ethnobotany should be conducted within this conservation-development framework because effective conservation of biological diversity depends on the individual or community’s ability to apply their own culturally rel- evant cost-benefit analysis of conserving or extinguishing the local biota in attempting to resolve local community problems. The best justification for conducting ethnobotanical research is simply that such research 1s one way of assuring that arguments in favor of conserving local biota are based upon local values. Here we provide one example of how ethnobotanical research can be used in integrating conservation and development. This example provides a short list of plant species with local value and recommendations of how such uses might be integrated into a community forestry project in a Bio- sphere Reserve in western Mexico. The Sierra de Manantlaén Biosphere Reserve (SMRB) is an integrated conservation and rural development project in southern Jalisco and Colima, Mexico (Graf et al. 1995; Jardel 1992). The SMBR developed out of the discovery of a wild perennial relative of corn (maize), Zea diploperennis II\tis, Doebley and Guzman. This species was discovered in a very restricted area of agricultural lands in the community of Ayotitlan in Manantlan (Benz et al. 1990; Iltis et al. 1978). The combination of the potential economic value of this endemic species, the extremely rich biota of Manantlan (JJardel 1992) and the social and economic problems of the rural populace led to the creation of a research institute by the University of Guadalajara de- voted to the administration of the Reserve in 1985 and the establishment of the Reserve by federal decree in 1987. This biosphere reserve extends across the community-held lands of twenty-eight ezdos or indigenous communities and the private properties of seventy families. Creation of this reserve did not change land tenure, but —. 4 Stipa 17(1) it did place limits on land use in certain areas. The reserve’s buffer zone has guidelines for natural resource use while natural resource use is prohibited in the core zone. The area’s biological diversity is especially significant with more than 2770 vascular plant species and 560 species of vertebrate fauna occurring in at least nine distinct types of vegetation (Vazquez et al. 1995). Agricul- ture and animal husbandry have been the dominant forms of livelihood in spite of the fact that the area is dominated by forest ecosystems. During the last century, utilization of Manantlan’s forests has increased in importance. Unfortunately, the area’s rural communities did not receive the economic benefits that resulted from forest exploitation during the last fifty years, because the concessionaires frequently avoided paying the appropriate tax, or the community as a whole was not the recipient of such a tax VJardel 1992). There can be little doubt that the area’s principal vocation is forest use. From a conservation and development standpoint it should be stressed that use of the forests must be primarily for the benefit of the local popula- tion which must counterbalance use with maintenance of existing biologi- cal diversity. At present only a single community in the Sierra de Manantlan Bio- sphere Reserve (SMBR) holds a permit for commercial timber extraction. Most community lands in the Reserve are subject to harvesting of forest products by the local population because these activities are alleged to have less impact upon the forests than commercial timber harvesting and they are not regulated by existing legislation. The single community holding a commercial permit, El Terrero, is an ejido of ca. 3000 hectares with a popu- lation of 360 which operates its own saw mill. The ejido of El Terrero is located on top (between ca. 2000 and 2500 meters above sea level) of the Cretaceous-age massif known locally as Cerro Grande located at the east end of the Reserve. Forests are composed of mixtures of pines and oaks while the highest altitude peaks are forested by fir (Abies spp.) and cypress (Cupressus lusitanica Mill.) and the protected drainages are blanketed by diverse and relictual stands of Montane Meso- phytic Forest (Rzedowski 1988). A management plan for the e77do’s forest was drafted by the University of Guadalajara’s Instituto ‘Manantlan’ de Ecologfa y Conservacién de la Biodiversidad (IMECBIO) under agreement with the community (Jardel et al. 1995). This management plan and accessory environmental impact statement are a prerequisite for commercial logging. This plan establishes the parameters of cutting, monitoring actions and mitigation alternatives in the event that exploitation has adverse effects on existing natural or cultural heritage. One of the appendices to the management plan included an ethnobotanical evaluation of the area’s flora and vegetation. This techni- BENZ ET AL., Ethnobotany serving society 5 cal document, which is the basis for this essay, was made available to the ejido assembly to complement the economic and ecological plans for har- vesting timber from the ej/do’s forests. The focus of the aforementioned appendix was to recommend forest product use from the results of ethno- botanical research that might satisfy some of the ejédo’s immediate needs. Readers interested in the specifics of timber harvesting are referred to the original management document (see Jardel et al. 1995). ETHNOBOTANICAL RESEARCH GOALS AND METHODS Ethnobotanical inventory in the Sierra de Manantlan was undertaken to assess the variation in cultural knowledge about the area’s flora, determine the effects of current utilization on the area’s flora, and to systematically record ideas, species and traditional management techniques that could suggest land use alternatives that would ensure a more rational use of re- sources (Benz et al. 1994, Jardel 1992:257-265). The needs of each local community were given first priority based on the premise that this focus would gain community support for any subsequent proposed actions that might be needed to mitigate conservation problems. Hence species occur- ring in the immediate vicinity of the communities - those occurring within a two kilometer walking distance - that were identifiable (i.e. had flowers or fruits), were used in interviews during our short but seasonal visits. The general inventory was concerned with recording and analyzing the diver- sity of ethnobotanical knowledge using a systematic method of data collec- tion (details of collection, interviewing, and data management are pro- vided in Benz et al. 1994). Persons providing information were informed of the goals of this project and of its relevance to the implementation of the Reserve's management plan. While not all informants were in complete agreement with the Reserve's objectives (principally because of potential conflicts over land tenure), all who agreed to collaborate provided information on plant use knowing that it was being collected to assure its conservation and would be made available to them upon request. Our own interest in the system- atic collection of this information included securing alternatives that might assist in resolving existing and future needs of the community, as well as for seeking solutions to the needs of other communities in the Reserve. We felt justified in considering alternatives for all communities because of the need to conserve the biota in the Reserve as a whole, not just that in any one community even though information discussed here was elicited from El Terrero informants using species that occur in close proximity to the community. Of the 517 plus species occurring in the forests of El Terrero, at least 152 of the 253 species used in informant interviews have been designated — 6 Sipa 17(1) as useful. A total of 43 informants (17 males, 46.3 + 14.4 years old, range 21 to 72 and 26 females, 37.3 + 17.5 years old, range 12 to 88) were interviewed over a four year period. On average each provided 8.4 + 10.6 reports of use for all of the species used in interviews (see Benz et al. 1994). Each species was recorded as useful by an average of 5.3 + 4.9 informants. Eleven species were recognized as useful by more than 15 informants and thirty-six species reported as useful only once and by a single informant. Importance values were calculated to provide each species with a relative measure of local use value based upon the assumption that the total num- ber of reports of use, the sum of the different types of use and the total number of informants agreeing that a species is useful provide appropriate measures of a species’ usefulness to the community (Benz et al. 1994; Phillips & Gentry 1993; Phillips et al. 1994; interested readers should consult the original articles for details on calculating these indices). These importance values have been utilized as a means for recommending species based on the belief that a higher value indicates the species has greater relative im- portance to community members and thus has greater potential for resolv- ing community needs. Relevant information of a species’ utility also comes from outside the com- munity. A local herbal remedy (or other specific use for a species) that is used throughout the country (hence our frequent citation of Mexico’s Instituto Nacional Indigenista publication on medicinal plants {INI 1995 }) leads us to suggest that potentially greater economic benefit might be available to the community if greater investment was made to increase production and/ or distribute the product more widely. Such occurrences need to anticipate demand and plan production to ensure the local resource is not over ex- ploited. In a similar fashion, when a particular species is locally preferred for a specific use, ¢.g., house construction, recommendations for its extraction or planting need to be contemplated in anticipation of community needs. Recommendations are offered for the edo of El Terrero in its entirety. The forest management plan proposes timber harvesting in only a fraction of the total edo area (Jardel et al. 1995). Because the habitat requirements of some plant species are found only in certain vegetation types and these will probably not be subject to timber exploitation, some recommenda- tions may seem out of place. It should be made clear that our recommenda- tions were offered for planning management of all the ey/do’s forests, not only those slated for timber harvest. Furthermore, the data on which these recommendations are based are derived from existing knowledge and pat- terns of resource use. Our recommendations attempt to integrate these tra- ditional patterns with recommended timber harvesting schedules. Recom- mendations for many species will have to be made compatible with the ejido’s forest management plan, restricting, for example, exploitation of BENZ ET AL., Ethnobotany serving society 7 species found only in Mesophytic Forest which is protected by standards outlined in the Reserve’s Management Plan. The forest management plan divides the ezdo’s 38 most common tree species into four management categories (Jardel et al. 1995). A single spe- cies of oak, Quercus crassipes Humb. & Bonpl. is included in Category I, the species considered most significant for timber production. One of the spe- cies included in Category 2 of the ejédo’s management plan and in our short list, Q. obtusata Humb. & Bonpl. (Table 1 and 2), is recommended for ex- ploitation, not for lumber, but for cellulose, firewood or charcoal. Three of the species listed in Table 2, Arbutus xalapensis McVaugh and Rosatti, Prunus serotina Ehrh. and Ternstroemia lineata DC. are included in Category 3 which are destined for producing firewood, charcoal, posts, cellulose and wood- working because the species occur on e7/do lands in low volumes. Other tree species considered below (Table 2) are placed in Category 4 of the ejdo’'s management plan. Exploitation of natural populations of these species 1s restricted because the species are rare, regeneration is low, or current legis- lation or reserve naa guidelines mandate special protection. The species listed in Tables 1 or 2 and included in this category are Q. laurina Humb. & Bonpl., Q. rugosa Nee, Alnus jorullensis H.B.K., Cercocarpus macrophyllus C. Schneid , Styrax ramirvezii Greenm., and Sy ee citrea Lex. pecies listed in Tables 1 and 2 and not mieneened above are not given additional management recommendations by Jardel et al. (1995). ETHNOBOTANICAL RECOMMENDATIONS FOR EL TERRERO It should be made clear that we have not yet validated the efficacy of the traditional remedies or determined the economic feasibility of marketing additional quantities. The former will require extensive toxicology and clini- cal evaluations which will be undertaken in the future. Marketing of plant products from the species described below has been a subsistence endeavor of the local people for many years. Unfortunately, harvest and sale of some may have given them an unfair market advantage that clandestine exploi- tation offers (i.e. no limits on harvest quantities, no payment of institu- tional overhead, sale at unofficial rates). Whether increased production of theses products through managed exploitation can be accommodated in existing markets will require a detailed market evaluation. However, inter- views with retail and wholesale vendors have established the existence of local, regional and national markets that are far from saturated with the products that might be available from El Terrero. Literature citations are provided to indicate that certain uses have persisted over time or that other markets may have existed in the past. Recommendations are offered to suggest that local processing of marketable plant products appears to be feasible because the market is already known to exist. 8 Sipa 17(1) Tasie |. Useful plane species from the ejido of El Terrero with values greater chan two standard deviation units from the mean for one or more indices of importance. Most species are included because their values on Phillips’ overall use value exceed 2s. The importance values listed include: number of Nae of use from all informants (Reports; see Benz et al. 1994), cotal number of infor- manes that identify the species as useful (Infos), che number of occasions when the species was pre- sented to an sie ant co clicit information (Events), the number of general categories the plant is used for (of thirteen possible), and use value and overall use value described in detail by Phillips and Gentry (1993:19) Taxon Reports Infos Events Uses Ise Overall Value Use Value Prunus serotina 73 21 24 69 OL.0 a9 Quercus laurina 18 19 2) 15 34.7 1.8 Symplocos citrea 39 14 18 37 28.5 2.0 tyrax ramirezil 39 17 24 38 26.2 1.5 Rubus adenotrichos 35 17 22 44 26.5 1.6 Ternstroemia lineata 42 12 14 26 23.5 2.0 Baccharis pteronioide: 30 19 19 27 27.0 1.4 Comarostaphylis discolor 30 12 16 30 21.5 1.8 -elastrus pringlei 28 18 23 28 22.) 1.2 Rubus pringlei 27 17 1s 24 23.0 1.4 rataegus pubescens 26 12 12 24 24.0 2.0 Smilax moranensis 25 16 20 24 19.5 Ld Quercus rugosa 22 9 10 22 20.0 2.2 Quercus obtusata 17 5 5) 13 13.0 2.6 Fragaria mexicana 16 16 16 16 16.0 1.0 Quercus scytophylla li 5 5 13 13.0 2.6 Cercocarpus mé le cal 11 5 5 10 10.0 2.0) Quercus Crassipes 6 5 3 6 6.0 2.0 Xylosma flexuosum* 4 2 2 { 1.0 2.0 Zancthoxylum arborescens* 2 | l 2 2.0 2.0 ‘ These species have been eliminated from Table 2 because they have importance values that al- rae significant appear to be an artificially high because the number of different uses is exactly twice the number of informants. The order of presentation of the species is based on Table 1, the order of local importance based on relative use values calculated from reports of use elicited from the inhabitants of El Terrero. Capulin, (Prunus serotina Ehrh.) is a widespread but locally rare tree occurring in the Mesophytic, Adzes-Pine-Oak and Pine-Oak Forests of the ejido of El Terrero. It is used for a wide variety of purposes by the local population (Tables | and 2). The fruit is highly appreciated for making fruit drinks and punches. The bark and less often the leaves and flowers are prized ingredients for preparing cough remedies, and treating abdominal pain and diarrhea among other ailments (INI 1995; Hill 1952). The wood has been worked into fine furniture and cabinetry in the pase (Hill 1952) but because of limited availability it has fallen into disuse. The bark and leaves from felled trees should be collected and processed for sale. The wood of this species was previously used for furniture and BENZ ET AL., Ethnobotany serving society 9 rE 2. Twenty-seven plant species recommended for incorporation in forest exploitation program BI for ‘ie ejido of El Terrero based upon importance values, their potential in ef/do forestry programs and the existence of a local market for some or all of a species’ products. The uses listed are arranged in ited order of the frequency in which they were elicitec Taxon Local Uses Prunus serotina Edible, medicinal, construction, firewood, furniture, forage, fence posts, instruments Quercus laurina Construction, firewood, instruments, furniture, forage, medicinal, ed- Symplocos citrea Edible, instruments, construction, forage, fence posts, furniture Styrax ramirezil Forage, construction, edible, firewood, instruments Rubus adenotrichos Edible, medicinal, forage Ternstroemia lineata Medicinal, construction, firewood, fence posts, instruments s pteronioides Instruments, medicinal, forage, edible Com: ae discolor — Edible, firewood, fence posts, pene furniture velastrus pringlei Forage, edible Rubus pringle Edible, medicinal, forage Crataegus pubescens Edible, medicinal, forage Smilax moranensis Edible, forage, ins n um Edible, firewood, instruments, forage, construction Construction, firewood, for: ig & fence posts, furniture Edible Construction, edible, forage, medicinal, furniture, firewood aay firewood, construction, fence posts, forage, ea il ble fedi cinal, instruments, firewood, furniture, forage, edi Sambucus mexicana Medicinal Lepechinia caulescens Medicinal Buddleia sessiliflora Medicinal Gnaphalium americanum — Medic Myrcianthes fragans Edible, medicinal, construction cabinetry and also as support bases for scientific instruments, for making patterns and turnery (Hill 1952:97) because of the beauty of its grain and the ease with which it is worked. While many of the uses of this wood are no longer competitive, the beauty and ease of working will always be ap- preciated. Local use in furniture manufacture would appear to be a promis- ing enterprise. Anticipating relatively low quantities in existing forests leads one to suggest that this species be incorporated into nurseries. The encino chilillo (Quercus laurina) is one of the oak species listed in Category 4 of the ej7do’s management plan that show potential but whose merits have not yet been evaluated. The species occurs principally but not exclusively in Mesophytic Forest. Apart from the wood being highly prized for construction timber, firewood, tools, furniture, and fence posts, its acorns are a preferred fodder and acceptable foodstuff, and its bark is widely ap- preciated in Mexico for treating tooth-aches, diarrhea and other less com- monly cited maladies (INI 1995). Exploitation of this species should consider harvesting the bark as soon as 10 Sipa 17(1) the tree is felled. It should be dried in the shade if sold for commercial purposes. Symplocos citrea Lex., or Chico, is a medium to large tree occurring most frequently in Mesophytic Forest. The most frequently cited use by the local inhabitants is for its edible fruits, consumed both by humans anc on domestic animals. Other reports refer to use of the wood which is suitable for tool handles, construction timbers and fence posts. No reference is made to its use as firewood. Felled timber might be used first to satisfy domestic construction and fencing needs. Consumption of the fruit is widely acknowledged suggest- ing that nutritional evaluation would be useful to determine its potential as a foodstuff. Fruit-harvesting does not appear to be intense enough to Warrant concern or regulation, Palo de Casa (Styrax ramirezii Greenm.) is a small understory tree found principally in Mesophytic Forest. Leaves and small branches are used as fodder for domestic animals and the stout branches and trunk are preferred for use in house construction, hence its common name. Populations of the species might need to be monitored if house construction increases. Two species of blackberry (zarzamora, Rubus adenotrichos Schlecht.and R. pringlei Rydb.) are actively collected in El Terrero. Both species occur in Mesophytic Forest while the latter also appears as an element of Oak Forest. The acid fruits are used in fruit drinks and liqueurs and the leaves, branches and roots are prepared to treat diarrhea (INI 1995). Both species are colonizers of forest openings and aggressively hold ter- ritory once established. Openings in Oak forest created by timber harvest- ing might be planted initially with blackberry while regeneration is un- on derway. Cultivated populations would probably have to be managed under such conditions to ensure that pine or oak seedlings are not affected. Col- lection and sale of fruits from natural populations is a very remunerative activity that would be greatly enhanced through cultivation. Harvest of leaves or the roots for sale as ingredients in traditional remedies would be possible once forest regeneration had begun to limit fruit production. Tila or Jazmincillo (Ternstroemia lineata DC.) is a common under- story tree of the Oak and Mesophytic Forests on eido land. The most com- mon use of the species is as a medicinal remedy (INI 1995) although its wood is also used for construction timber, firewood, fence posts and tool handles. Fruits and flowers (actually the sepals and pistil that persist after the corolla has fallen) are collected during the dry season from trees in forest clearings, along roadsides and in house gardens, dried, and sold. Al- though collection may involve all members of the family, women and chil- dren devote substantially more time to all aspects of commercialization than men. A large national market exists for the flowers and fruits of tila. Tila is a shade tolerant species that is widespread in the forests of the BENZ ET AL., Ethnobotany serving society 1] ejido. Exploitation of the wood is contemplated in the ejédo’s management plan. The wood is reported to be useful for construction and firewood, less so for fence posts or tool handles. Cultivation of the species for its wood does not appear to be a priority (Jardel et al. 1995) due probably to its ecological requirements and a lack of known marketability. Integrating the species into management of the forests should be sought knowing that a national market exists for the fruits and flowers, and that members of the community already devote considerable time to collection and commer- cialization. Integration may be difficult because tila is a late successional species and does not produce well under such natural conditions. One al- ternative might be to plant it in openings along logging roads and along the margins of clearcuts. Cultivation in such open areas might permit greater fruit production earlier than normal allowing flower and fruit collection during periods when forests in adjacent areas are regenerating. A second alternative might be to experiment with mixed plantings of oak and tila thinning tila as the oaks begin to dominate. Baccharis pteronioides DC. (cuaracata) is an understory shrub of Adzes Forest. It is locally prized for making brooms and for use in medicinal remedies. Domestic needs appear to be satisfied without undue pressure on local populations. Insufficient information is available to determine whether the species might be cultivated for medicinal purposes. Madrofillo (Comarostaphylis discolor (Hook.) Diggs), is an under- story shrub that occurs in Mesophytic Forest. The fruit is edible and the wood is commonly used for firewood and fenceposts. Fruit palatability ap- pears to be quite variable suggesting that nutritional profile and palatabil- ity studies be made if the species is considered a candidate for managed production. Celastrus pringlei Rose (bejuquillo relumbroso) is a canopy liana oc- curring principally in Mesophytic Forest. It often appears in clearings and along roadsides throughout the e7do as well. It is one of the locally pre- ferred forage plants and the aril is also edible. The bark is infrequently reported to have medicinal properties. The importance of the plant as a forage is widely recognized suggesting that when stands destined for logging that contain the species are located, harvest of the foliage should take place prior to logging to ensure that the felling of large canopy trees acting as supports for this and other lianas (e.g. see Smilax moranensis below) will not disturb subcanopy trees. Such harvest- ing could satisfy some of the forage needs of families that possess livestock. The edible fruits of teyjocote (Crategus pubescens (H.B.K.) Steud, also known locally as manzanillo) are widely appreciated in Mexico as an ingre- dient in holiday punches. The fruits are widely sold in markets though price differentials for larger cultivated varieties and wild collected fruit are 12 Sida 17(1) not apparent. This species occurs in disturbed areas, as well as in Oak and Mesophytic Forests. The roots of tejocote are also very widely harvested for medicinal purposes for a vast array of illnesses and extracts are known to have cardiotonic activity (INI 1994), This species could be cultivated more widely in the ejido especially in abandoned mé/pas and pastures. It has a significant though seasonal local market that is otherwise satisfied with fruit transported from regional mar- ket centers. Local focus on extraction of roots for medicinal purposes may have had a negative impact on existing populations. Determination of the efficacy of extracts from other parts of the plant could be an especially timely undertaking to anticipate and understand the impacts of root harvesting. Asierrilla (Smilax moranensis Martens & Galeotii) is a canopy liana apparently restricted to Mesophytic forest. It is highly esteemed locally as an edible green (Cevallos 1992). Medicinal use of the roots is well-known in other parts of Mexico (INI 1994) as is that of tropical relatives (Arvigo & Balick 1993) If restricted to Mesophytic Forest, it is unlikely chat this species will be encountered in forests destined for timber harvest. Cultivation of the spe cies in clearings and house gardens is suggested should collection of local stands ever intensify. The root of the species is commercialized throughout Mexico. Harvest of populations existing in logging tracts should be a in anticipation of cutting for reasons already mentioned in the case of beju- quillo. Market evaluation should be undertaken if pharmacological activ- ity shows promise. The species is extracted from natural stands in other parts of Mexico suggesting that protection may be necessary in the future. Two additional species of oak (avellano, Quercus rugosa Nee and roble negro, Q. obtusata Humb. & Bonpl.) are widely recognized as useful by the people of El Terrero. Jardel et al. (1995) report both from a variety of forest types in addition to Mesophytic Forest. Avellano (roblillo in Jardel et al. 1995) is most frequently reported for its edible acorns and less fre- quently for the multiple uses of its wood while roble negro is more often identified for the usefulness of its wood or the medicinal value of its bark (INI 1995). The latcer species is included in Category 2 in the ejdo’s man- agement plan and will probably be exploited commercially. Harvest and commercialization of the bark of Q. rwgosa should be con- sidered if the species is incorporated into timber harvesting cycles. Use of oak barrels for aging tequila or mescal might be pondered as a viable mar- ket for the wood of these oaks. The face that Q. rwgosa has edible acorns implies that acorn production of this species and perhaps other oak species might be systematically harvested by livestock (cf. Peattie 1950:210—211). Fresa cimarrona (Fragaria mexicana Schlect.) is an herb found princi- pally in Adves or Pine-Oak Forests. Its edible fruit are widely appreciated — BENZ ET AL., Ethnobotany serving society 13 but the plant does not produce sufficient quantities nor large enough fruits to merit large-scale sytematic harvesting nor commercial production. The populations found in El Terrero may be useful as a source of germ plasm. Roble negro (Quercus scytophylla Liebm.) is a canopy tree that occurs principally in Mesophytic Forest. The utility of its wood is well-known though it is just as likely to be mentioned as a source of edible fruit or for its medicinal bark. The same recommendations made for Q. rugosa or Q. obtusata can be made for Q. scytophy/la though its distribution in Meso- phytic Forest will probably restrict its exploitation. Aile (Alnus jorullensis H.B.K. subsp. jorullensis) is a medium-sized tree occurring in Adies, Pine-Oak Forests and Oak Forests. It is prized lo- cally for tool handles fabricated from its wood as well as for its use as a construction material and for fuel. Leaves and wood are used medicinally in Puebla and Sonora (INI 1995). It is reported least frequently as a forage in El Terrero. The species might be useful in restoring degraded areas as a forage be- cause it is known to grow with associated nitrogen fixing organisms (Langenheim & Thimann 1982; Noggle & Fritz 1976). Madrofio (Arbutus xalapensis McVaugh & Rosatti) is a locally rare but widely occurring medium-sized tree found in most all forest types in the ejido. It is recognized locally for the medicinal properties of its bark. A related species is used medicinally in the state of Baja California Sur. The wood is very hard and has recognized potential for tool handles, furniture or firewood. This species is included in Category 3 of the ezdo’s management plan, hence exploitation of its wood is anticipated. Local use of the bark for me- dicinal purposes might signal economic potential. —* Five of the six species added to Table 2 are discussed here because most have known market value and are commercially exploited for the national medicinal plant market (INI 1995) even though their use value indices are not significantly greater than the norm. Local vendors in Jalisco and Colima obtain these plants from wholesale markets (Los Abastos, in Guadalajara) or from local collectors. All five species occur in forest openings or dis- turbed areas. Three of the five (Marrubinm vulgare L. » Lepechinia caulescens (Fernald) Epling, Buddleia sessiliflora H.B.K.) are aggressive perennial weeds and could be cultivated in forest clearings prior to or during replanting and regeneration and harvested very intensely once the regenerating tree species establish themselves. The last species in Table 2, (Myrcianthes fragrans (Sw.) McVaugh) is a small to medium-sized tree that occurs in Mesophytic Forest. The fruits are consumed fresh or used to make fruit drinks. The species is highly esteemed as a flavoring agent and medicinal species and shows great potential for commercial use in herbal teas. Consideration might 14 Stipa 17(1) be given to intensifying fruit production in areas that are opened up through clear-cutting by planting the understory after the forests have regenerated. It should be noted that many of the species listed in Table 2 that have economic value individually, could be combined and commercialized as herbal teas or as traditional remedies. For example, Sauco (Sambucus mexicana Presl.), ts a widely sold medicinal species that has been combined with other species, e.g. Quercus rugosa and Crataegus pubescens as a cough remedy (cf. INI 1995). Other combinations from the list of 152 species are possible but should be evaluated to determine whether a market exists. CONCLUDING REMARKS Recommendations offered to the edo of El Terrero focus on a few se- lected products available from local forests whose importance is already recognized by community members. Our suggestions attempt to integrate existing patterns of use with a more focussed and systematic commercial- ization of timber and nontimber forest products into the ej/do’s forest man- agement plan. Benefits derived from these recommendations will be avail- able first to members of the community; thus investments made by the ejido itself to evaluate the potential of certain resources in greater detail, e.g. nutritional quality or pharmaceutical activity of certain species, will confer benefits over the long-term eee to the community. Decisions to implement these recommendations will be a function of the e7do's own understanding of potential costs and ae The ethnobotanist’s input in helping the ej/do to understand these costs and benefits is crucial because both social and economic impacts of implementing these recommenda- tions require both the perspective of the community and an independent ecological and economic assessment on the part of the researcher. Loss of biodiversity through inappropriate use of natural resources might not be readily apparent to community members but its impact will be felt glo- bally. Recommendations for use of forest resources need to be evaluated periodically to ensure that benefits to community and environment out- weigh their costs. Ethnobotanical work in the ejido of El Terrero is only beginning. We now need to follow up on the community’s evaluation and implementation of our recommendations and those made on the remain- ing 127 species of plants that the community recognizes as useful. ACKNOWLEDGMENTS We are grateful to the numerous individuals from the ejdo of El Terrero who provided valuable personal knowledge about the local flora. We would like to dedicate this article to the memory of Fructuoso Ayala who devoted his energies toward the conservation and sustainable use of the ejido’s for- ests. We thank A. de Avila B, P. Gerritsen and G. Martin who provided BENZ ET AL., Ethnobotany serving society 15 valuable comments on an earlier version. This research was supported by the Universidad de Guadalajara, Mexican Secretario de Educacién Publica, the Biodiversity Support Program and Earthwatch Volunteers. REFERENCES ALCORN , J.B. 1995. Economic ioe. conservation, and development: What's the connec- . Ann. Missouri Bot. Gard. 82:34—46. Reais, - a. a Batick. 1993, eee remedies. Lotus Press, Twin Lakes, Wisconsin. Batic rere ieee drug development and biodiversity conservation—ex- cee eee In: D.J. Chadwick and J. Marsh, eds. Ethnobotany and the search Hot new drugs. Ciba Foundation Symposium 185. Joh iley & Sons, New Yor 4 Benz, B.F., L.R. SANCHEZ-VELASQUEZ, and F. re MicHeL. 1990. i aad: ethno- botany of Zea diploperennis: Pile investigations. Maydica 35:85— Benz, B.F., F SANTANA MicHeL, M. R. Pinepa-Lopez, J. Cevaos E., L. ee H. and D. DeNiz L. 1994. C haracterizacion of Mestizo plant use in the Sierra de Manantlan, Jalisco- Colima, Mexico. J. Ethnobiol. 14:23-41 Cox, P.A. 1994. a and drug development. In: D.J. Chadwick and - Marsh, eds. Ethnobotany and the search for new drugs. Ciba Foundation Symposiu 185. John Wiley & Sons, New York. Pp. 25—41. Ceva.tos Espinoza, J. 1992. El papel de los quelites en las comunidades de la Reserva de la Biosfera Sierra de Manantlan. Tesis de Licenciatura, Faculrad de Ciencias, Universidad de Guadalajara. EIsNER, T. 1990. Prospecting for nature’s chemicals. Issues Sci. Tech. 6:31—34. Eisner, T. and E.A. BEIrING. 1994. Biotic ew Mg fund—Protecting biodiversity through chemical prospecting. Bioscience 44:95— FarnswortH, N.R. 1990. The role of ethnoph pee er in drug development. In: D.J. Chadwick ana i Marsh, eds. Bioactive sn a from plants. Ciba Foundation Sym- posium 154. John Wiley & Sons, New York. Pp. 2-21. Farnswortn, N.R. 1994. Echnopharmacology and on development. In: D.J. Chadwick and J. Marsh, eds. Sneed: and the search for new drugs. Ciba Foundation Sympo- sium 185. John Wiley & Sons, New York. Pp. 42-59. Grat M.S., E. SANTANA ee E. Jarper PeiArz, and B.F. BENz. 1995. La Reserva de la Biosfera Sierra de Manantlan: un balance de ocho afios de gestién. Rev. Univ. Guadalajara. March-April: 55-60 INI (Instituto NACIONAL INDIGENISTA). 1994. Atlas de las plantas de la medicina tradcional Mexicana. Vols. I, II and III. Insituto Nacional Indigenista, Mexico, D.F. Jarpet P., E. (Coord). 1992. Estrategia para la conservacion de la ae de la Biosfera one ts Manantlan. Universidad de Guadalajara, Guadalajara, Mexico. JarpeL P., E. ec alia 1995. Programa de manejo forestal del ejido El Terrero, Mpio. ee Colima. Informe Técnico, IMECBIO, Universidad - Guadalajara. Hu, A.E. 1952. Economic botany. McGraw-Hill Book Company, Inc., New York. Intis, H.H., J. Dorstey, R. Guzman, and B. Pazy. 1979. Zea diploperennis (Gramineae): a new teosinte from Mexico. Science 203:186-188. Kino, S.R. and M.S. Temprsta. 1994. From shaman to human clinical trials: the role of eae in rer ytany, conservation and community reciprocity. In: D.J. Chadwick . Marsh, eds. Ethnobotany and the search for new drugs. Ciba Foundation Sympo- um 185. John Wiley = Sons, New York. Pp. 197-2 ican J.H. and K.V. THimann. 1982. poe Plane biology and its relation to human affairs. John eee & Sons, fie. New Yor 16 Stipa 17(1) Martin, G.J. 1994. Conservation and ethnobotanical exploration. In: D.J. Chadwick and Marsh, eds. Ethnobotany and the ic. for new drugs. Ciba Foundation Symposium 185. John Wiley & Sons, New York. 229-239 Reib, W.V.,S.A. Lainb, C.A. Meurr, R. ee A. SITTENFELD, D.H. JANZEN, M.A. GOLLIN, and C. Juma. Biodiversity prospecting: Using genetic resources for sustainable develop- ment. World eee oe Washington. Noccie, G.R. and G.J. Frrrz 1976. Introductory plant physiology. Prentice-Hall, Inc., br ed Cl . So _———— ae SS nN un 20 ee RRR SE Se SEE 4 Ee SSS ees ees a PRP = ea) peed eee. EE Seay Soe $B iis ue / if as 0 va eds = tee, See ore See no. of plants J 1 2 3 4 5 6 7 8 9 10 11 12 13 N transect number S Fic. 3. Distribution of black oak taxa on linear transects at Ferrerfa. With histogram bars representing (front to back) Q. edwardii, individuals of hybrid origin, and Q. conzattiz, re- nernmost and transect 13 is southernmost. Transects are — spectively. Transect | is nort oriented approximately E-W. —_— tically verifiable. Beyond the southernmost transect no Q. conzattii and only an isolated hybrid individual were present. On the transects we found 163 black oaks (Fig. 3). Putative F, and later generation individuals were distributed unevenly throughout the site ex- cept on the south end, where only Q. edwardii occurred. Q. conzatti is the most abundant black oak in the hybrid swarm area, represented by 82% (n = 133) of the individuals. Hybrids made up 14% of the population (” = 25), and, among these, only 4% (7 = 7) of the total number of black oaks sampled were probable F, plants (morphologically intermediate without apparent recombination of characters). The frequency of later generation hybrids (non-F,) was equal (5.5% each) for Q. conzattii-like hybrids (7 = 9) and Q. eduardii-like hybrids (7 = 9). Three percent (7 = 5) of the black oaks within the hybrid zone were considered relatively pure Q. eduardiz. The relatively small percentage of hybrid individuals recorded becomes more impressive when one considers the extended population of Q. eduardii to the south. This suggests that post-zygotic selection resulting from subtle differentiation of habitat is probably a key factor isolating these species, as Muller (1952) proposed in his evaluation of several hybridizing oak species pairs. The distribution of Q. edwardii almost exclusively at the south end of the site and the lack of Q. conzattii and hybrids beyond the southernmost transect indicates that such ecological isolation is occurring at Ferreria. 24 Stipa 17(1) Taste 1. Characters used in calculation of Anderson hybrid index values and descriptions of states considered “typical” for each taxon evaluated. Anderson hybrid index values for each taxon are indi- cated in parentheses below each taxon name. character Quercus conzattit hybrids Quercus eduardti (2) ) leaf size (length in mm) >98 58-98 <58 leaf shape (engch: width) < 2.1 2.1-3.1 23,1 rugosity of leaves veins strongly veins moderately veins not impressed impressec mpressed luster of adaxial leaf surface dull mod. lustrous lustrous pubescence type on woolly & dense moderately woolly — short & sparse adaxial surface & dense pubsecence type in no dense clusters lense clusters dense clusters leaf vein axils barely visible crete on 1 abaxis al present barely visible absent idert prominence of reticulation strongly prominent some reticulation — absent of leaf veins bullateness of abaxial epidermis — strongly bullate mod. bullate smooth SALIENT CHARACTERS OF PARENTS AND HYBRIDS In this section and following, specimens are cited by collection number, fully documented and explained in the appendix. Collection numbers are those of Bacon and associates unless otherwise indicated. Quercus conzattit:—Shrubs to trees; leaves thick and very stiff, to 14 cm long, strongly convex (often folding when pressed), broadly obovate to round; adaxial surface olive green, semi-lustrous to dull, nearly glabrous, rugose by impression of larger veins and veinlets; abaxial surface gray (in Durango), pilose, the pubescence of long-stipitate-fasciculate trichomes (terminology of Hardin 1979) with stipes 1—1.2 mm bearing 6—7 branches to 1 mm long, nearly or completely obscuring the strongly bullate, mi- nutely papillate epidermis, abaxial vein axils without marked differentia- tion of trichomes; acorns small, in groups of 1 or 2 and nearly sessile or, usually, up to 4—5 on short peduncles 1-1.5 cm long, maturing in late summer or early fall. General characters of the leaves and trichomes of Q. conzatti are evident in Figs. 2 and 4C, respectively, and qualitative and quanti- tative characters of plants examined are summarized in Tables 1 and 2 Quercus conzattit X Quercus eduardii F; and segregates:—Hybrid plants are intermediate between the parental species in leaf morphology (size, shape, texture, bullateness, convexity, adaxial luster, vein reticulation, BACON AND SPELLENBERG, Hybridization in Quercus Taste 2. Characters used in multivariate analyses. All variables were used in analyses involving the Full Data Set. Annotations: 1 = data used in Abbreviated Data Set; 2 = no significant difference in data used in Hair Data Full Data Set; 3 = no significant difference in Abbreviated Data Set; and 4 = Set (see text for explanation of data sets) A variable (units) Quercus conzatti1 hybrids mean (s.d.) Quercus eduardit mean (s.d.) mean (s.d. leaf length (mm)! a 3(12.3) 68.8(21.8) 35.4(6.3) maximum leaf width (mm)! (9.0) 37.6(15.9) 12.2(2.3) dist. from apex to pnt on midrib 3 . 3.4(9. 6) 32.5(10.1) 17.0(5.4) that eae to widest pnt on leaf (mm)! leaf Se at basal 1/3 of leaf(mm) 46.5(10.4) 32.7(13.9) 11.1(2.1) leaf width at apical 1/3 of leaf (mm) 50.4(7.4) 33.4(16.0) 10.6(2.2) no. veins per leaf? 17.3(2.1) 18.2.0) 16.2(2.9) no. teeth per 3 leaves 37.8(11.9) 28.2(14.2) 16.3(11.6) width of midvein adaxally (mm)! 2.0(0.6) 1.8(0.6) 1.2(0.3) width of midvein abaxally (mm)! 5.2(1. es 4.001 2 2.1(0.4) % coverage by pubescence of 98.4(2.7 88.8(15.8) 66.3(24.6) abaxial leaf surface! % coverage by pubescence of adaxial leaf surface!” 5.9(17.8) 2.8(33.6) 32.4(40.6) % coverage by pubescence of first 100(0) 100(0) 100(0) year stem maximum tooth ht. on leaf (mm)! 1.0(0.4) 0.8(0.4) 0.5(0.4) leaf apicule length (mm) 0.6(0.4) 0.5(0.4) 0.4(0.2) petiole length (mm) 5.4(2.8 4.2(1.7) 2.9(1.0) leaf surface rugosity (%) 18.8(19.0) 12.4(19.0) 4.1(15.8) twig di (m 2.2(0.41) 2.7(2.2 1.2(0.38) petiole diameter (mm) 5.7(1.86) 3.71.54) 2.5(0.58) cordateness of leaf base (mm) 2.2(1.41) 0.7(0.66) 0.2(0.28) vein angle at midleaf (degrees)? 137.7(8.8) 143.4(8.5) 127. 127.7) leaf apex angle (degrees PI2Z20 5, 79.6(17.3) 75.7(23.4) stipe length of trichomes of abaxial 0.096(0.016) 0.058(0.024) 0.005(0.008) leaf surface (mm)' maximum branch length of 0.029(0.007) 01(0.143) 0.64(0.248) trichomes of abaxial leaf surface (mm)* number of branches of trichomes 6.6(0.8) of abaxial leaf surface“ 9.8(4.0) 15.5(2.7) density of abaxial pubescence, and length of abaxial trichomes [Fig. 41), and other intermediate characters (see data analyses below). Later genera- tion segregates or backcrosses vary continuously from more or less exact intermediacy to one or the other parents (Fig. 2 and Tables 1 and 2). Hybrid individuals (see appendix) were found in nearly all locations where Q. conzattii and Q. eduardii intermixed (Fig. 1). Hybrids were also found in areas where both species were not intermediately intermixed, but where the second putative parent occurs within 0.5—2 km. Such was the case north of Boca del Mezquital, where both Q. conzattiz (1290) and 26 Sipa 17(1) Fic. 4, 4C. details of abaxial tric nome? ona leaf of Q. eduardii (Spellenberg 1097412); note that dense, tangled branches obscur e] the presence of a long stipe at top, right center. 4B. details of trichomes on a leaf of a hybrid with Q. edvardii-like morphol- rt =) ~ ogy (Spellenberg, et al. 10971—9); note partial visibility of epidermis and presence of s stipes (trichome at top center and at bottom right). 4A. ak of abaxial mee ona leaf of Q. conzattii (Spellenberg 1097312), note visibility of epidermis and sessile nature of rosulate trichomes. BACON AND SPELLENBERG, Hybridization in Quercus ay intermediate individuals (7293) occurred, and Q. eduardii (1048) occurred on nearby (1 km) hills and arroyos. Quercus eduardii:—Shrubs to trees; leaves to 5.2 cm long, thin and stiff, but flexible, flat, oblong, the adaxial surface dark green, lustrous, nearly glabrous; abaxial surface light green, sparsely pubescent, the pubescence of rosulate trichomes (terminology of Hardin 1979) with stipes 0.05—0.15 mm bearing 15—16 contorted or sinuous branches 0.32—0.34 mm long , not obscuring the smooth epidermis (Fig. 4A), the vein axils with clusters of dense, stout, conspicuously stipitate, sinuously-branched hairs; acorns small, in groups of 2—3 ona slender peduncle 0.5—2 mm long, maturing in mid- to late summer. Acorns and general leaf characters are visible in Fig. 2. ANALYSIS OF POPULATIONS OF PARENTAL SPECIES AND HYBRIDS We collected from sites where intergradation was obvious and from other sites where each of the parental species was away from genetic influence of the other. At four sites (exact localities and specimens cited in appendix and illustrated in Fig. 1) the following situations were found. Material rom 93 plants was collected, pressed, dried, and later measured: 1) O. conzattii and Q. eduardii hybridizing extensively south of Ferreria, represented by collection of: a) 18-intermediate specimens (1042-{1—18 (see Appendix for explana- tion of specimen numbering)}), b) 19-Q. eduardii specimens (1044-[1—19]), and c) 18-Q. conzattit specimens (1045-[1-18}); 2) Q. eduardii away from any other black oak at La Mona represented by collection of 11 specimens (1049-[1—11]), 3) Q. conzattii away from any other black oak (the nearest plant of Q. eduardi found more than 500 m from the population) east of the village of Boca del Mezquital represented by collection of 10 specimens (1046-{1—10], and 4) both parental species with very limited hybridization at Mezquital (at first we thought none; after several visits to the site a single, small interme- diate individual was discovered {Spellenberg & Bacon 11081] that was not used in analyses) represented by collection of nine Q. conzatti specimens (1047-[1—9}) and eight Q. eduardii specimens (1048-{1—8}). Further investigation revealed that hybridization between these species was widespread but sporadic in this region, but only at one site (Ferreria {1042}) did the situation represent a hybrid swarm with many individuals forming morphological continuum between parental species. Hybrid populations were documented at a number of other locations in southern Durango (/ 286, 1400, 1293, 1042, Spellenberg & Bacon 11681), and a single hybrid indi- vidual (1483) was found in Jalisco (see appendix). 28 Sipa 17(1) Vegetation at the sites varied from highly disturbed, arid savanna (lower elevational extreme of “bosque de coniferas y de Quercus” of Rzedowski 1978) at the hybrid swarm site near the agricultural village of Ferrerfa, to mature Quercus-Pinus-Arctostaphylos woodland at Mezquital. Vegetation at La Mona was Quwercus-Pinus forest. Vegetation at Boca del Mezquital was Quercus savanna, where Q. conzattii occurred high on the slope and Q. edvardii occurred more than 500 m away in the arroyos and on the bajadas below. Grazing and associated habitat disturbance probably occurred at all the sites, but was most likely much less on the very steep slopes of the Mezquital population. ANDERSON HYBRID INDEX Morphological intermediacy or a combination of suspected parental char- acter states are good indicators of hybridization in plants (Gottlieb 1972; Phipps 1984; Wilson 1992). Data were collected from specimens from the four sites previously mentioned to verify observations of intermediacy and inquire into the intermediacy of other morphological and micromorpho- logical characters. In populations where hybridization is evident, the Anderson Hybrid Index (Anderson 1949; 1953) is useful for identifying intermediate indi- viduals (Davis & Heywood 1963), showing backcrossing (or later genera- tion segregation), and showing the structure of a hybrid swarm (Wilson 1992). This technique is the most effective non-multivariate method for separating the parental taxa in a situation involving comparison of puta- tive hybrid taxa (Namkoong 1966; Wilson 1992), and the histogram it yields provides the advantage of an easily assimilated graphic representa- tion of the variation within each of the populations (Wilson 1992). For each plant specimen from the four study sites, an Anderson hybrid index value was calculated based upon nine characters that were consis- tently different between the two parental species (Table 1). These data con- sisted of measurements or subjective evaluations that placed each character into a ranking system (Anderson 1949, 1953), based upon its similarity of a single specimen to the character condition in the parental species. Char- acters representative of Q. conzatti received a rank of 2; characters represen- tative of Q. edwardii received a rank of 0; and characters that appeared inter- mediate between the two taxa received a rank of 1. Ranking method followed that of Wilson (1992), The Anderson hybrid index histogram for the Ferrerfa population re- veals a more or less continuous distribution of individuals between hybrid index values ranging from 0-17 (Fig. 5), suggesting that the black oak population there is the product of hybridization and subsequent backcross- ing involving Q. conzattii and Q. eduardii and/or later generation segrega- BACON AND SPELLENBERG, Hybridization in Quercus 29 12 Ferreria | Mezquital swarm 6 2 Cc } & a. Lom 0 4x = 1a S | Boca del Mezquital |; La Mona > | } c 6r L a: a: ial 0. 9 48 Anderson hybrid index value Fic. 5. Anderson hybrid index histograms showing frequencies of index values at four study sites in southern Durango. Plants of Q. edvardii have low values (ca. 0-4), and plants of Q. conzattii have high values (ca. 15-18). Intermediate values represent hybrids, back- crosses, or later generation segregates. tion. This range of variation is similar to variation in other oak hybrid swarms thought have structure which permits introgression (Stebbins et al. 1947; Cooperrider 1957; Silliman & Leisner 1958; Bensen et al. 1967; Maze 1968; Hardin 1975). No other black oaks are at this site; a white oak, QO. chihuahuensis is infrequent. The histogram for Mezquital reveals a similar range of values (1-18) but is bimodal, with perfectly intermediate plants missing excluding one weak putative F,; discovered too late to be included in same-year collections, as mentioned above). The population east of the village of Boca del Mezquital had values ranging only from 1 to 4, indicative of individuals exhibiting characteristics only of Q. conzattii. Plants from the La Mona site had values ranging from 15-18, values characteristic only of Q. eduardi1. To further examine variation in these populations, a multivariate analy- sis of variance was used to compare means between taxa within and be- tween sites. For each collection examined, initially a single measurement of each character was taken on the largest leaf (for number of teeth, three largest leaves were used). Twenty four characters were measured (Table 2). hese data are hereafter referred to as the Full Data Set. 30 Sipa 17(1) Based upon preliminary analysis of the Full Data Set, it was apparent that the taxonomic utility of some characters was greater than that of others. Consequently, 12 of the characters that demonstrated the greatest taxo- nomic value (annotated with a “1” after the name of the character in Table 2) were measured on up to 14 additional leaves on each specimen (fewer were measured on the few specimens with less than 14 intact leaves). These data were combined with data from the first series of measurements for a total of up to 15 repetitions on complete specimens. This data set in multivariate analyses (described below) was referred to as the Abbreviated Data Set. Dimensions of trichomes are variable but are commonly used as reliable indicators of taxonomic affinity of individuals. Consequently, data used to calculate means for trichome stipe length, branch length, and number of branches for the Full Data Set were used to examine micromorphometric differences between populations. This data set was referred to as the Hair Data Set. The character “pubescence on first year stems” was excluded from analy- ses because it did not vary significantly from 100% coverage of the stem surface. For the rest of the data, multivariate analysis of variance (MANOVA) (SAS GLM procedure, SAS Institute 1992) revealed significant differences (P < 0.05) between taxa within sites and among all sites for each of the three data sets for all characters except for vein number per leaf (P = 0.07) and vein angle at mid-leaf (P = 0.11) in the Full Data Set, and adaxial pubescence (P = 0.10) in the Abbreviated Data Set. Means for 20 of 21 (95%) characters (exhibiting significant differences between populations) were intermediate in plants of the hybrid swarm. The non-intermediate character “twig diameter” exhibited more variation in the hybrid swarm (mean = 26.9, sd = 22.2) than did either parental species population (Q. conzatti mean = 25.2, sd = 9.9; O. eduardii mean = 11.6, sd = 3.8), suggesting the formation of novel genotypes, a factor thought to be important in the development of new evolutionary trajecto- ries in hybridizing species pairs (Lewontin & Birch 1966; Arnold 1992: Grant, PR. & B.R. Grane 1992) or in permitting acceleration along exist- ing trajectories (Grant PR. & B.R. Grant 1994). CLUSTERING RELATIONSHIPS BETWEEN POPULATIONS Utilizing the same three data sets from MANOVA, cluster analysis (SAS CLUSTER procedure, SAS Institute 1992) was used to examine the clus- tering relationships among a priori recognized taxa within sites. Centroid distances (Dc) between groups within sites were calculated and are repre- sented in three cluster diagrams (Fig. 6). The cluster analyses were robust in joining the populations of Q. conzatti from east of the village of Boca del Mezquital and above Mezquital. This clustering relationship was weakest — BACON AND SPELLENBERG, Hybridization in Quercus pal Hair Data Set Abbreviated Data Set swe BdMc Full Data Set | BdMC Fic. 6. Cluster diagrams showing clustering relations of @ priori taxonomic populations at sites in southern Durango. BAMC = Boca del Mezquital _ population) (Q. conzattit), LME= La Mona (Q. eduardii), MC = Mezquital (Q. conzattiz), ME = Mezquital (Q. ea duardit), SWC = swarm site (Q. conzattii), SWE = swarm site (Q. ane Se swarm site (hybrids, backcrosses, and later generation segregates Sipa 17(1) eo No in the Hair Data Set (Dc = 0.05) and strongest in the Full Data Set (De = 0.82). This cluster did not group closely with the third Q. conzatti/ popula- tion, that of the hybrid swarm site south of Ferrerfa. Instead, Q. conzattii from here clustered more closely with the cluster of Q. edwardii from the site and intermediate plants (Hair Data Set Dc = 0.52, Abbreviated Data Set De = 0.37) or with the cluster of the Ferrerfa hybrid and other Q. eduardii populations (Dc = 0.35). To determine why the Q. conzatti/ population at Ferrerfa joined with the hybrid swarm population instead of with other Q. conzattii populations, populations of each taxon were compared to this Q. conzattii population by discriminant function analysis (SAS IML procedure, SAS Institute 1992). Five of the plants from this population were grouped with plants from the hybrid swarm (1042-41, -44, -49, -50, and -5/), accounting for the unex- pected clustering. The other 11 individuals grouped with individuals of Q. conzatti! populations, as expected. Two factors contributed to this unexpected result. First, a fire at Ferrerfa had set back growth of leaves on the site’s north end, where Q. conzattii collections were made, and collections included twigs with a late-season burst of growth (1042-41, -44). Immature leaves of Q. conzattii tend to be small, narrow, thin, less rugose, and sparsely poe abaxially, and im- mature twigs tend to be thinner. All of these characteristics are in the di- rection of Q. eduardi1. Secondly, of the remaining 16 plants identified as Q. conzatty at Ferreria three had small, narrow leaves and thin twigs (1042— 49, -50, -51), and in these respects were more like Q. edvardii. One of these had relatively thin pubescence on the abaxial leaf surface (1042—50), also more like Q. eduardii. Otherwise, Q. conzattii individuals at Ferrerfa grouped with Q. conzattii from other populations of that species. The analysis indi- cated that an @ priori classification was not completely correct, suggesting that these plants were of hybrid origin but were very close to Q. conzatti1. In the three data sets, the hybrid swarm population always joined most closely with one or more populations of Q. eduardii (De < 0.35 in all data sets), but the relationship between these populations was weak and vari- able between cluster diagrams. This suggests that hybrid plants sampled in this study are more like Q. eduardii than Q. conzattii. Principal components analysis (PCA) is useful for the identification of individuals within a population and placement of those individuals with- out their a priori identification. PCA (StatMost PCA, DataMost 1994, 1995) of the variance/covariance matrix was performed on the Full Data Set (Fig. 7), leaving out characters whose means were not significantly different be- tween populations (vein number per leaf, vein angle at mid-leaf, and adaxial pubescence) and character sets missing one or more data points (petiole diameter, leaf apex angle, number of teeth per three leaves, and vein angle — — BACON AND SPELLENBERG, Hybridization in Quercus ae) 150 100 }- LJ —_ Cc ® S Q s0F [1 = [e) ) 1S) So} ‘oO = So. -50 -100 | 1 | t L . 1 -250 -200 -150 -100 -50 0 50 100 150 principal component 1 Fic. 7. Principal component plot of PC1 and PC2 of all populations sampled (1042, 1044, 1045, 1046, 1047, 1048, 1049). Loading of axes is explained in text. A priori identification of individual plants is represented by squares = Q. conzattii (filled squares from Ferreria swarm site, 1042); diamonds = hybrids, backcrosses, and later generation segregates; and circles = Q. eduardit. Numbers labelling points refer to specific collections (1045-41, -44, -49, -50, -51; 1042-9) referred to in the text. at mid-leaf). Axis PC1 accounted for 71.0% of the variance, loading charac- ters as follows: cordateness of the leaves (-0.50), trichome maximum branch length (-0.39), and leaf length (-0.35). Axis PC2 accounted for only 8.6% of the remaining variance, with cordateness (0.70), trichome maximum branch length (-0.41) and petiole length (0.32) having the greatest load- ing values. The two axes jointly account for 79.6% of the total variation. The PCA plot shows Q. edwardii from all sites form a tight group. Quercus conzattii, however, is much more loosely grouped, extending from the up- per left corner of the diagram to the lower center. Quercus conzattii plants from Mezqguital fall approximately in the center of this distribution and do not extend to the extremes of distribution, as do plants from the Boca del Mezquital and Ferrerfa sites. The points representing plants we identified (a priori) as hybrids are mostly between Q. conzattii and Q. eduardii, overlap- ping somewhat with the former. This reinforces the discriminant function analysis assignment of some plants of Q. conzattiz to the hybrid swarm (marked 49, 50, and 51 in Fig. 7). It likewise accounts for discriminant function analysis inclusion of two plants with immature leaves (marked 41 and 44 in Fig. 7) in the hybrid swarm group. The point designated “9” on 34 Sipa 17(1) the PCA plot represents a plant originally identified as a hybrid (second pair of leaves in Fig. 2 is from this plant), but it has large, deeply-notched, rugose, reticulate, obovate leaves, and dense abaxial pubescence, as one would expect to find on Q. conzatti7. Data from this plant probably played a role in the clustering of Q. conzattii at Ferrerfa with the hybrid swarm, rather than with other populations of Q. conzattii. Even excluding these individual speci- mens, there still is considerable overlap between distributions of Q. conzattii and hybrids. This pattern, in which one species overlaps with hybrids and one does not, is similar to that found by Rushton (1978) in hybridizing Quercus robur L. and Quercus petraea (Matt.) Liebl. EVALUATION OF FERTILE STRUCTURES On 3 and 4 July, 1992, 63 plants were visually ranked ona scale from 1— 5, based on their vegetative similarity to either Q. conzatti (rank 5) or Q. eduardi (rank 1). Intermediate individuals were ranked as three (3), and other intergrades were ranked as 2 or 4. After ranking, inflorescences on each plant were examined to determine the stage of acorn development. They were ranked from 1—5, with ranks ranging from pistillate flowers just appearing on the peduncles (1) to acorns fully mature (5). Number of acorns per peduncle were counted for each peduncle. Correlation coefficients were calculated for mean acorn maturity rank and mean number of acorns per peduncle against the initial ranking of hybridity. Plants with Q. conzatti-like morphology generally had 4—5 immature acorns per peduncle in early July, and plants with Q. edvardii-like mor- phology generally had peduncles with 2—3 more mature acorns (see ex- amples in Fig. 2). Plants with intermediate vegetative morphology exhib- ited intermediacy in these characteristics (3-4 acorns per peduncle). Taxonomic rank of trees was a ue correlated (P < 0.05) with both number of acorns per peduncle (r = 0.62) and stage of acorn development (r = 0.66). Consequently, it appears oe acorn morphology and phenology correspond to vegetative morpholog Rushton (1978, 1988) showed that morphological intermediacy is ac- companied by reduced pollen viability in some plants of Q. robur and Q petraea hybrids, and Olssen (1975) found similar results, but wich rela- tively high pollen stainability in most F, hybrids. Only four plants with pollen were available to us, and two did not provide the 200 grains usually assessed. Nevertheless, pollen in each of the four specimens showed high stainability with cotton blue in lactophenol (Radford et al. 1974). The stainability in each specimen was as follows: Q. conzattii (Gonzdlez E. 5067) 98% (n = 59 grains); Q. eduardii (5056) 83% (n = 122 grains); Q. conzattit- like hybrid (5079) 97% (N = 260); and Q. eduardii-like hybrid (5081) eam — BACON AND SPELLENBERG, Hybridization in Quercus 35 97% (N = 230). This limited sample suggests that pollen fertility in both parental species and the hybrids is high. Consequently, it appears that hy- brids, backcrosses, and Fy segregates are capable of contributing pollen- mediated genes to future generations. DISCUSSION Examination of other populations in south-central Durango, northern Jalisco, and extreme western Zacatecas shows that hybridization is not unique to the Ferreria site. Occasional hybrids were found (sometimes only after extensive searching) where Q. conzattii and Q. eduardii occur sympatri- cally. Such hybridization appears to be more frequent in the relatively xeric habitat at the northern end of Q. conzattii’s northern distribution, such as at Pilar de Zaragosa (1286), Nombre de Dios (1400), and north of Boca del Mezquital (7293), all near the south edge of the Valle de Guadiana near Cd. Durango. Even in these places, the level of hybridization never ex- ceeded that which we found at Ferrerfa (/045). Variation in the amount of hybridization in these species ranged from that found in other species pairs thought to be introgressed (Stebbins et al. 1947; Cooperrrider 1957; Silliman & Leisner 1958; Bensen et al. 1967; Maze 1968; Hardin 1975), as exemplified by the Ferrerfa hybrid swarm (with ca. 14% plants of hybrid origin) to very little (as in the few plants found in Pilar de Zaragosa {1286}, Nombre de Dios {7400}, and north of Boca del Mezquital [/293}), as Dupouey (1983) found with Q. robur and Q. petraea. There are also sites where hybridization seems limited to the occurrence of probable F, indi- viduals (as in Mezquital [7483] and Bolafios 1/681), as Tucker and Bogert (1973) found with Q. gambelzi Nutt. and Q. turbinella Greene. Significance of hybridization between Q. conzatti and Q. eduardii in this region could be better assessed if it were known whether genetic markers cross species bound- aries and spread beyond where hybridization now occurs. Several authors have pointed out the importance of ecological factors in the control of interspecific hybridization between oak species (Muller 1952; Tucker 1961; Rushton 1979). Q. conzattii and Q. eduardii appear to be sepa- rated by soil type, and perhaps slope (affecting exposure) at the margin of the range of Q. conzattit. Quercus conzattii occurs on rocky soil, and often steep slopes. Quercus eduardit, commonly occurs on somewhat finer soils, often in gently rolling terrain. Climatic factors may play a role in limiting (or stimulating) hybridization. In less xeric environments, farther from the northern extreme of Q. conzattii’s range, hybridization between the two parental species seems quite limited, as near Mezquital, Durango (/483), and near Bolafios, Jalisco (1/68/). Whenever we encountered a plant of hybrid origin in a mature forest where environmental conditions appeared 36 Stipa 17(1) less extreme (in terms of aridity), it was solitary and intermediate in char- acter to the parents, suggesting that it was an F, individual. Apparent backcrossed individuals were absent. The difference between populations where only Fys occur and where later generation segregation occurs may relate to the degree of disturbance these sites have experienced, as suggested by Grant (1981) and Silliman and Leisner (1958). The sites near Ferrerfa (J042), Nombre de Dios (1400), Boca del Mezquital (northern population) (1293), and Pilar de Zaragoza (7286) all occur on rolling hills. They are also near villages, and frequent disturbances from livestock and human activities such as wood cutting and other habitat disturbances are common. Dodonaea viscosa, a weedy shrub of poor, disturbed soil, is abundant at all of these sites. The site at Ferrera, for example, has both open and disturbed conditions that Grant (1981) and Silliman and Leisner (1958) mentioned as important in hybrid swarm for- mation. It is near an area that has been utilized by humans for 3,000 years (Hendricks 1961) and is along an old road apparently many dec -ades old. Despite hybridization in Q. conzattii and Q. eduardii, the two species have remained morphologically distinct. As Stebbins et al. (1947) con- cluded for the Q. s/icifolia Wangenheim x Quercus marilandica Muench. population in the eastern United States, several isolating barriers may be operating. Just as in their example, seasonal isolation appears to be of mi- nor importance here. Flowering time in both species is poorly documented, but our few observations indicate that they overlap (late spring, but per- haps peaking two or three weeks apart). The flush of growth that occurs after a fire also results in at least a few plants blooming after the normal blooming date, perhaps serving to reduce barriers to cross pollination. Unfortunately, there is little published information concerning the affect of fire on hybridization. High pollen stainability indicates that hybrid in- fertility is not an isolating factor. Neither is failure to produce acorns, but the comparative viability of embryos is not known The percentage of putative F, hybrids in the population is relatively small (4%) within the area we defined as the hybrid swarm, and even the percentage of all putative hybrids (including Fy individuals) in the popu- lation seems rather small (14%) (Fig. 3). These numbers become very much reduced if one considers the extended population of Q. eduardii to the south. This suggests that post-zygotic selection is probably a key isolating factor between these species. Although introgression beyond the hybrid swarm was not detectable in the morphological characters examined in this study, the frequency of hy- bridization events between Q. conzattii and Q. eduardi1, the extent of hy- bridization at Ferrerfa, and the continuous variation encountered there sug- gests that it should play a role in evolution of these species as a means of BACON AND SPELLENBERG, Hybridization in Quercus a7 introducing genetic diversity. This may be particularly important at the edge of the species’ ecological ranges, a factor that could facilitate direc- tional evolutionary change (P.R. Grant & B.R. Grant 1994 and references therein). Further examination of the species, by other means, could provide more insight into gene exchange between these species. ACKNOWLEDGMENTS A New Mexico State University (NMSU) Arts and Sciences mini-grant #92—027 financed SEM work done by Hank Adams and Mark Cunningham at NMSU and Angela Welford at the University of New Mexico. Consider- able assistance was provided by the NMSU Foundation “Friends of the Herbarium” Fund. The NMSU Department of Biology also provided funds for travel. Dr. Socorro Gonzalez E. sent flowering oak specimens to NMC and provided lodging. Dr. Jorge Nocedal also provided lodging. Dr. Cecilio Barrera and Dr. Jorge Bretado V. proofread the Spanish abstract. Abel Garcia Arevalos and Sergio Rodriguez T. assisted us on several occasions 1n the field. The support and encouragement of A. Araceli Arias de Bacon is deeply appreciated. This paper is based on a master’s thesis at NMS APPENDIX Documenting specimens. Collections, locations of samples and characteristics of oak species and hy- brids in Mexico. Herbaria are abbreviated — to Holmgren et al. (1990). Specimen numbers listed are as they appear on specimen labels (e.g., Bacon, Spellenberg, & Garcia A 1290), which, in the e of specimens of one or both of the authors, may include a hyphenated suffix (e.g., Spellenberg & cas Bacon 1097312), indicating individual tree number within a collection of individuals of a popula- tion EXICO. Durango: Cerro C oe 1 km S of Puerto de la Cantera and Ww of Pilar de Zaragosa @ Rio Santiago, 104°W,24°N, 4 Jul sce Bacon & Paeeaed 1287 (CHDIR, NMC); ca. 28 air km E of ai ae and 2 km S$ of Fiwy 45 r N end of Ist mountain range E of neers ” 104024" he N, elev. 1860 m, 7 Jan 1993, Bacon 7 Spellenberg 1397 (CAS, IBUG, INI ou 11 road km of Nombre de Dios on Hwy 45, et W,23°54'N, elev. 1950 m, 7 Jan oe ngG Sle 1398 (BH, CHDIR, IEB, NMC); ne N of Boca del Mezquital, eee O'N, 5 Jul 1992, Bacon, » aaads & Garcla- A. i 0c AS, CIIDIR, F IBUG, MEXU, , TEX); near km marker 12, ad km S of Ferrerfa along the hwy to La Flor, 104°42'W,23°56'N, ee een m, 5 i Te Bacon, fe Ee ane T. 1045 (CIIDIR, F, ae eras qe X, TEX); 2 km E of the village of Boca del Mezquital, near km marker 31, om ere to Mezquital, 104°42'W,23°56'N, elev. 1920 m, 5 Aug 1991, Bacon, Cee Rade. & Garcia-A. 1046 (CUDIR, IEB, MEXU, NMC, TEX, ); 10 road km SE of Mezquital, 104°22'W,23°27'N, elev. 1,860 m, 5 sone 1991, Bacon, Spellenberg, Rodriguez-T. , & Garcia- A. 1047 Gon IEB, MEXU, NMC, TEX); 3 km S of Ferrerfa, along the road to La Flor, eee ne 7'N, 9 Apr 1992, Gonzdlez-E. ave (CHDIR),; 2 km E of village of Boca del Mezquital, 104°26'W, 23°46'N, elev. 1950 m, | Nov 1992, ellalers G Bacon 10973—12 (NMC). “ alisco: 9 air km NW of Bolafios in the center of the northern extension of Jalisco, 20 road km W of olafios on the asce 5 the mountain, 103°5 1'W,21°53'N, elev. 2070 m, 23 Jan 1993, Bacon G eee 1482 On MEXU, NMC) uercus conzattit Trel. x ° comnane Trel. cen CO. Durango: Cerro Chiquihuitillo, lkmS of Puerto de la Cantera and s e W of Pilar de Zaragosa at Rio Santiago, 104 W,24 N, 4 Jul 1992, Bacon & Spellenberg er ae F, ce IEB, INIF, NMC); 11 road km W of Nombre de 38 Sipa 17(1) Dios on Hwy 45, 104°20'W, 23°54'N, elev. 1950 m, 7 Jan 1993, Bacon & Spellenbere 1400 (BH, CAS, CHIDIR, IEB, MEXU, NMC, NY, RSA); 35 road km S$ of Cd. Durango, along a road from Duras? to Mezquital, 104930! W,23°40'N, 5 Jul 1992, Bacon, oes & Garcta- a 293 (CHDIR); 4.3 road km S of Ferrerfa along the hwy to La Flor, near km marker 12, 104°42'W, ee N, elev. 1920 m, 4 Aug 1991, Bacon, Spellenberg & Rodrignez-T. 1042 (CIIDIR, F, IEB, MEXU, NMC, OAX, TEX); 3 km S of Ferrerfa, along the road to La Flor, 104°22'W,23°27'N, 9 Apr 1992, Gonzdlez-E, 5056 (CIIDIR); 3 km S$ se Ferrerfa, along che road to La Flor, 1( J4°22'W2 seen 9 Apr 1992, Gonzdlez-E, 507 ie (CIIDIR); 10 road km SE of Mezquital on road up mountain to Las Mi argaricas, 104°22'W,23°27'N, elev. 1860 m, 5 Jan 1993, Spellenbere & Bacon 11681 (NMC); ca. 9 km S of sed sons hwy to La Flor, 104°42'W, 23°56'N, elev. 1860 m, 30 Oct LOOL, Spellenberg, ape & Sooby 71 (NMC, MEXU); ca. 10 air km S of Cd. Durango on road to La Flor, 21 Jul 1990, ae : Gonzdlez 10206 (MEXU, NMC), Jalisco: 20 road ee W of Bolanos up ascent to moun- tains toward Los Amoles, 103°5 1'W,21°53'N, elev. 2070 m, 23 Jan 1993, Bacon & Spellenbere 1483 (CAS, MEXU, NMC). (a) | [rel —MEXICO. Durango: Cerro Chiqui! a 1 km S$ of Puerco de la Cantera and somewhere W of Pilar de Zaragosa at Rio Santiago, 24°N, 104 4 Jul 1992, Bacon & Spellenberg 1288 (CHDIR, NMC); 11 road km W of Nombre de Dios on a 45, 104°20'W,23°54'N, elev. 50m, 7 Jan 1993, Bacon & Spellenbere 1401 (CIIDIR, NMC); 5 road km S of Ferreria along the hwy o La Flor, near 7 marker 13, 104°42'W,23°56'N, elev. 1920 m, 4 cae 1991, Bacon, Spellenberg, & Rodriguez-T. 10 CIIDIR, F IEB, MEXU, pe OAX, TEX); 10 road km SE of Mezguital along oa road to Las ere is, 104°22'W/,23°27'N elev. 1,860 )m,5 ue ee Bacon Spellenberg Rodrienex- ,& Garcia-A. 1048 (CIHDIR, IEB, MEXU, ey — ca. 9 air aa S of sere at top of the ae ascent of ae road S to La Flor out of the valley where Cd. Durango is situated, 2 air km SW of Ferrerfa, 104°42', 23°50'N, elev. 1920 m Spellenberg ae 11692 (MEXU, NMC) “3 S road km S of Ferrerfa along the road to La Flor, across the hwy from the monument now as La ere 104°42'W/23°56'N, elev. 1,860 m, 6 ae 991, Bacon, Spellenberg, Rodrieuez-T., & Garcta-A. 1049 (CIHDIR, IEB, MEXU, NMC, TEX); 3. km S$ of Ferrerfa, along the road to La Flor, 9 Apr Gonzdlez SOS 1 (CHDIR); 6 km S of Ferrerfa, elev. 1860 m, 2 Nov 1991, Spellenberg, Boecklen, & Sooby 10974, (NMC), Jalisco: ca. 21 road km NW of cnee on the road to Los a 103°52'W,21°53'N elev. 2150 m, 24 Jan 1993, Spellenberg & Bacon 11752 (CAS, IBUG, NMC REFERENCES ANbDERSON, E.A. 1949. Introgressive hybridization. John Wiley and Sons. es York. ANDERSON, E.A. 1953. Introgressive hybridization. Biol. Rev. 28:280—30 ANDERSON, E. aaa G.L. Steppins. 1954, Hybridization as an evolutionary stimulus. Evolu- tion 8:378—388. ARNOLD, M.L. 1990. 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Quercus robur L. and Quercus petraea (Matt.) Liebl.: a multivariate approé ch to the ee problem, 2. The geographical distribution of population types. 24 Nol Watson RUSHTON, B. S. ne Pollen ees and the hybrid oak (Quercus robur L. xX Q. petraea (Matt) Liebl.) controversy. In: rty, ed. Taxonomy, Putting Plants and Animals in their Place. R. Ir. Acad., oe Pp. 89-110. RzEpowskl, J. 1978. Vegeraci6n de México. Editorial Limusa, México SAS Institute, Inc. 1992. SAS System, Release 6.07, for the VMS Bagiconmene Cary, North Carolina. SILLIMAN, FE. and R.S. LeisNer. 1958. An analysis of a colony of hybrid oaks. Amer. J. Bot. 4 2 5:730-736. SPELLENBERG, R. 1992. A new species of black oak (Quercus, an . le Fagaceae) from the Sierra Madre Occidental, México. Amer. J. Bot. 206 BACON AND SPELLENBERG, Hybridization in Quercus 41 SPELLENBERG, R. and J.R. BAcon. 1996. Taxonomy and distribution of a natural group of black oaks of Mexico (Quercus, section Lobatae, subsection Racemiflorae). Syst. 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Bos ERT. 1973. a of a progeny test of a hybrid oak, Quercus gambelii x Quercus turbinella. Madrono 22:1— VaN DEVENDER, he R. ae eat climates ane endemism in the Chihuahuan Desert flora. In: we Je . Powell, and B.N. Timmermann, eds. Transactions of the Second — on Resources 7 the Chihuahuan Desert. Chihuahuan Desert Research Institute, Alpine, Texas. 19. Van Devenper, T.R. 1990. Late eae vegetation and climate of the Chihuahuan Desert, United States and México. In: Betancourt, J.L., T.R. Van Devender, and PS. Martin, eds. Packrat Middens: the Last 40,000 Years of Biotic Change. The University of Arizona Press, Tucson. Pp. 104-133 VAN DEVENDER, T.R., R.S. THOMPSON, anda L. Betancourt. 1987. Vegetation history of the deserts of southwestern North America; the nature and timing of the Late Wiscon- E. Wrig sin-Holocene transition. In: Ruddiman, W.F. and I ight, Jr., eds. The geology of North America, Vol. K-3, North America | adjacens oceans during the last eee The Geological Society of America. Pp. 3 VAN VALEN, L. 1992. Ecological species, multispecies, and Bake, In: M. Ereshefsky, ed. The units of eston st on the nature of species. Bradford Book, MIT Press, Cam- bridge, Mass. Pp. 65-77. Wison, P. 1992. On inferring hybridity from morphological intermediacy. Taxon 41:11—23. Wuitremore, A.T. and B.A. ScHaaL. 1991. Interspecific gene flow in sympatric oaks. Proc. Acad. Sci. 88:2540—2544. ZAVALA C.F. 1989. Identificacién de encinos de México. Universic México, México. ad Autonoma, Chapingo, 42 Stipa 17(1) BOOK NOTICES GoopMAN, GeorGeE J. and CuHeryt, A. Lawson. 1995. Retracing Major Stephen H. Long’s 1820 Expedition. The Itinerary and Botany. (ISBN 0-8061-2703-1, hbk). University of Oklahoma Press, P.O. Box 787, Norman, OK, 73070-0787. The American Exploration and Travel Series, Vol. 73. $38.50. 366 pp. In Part I, The Itinerary, Goodman and Lawson document the day by day account of the rouce and the location of each night’s campsite. From the Missouri River in Nebraska near orado, New Mexico, Texas, and Oklahoma ending at Fort Smith, =— Council Bluffs through Co the authors traveled and retraced the route themselves noting the vegetation anc and form ante during that interval. In Part H, The Botany, the collections of Edwin James, M.D. are studied alphabetically by family a in conjunction with their own collections from the same sites. Appendix One ts a list of plant names commemorating James based on collections made along the route of the expedition. Appendix Two lists the types collected by James. The remaining indices include Sources Cited, Index to Scientific Names, Index to Common Names and General Index This is an excellent book based on field work by the two authors over 150 years later . With the maps and selected illustrations of then and now, it shows the effects of erosion and vegetation changes over this time period. Their study of the plants and types will aid the researcher in having an analysis readily available. This is a book that should be in everyone's library.— Wm. F. Mahler. WELSH, STANLEY L. 1995. Flowers of the Canyon Country. (ISBN 0-87480 -486-8, pbk). University of Utah Press, 101 University Services Bldg., Salt Lake City, UT 84112. Photography by Bill Ratcliffe. Paperback $15.95. 85 pp. Color photographs. Copyrights: 1971, 1977, 1986. The canyon lands in this text refer to the Four Corners area where Arizona, Colorado, New Mexico, and Utah meet. The Foreword was written by Glen Moore, The nontechnical text was written by Welsh for Using the Book, The Canyon Country, and The Flowers sections. The plo ea shy of Bill Ratcliffe is outstanding and has been reproduced from 4.25 by 4.75 inches to more than 8.5 by 11 inches in at least one photo. Everyone should — enjoy just browsing oa pondering the photographs. The text of the taxa includes a lot more information than just the typical species descriptions. —Wm. F. Mahler. Sipa 17(1): 42. 1996 INVASION AND SPREAD OF COINCYA MONENSIS (BRASSICACEAE) IN NORTH AMERICA ROBERT EC. NACZI and JOHN W. THIERET Department of Biological Sciences Northern Kentucky University Highland Heights, KY 41099, U.S.A. ABSTRACT Invasion and spread of che European native Co/ncya monensis (Brassicaceae) 1n North America are traced. In the late 1800s the species was found in the Philadelphia, Pennsyl- vania/Hoboken, New Jersey area on ballast, where it has not persisted. Then, in the mid and late 20th century, naturalized populations of unknown source were documented in North Carolina, Pennsylvania, New York, Michigan, and Kentucky. RESUMEN Se esboza la invasién y dispersion de la planta nativa de Europa C oimeya MONENSIS (Brassicaceae) en Norte América. A fines del siglo se encontré la especie en el drea de Philadelphia, Pennsylvania/Hoboken, New ae sobre cascajos, ive ade no ha periciee: ia a mediados y as del siglo veinte, han sido documentadas poblaciones origen de: n Carolina del Norte, Pennsylvania, New York, Michi- gan, y Kentucky. INTRODUCTION In August 1992 JWT noted an abundant, robust, showy, yellow-flowered crucifer unknown to him along interstate highway 80 in central Pennsyl- vania. After making a collection of the plant, he drove on, observing the plant for many more miles at the roadside. Later, at Northern Kentucky University, work on identification of the plant was begun. That the species is not included in the standard north- eastern North American manuals (Fernald 1950; Gleason & Cronquist 1963, 1991) soon became obvious. Referral to Flora Enropaea (Tutin et al. 1964) finally brought an answer: JWT identified the plant as Rhyuchosinapis monensis (L.) Dandy ex Clapham in Clapham, Tutin & Warburg, one of several bino- mials recently applied to it. Wherry et al. (1979) included it as RAynchosinapis cheiranthos (Vill.) Dandy. Al-Shehbaz (1985) treated it as Hutera cheiranthos (Vill.) Gémez-Campo, the name under which it appears in Rhoads and Klein (1993). The second edition of Flora Europaea (Tutin et al. 1993) uses Coincya monensis (L.) Greuter & Burdet, the name determined to be correct by Leadlay and Heywood (1990). As the common name for this species we suggest and will use “coincya.” Stipa 17(1): 43-53. 1996 44 Sipa 17(1) During our joint field work in Pennsylvania in 1994, we made a special effort to collect coincya from as many counties as possible. Then the next year RFCN and JWT separately made additional collections of the species in Pennsylvania, the state in which it appears to be most common and where it is spreading. In this paper we trace the species’ current range in North America and summarize its collection history on the continent. MATERIALS AND METHODS To document the history of invasion and the species’ current range in North America, we requested loans of herbarium specimens of North American Cozncya and of species under which we thought coincya might be filed, incorrectly identified. We also asked for loans of unidentified cruci- fers resembling the species (photocopies of several of our specimens of coincya were included with each loan request). Of the 126 United States and Cana- dian herbaria to which we wrote, 90 responded. Many had no specimens of the species, but overall we received 353 sheets of miscellaneous weedy, silique-bearing crucifers. Among these specimens—including sheets of Arabis, Brassica, Erucastrum, Erysimum, Raphanus, S inapts, Sisymbrinm, Tropi- docarpum, and, especially, Diplotaxis—we located 62 sheets of coincya rep- resenting 22 separate collections (some were widely distributed duplicates). When to these 22 collections are ac eos ded the 23 that we made, the total is 45 collections of C. monensis we have seen from North America (see appendix). IDENTIFICATION, TAXONOMY, AND NOMENCLATURE Coimcya monensis 1s best distinguished by its combination of conspicu- ously glaucous and distinctively shaped leaves (Figs. 1, 2); a usually ebracteate inflorescence (occasionally an elongate, entire bee subtends the most proximal pedicel) (Fig. 3); bright yellow petals with pale brown or violet veins (the blooming season is long: late April through October); a single row of seeds in the siliques, which have clearly 3—5 veined valves and a somewhat flattened, 1—6 seeded beak (Fig. 4). The mature silique is 3.5—8.5 cm long (including the beak); the beak is 15-20 mm long and ca. 2 mm wide at its base. Most North American botanists are certainly as unfamiliar as we were with Corncya monensis. In Table 1 we detail ways to separate C. monensis from those crucifers that superficially may resemble it in eastern North America. Comcya belongs to the tribe Brassiceae of the Brassicaceae (Cruciferae). This tribe is characterized by fruits being usually beaked, the beak typi- cally 1- or few-seeded, and the cotyledons being conduplicate (Al-Shehbaz 1984). The genus is included as Hwtera in the key to southeastern United States genera of Brassicaceae (equally useful in northeastern United States) Naczi AND THIERET, Coincya monensis in North America 4S IZ AMZ ATA Fics. 1-2. Coincya monensis. Fig. 1 (left). Basal portion of plant. Fig. 2 (right). Selection of leaves from a single plant of Coincya monensis. The amount of lobing of the leaves decreases from the base of the plant to the base of the inflorescence. Vertical bar represents 5 cm for both figures. by Al-Shehbaz (1984) and in his key to southeastern United States genera of Brassiceae (Al-Shehbaz 1985). Taxonomy and nomenclature of Coincya were discussed by Leadlay and Heywood (1990), who recognized five subspecies of C. monensis. Further discussions of the species’ nomenclature are in Kartesz and Gandhi (1994) and Rollins (1961, 1993). Our plants are subsp. recwrvata (All.) Leadlay, a taxon burdened by no fewer than 129 synonyms. This subspecies has the same indigenous range in the Old World as that of the collective species: Corsica, France, Germany, Great Britain, Italy, Morocco, Portugal, and Spain (Leadlay & Heywood 1990). INVASION AND SPREAD Except for the species’ convoluted nomenclature, the invasion history of C. monensis in North America is straightforward. Apparently there are five areas in eastern United States in which the plant has appeared (Fig. 5), one of these in the late 1800s only, the others after the middle of the 20th century. In the Philadelphia/Hoboken area it was collected on ballast in AG Sipa 17(1) Fics. 3-4. Coineya monensis. Fig. 3 (left). Fruiting branch. Vertical bar represents 5 cm Fig. 4 (right). Fruits, one with one valve removed. Note conspicuous beak and the one row of seeds. Vertical bar represents | cm. the 1870s to at least 1890. Then, in 1958, it was found in far western North Carolina, where it has persisted at least into the 1990s. Then fol- lowed Pennsylvania and southern New York, where the species was first collected in 1964 and where we made 22 collections of it in 1992, 1994, and 1995. In southwestern Michigan it was first collected in 1989 and again in 1993. Finally, the species was found in Kentucky in 1996. Philadelphia/Hoboken area on Ballast (1870s to at least 1890) Coitcya monensis was one of many species introduced in ships’ ballast in ye late 1800s. Soil, sand, and rocks incidentally containing seeds were — i loaded onto ships in Europe and then exchanged for cargo in the United States. Studies of these ballast floras recorded at least 386 species associated with ballast in New Jersey, New York, and the Philadelphia area alone (Muehlenbach 1979), The earliest North American reports of C. monensis we have located—all from ballast in the vicinity of Philadelphia—are those of Burk (1877) and Martindale (1876, 1877). Further ballast collections were made by Brown (1879, 1880) from Hoboken, New Jersey. In these early reports the species Naczi AND THIERET, Coincya monensis in North America 47 Taste 1. Characteristics distinguishing Co/ucya from selected genera of Brassicaceae with which it has most often been confused in eastern North America. From Brassica Coincya Brassica Valves of fruit with 3—5 veins Valves of fruit with 1 vein From Diplotaxis oincyd rine Seeds in 1 row in fruit eeds in 2 rows in fruit Beak of fruit broad, somewhat flattened, oo of fruit style like, 1.5-3 mm long 15-20 mm long, ca. 2 mm wide at base From Erucastrum Coincya Erucastrum Inflorescence ebracteate (occasionally an Inflorescence bracteate at least at the most elongate, entire bract at base of most proximal Ss ide the bracts pinnately lobed proximal pedicel) Beak of fruic broad, somewhat flattened, Beak of fruit slender, style-like, co 4 mm 15-20 mm long, ca. 2 mm wide long at base From Sfnapis Coincya Sinaprs Veins of petals darker in color than the rest — Petals uniform in color the blade Sepals erect, connivent, the inner two Sepals tend to be spreading (rarely saccate at reflexed), not connivent, not saccate at base was called “Brassica monensis” or “B. cheiranthus.” It was present at Philadel- phia uncil at least 1890 (the date of the most recent ballast specimen we have seen). At the 19th century ballast sites, C. monensis was described as being “scarce” (Brown 1879) to “quite common” (Martindale 1877). Changes in regulations regarding the disposal of ballast eventually closed this avenue of introduction. These early introductions on ballast apparently did not lead to natural- ization of the species in North America. Such naturalization followed only after the post-1950 appearances of the species on the continent. Far Western North Carolina (1958—Present) After a hiatus of about seven decades, the plant was again found in North America: collections of it from far western North Carolina (Yancey County) made in 1958 were reported by Ahles and Radford (1959) but under the misidentification Diplotaxis muralis; this misidentification was “corrected” in the Vascular flora of the Carolinas (Radford et al. 1968), where the species appears as Brassica erucastrum L., a misapplied name (Al-Shehbaz 1985; Leadlay & Heywood 1990). The North Carolina collections were discussed by Rollins (1961). 48 Sipa 17(1) ue ,500km if Coincya monensis OO 1879-1890 O 1958-1968 @ 1975-1996 ie) A seunverified report Fic. 5. Commeya monensis. Documented range in North America. Symbols indicate year of -_ first collection from a county At the sites of this first United States collection in the 20th century, the species was said to be “very abundant, turning roadsides and fields into a yellow haze for miles around when in flower, particularly in the Cane River Valley. As a weed, it would seem to have the potential of becoming a very serious pest” (Ahles & Radford 1959) The most recent collection of coincya we have seen from North Carolina was made in 1994, again in the far western part, about 15—20 miles south of the sites of the original collections from the state. Pennsylvania/New York (1964—Present) Pennsylvania.—Vhe earliest 20th century collection of C. monensis we have located from Pennsylvania (Luzerne County) is dated 1964. The first pub- lished Pennsylvania record is in Atlas of the flora of Pennsylvania (Wherry et al. 1979). The updated Pennsylvania atlas (Rhoads & Klein 1993) reported the plant from two counties (Clearfield & Luzerne), mentioning chat it is “apparently spreading along Interstate 80.’ Our Pennsylvania collections, made in 1992, 1994, and 1995, are from 19 counties. We suggest that coincya is spreading into New York from Pennsylvania and that the species will become more common in New York than it seems to be now. We know of two collections and one undocu- mented report from the state, all near the Pennsylvania border. Both of the New York specimens are from the 1990s: a JWT collection from a roadside near Port Jervis in Orange County; and a Chapman collection from “steep rocky walls and small cliffs” at South Addison in Steuben County. The report is in notes accompanying W. Chapman’s Bradford County, Pennsyl- Naczi AND THIERET, Coincya monensis in North America 49 vania, collection, which mention that the species is “rare in New York, though it occasionally shows up in the south-central (Binghamton) area.” We located no verifying voucher, although we have no reason to doubt the report's accuracy. Michigan (1989—Present) Swink and Wilhelm (1994) reported collections of the species from south- western Michigan (Berrien County) in 1989 and 1993. The sites where the plant was found are only about 30 miles north of Interstate 80, the high- way along which JWT first noted the plant in Pennsylvania ca. 400 miles to the east. Kentucky (1996-present) The first collection of the species from Kentucky was made in 1996 in Rowan County ona rocky roadside slope. Our map shows not only the documented occurrences of C. monensis in North America but also clearly where we made a special effort to search out the species in 1994 and 1995 in Pennsylvania. The advance of coincya in Pennsylvania may have been rather rapid: from one county in 1964 to at least 20 counties ca. 3 decades later. We predict with confidence that the range of C. monensis will increase. Indeed, the plant is probably already in some states additional to those reported here, having been overlooked or simply ignored by collectors, many of whom do not collect roadside weeds (especially those resembling Brassica, so commonly seen but not an es- teemed prize for collectors). Indeed, we suggest that in some herbaria from which we did not request loans—and maybe even in those 36 herbaria from which we received no response to our loan request—there might be specimens from states other than those we have verified. That seeds of C. monensis were introduced via ballast at the 19th century sites is clear. The source of the disseminules for the post-1950 introduc- tions is unknown. Two patterns of spread by invading plants were distinguished by Baker (1974). In the first, “echelon” movement, a species has a single point of introduction followed by movement across the landscape in wave- or step- like progression. In the second, “spot and fill,” several centers of introduc- tion may be followed by a filling in of the intervening open spaces. The pattern exhibited by C. monensis in North America in the 20th century would appear to be “spot and fill,” with the “filling in” yet to develop fully. The European range of C. monensis subsp. recurvata is from southern Spain north to western Germany and northern Scotland, covering ca. 21° of latitude (Leadlay & Heywood 1990). This range extends from hardiness zone 10 (minimum 30° to 40° F) to zone 7 (minimum O° to 10° F) 50 Sipa 17(1) (Kriissmann 1984). Considering temperature alone, we note that, in North America, the species already occurs in colder areas (zone 5, minimum -20° to - 10° F) than in Europe. ECOLOGY One Pennsylvania herbarium specimen (Bradford County, near Wyalusing, “late July” 1983, Chapman s.n. [BH}) has attached to it copies of two letters in which are useful data. Chapman wrote in 1983 that the species’ preferred habitat is “along the edge of rock ledges .. . and at the bases of such cliffs . .. especially in those areas where small cliffs are formed by the highway department’s cutting through a hill to make a road.” He further noted that the plants “often live 2—3 years and set a full crop of seeds each year.” The habitats mentioned by Chapman are precisely those from which most of our collections came; particularly impressive is the plant’s abun- dance in some of those habitats. In addition, we have seen the plant among cobbles placed on roadside slopes to retard erosion, along railroad tracks, at weedy edges of roadside rest areas, and at disturbed sites at service stations and motels. Many of the sites in which it grows are gravelly or rocky. The synanthropes associated with coincya in its weedy habitat are fre- quent to common along highways over much of northeastern United States. We noted Capsella bursa-pastoris, annual species of Cerastium, Coronilla varia, Erigeron canadensis, Erysimum repandum, Lotus corniculatus, Polygonum aviculare, Silene antirrhina, 8. cucubalus, and §. pratensis (Lychnis alba). Some associates of coincya in southwestern Michigan are listed in Swink and Wilhelm (1994). We noted in Pennsylvania, though, that coincya often dominated its habitat to the apparent exclusion of other species. A single, vigorous plant of C. monensis, sometimes nearly 1.5 m long (individuals that large are sprawling, not erect), is seemingly a formidable competitor, capable of covering a large area. Occasional plants of C. monensis may be found grow- ing in the midst of otherwise continuous Coronilla varia. Additional data on biology of the species are in Hegi (1918 funder Brassiwella erucastrum}, 1986 tunder Rhynchosinapis cheiranthos}) and Leadlay and Heywood (1990). We hope that our summary of data on this species will provide baseline data of some use to future investigators of the spread of the plant. ACKNOWLEDGMENTS We thank the herbaria that responded to our request for loans: United States: BH, BHO, BRL, BUT, CCNL, CHRB, CLEMS, CLM, CM, CONN, DAV, DEK, DOV, FE, FLAS, FSU, GH, HNH, JA, ISC, KANU, KE, KSC, LSU, LYN, MASS, MICH, MIN, MO, MOR, MSC, MU, MUHW, MW, NACZI AND THIERET, Coincya monensis in North America 31 NCBS, NCSC, NCU, NDG, NHA, NLU, NY, NYS, ODU, OSC, PAC, PH, PUR, RSA, SDC, SIU, SLRO, TEX, UARK, UC, UNA, UNCC, S, USCH, UTC, VDB, VPI, VT, WILLI, WIS, WVA, YU. Canada: ACAD, ALTA, CAN, DAO, HAM, MMMQN, MT, le NBM, NELD, NSPM, OAC, QFA, QK, QUE, SASK, SCS, SFS, TRT, UBC, USAS, UWPG, WAT. The Lloyd Library, Cincinnati, was invaluable for our lit- erature searches. The Spanish translation of the abstract was provided by Dr. Miriam Steinitz-Kannan. Our wives, Mary A. Naczi and Mildred W. Thieret, assisted with field work. APPENDIX Coincya monensis—Herbarium pci Seen NTUCKY. Rowan Co.: 3.5 air mi NE of center of Morehead, along W side of route 32, 1 Jun i658. Naczi & Trauth 5561 enir KNK, US, VDB). MicnHiGAn. Berrien Co.: Coloma, sandy soil ee railroad, 29 Aug 1993, seca & Dritz 21806 (MICH, MOR); Coloma, weedy hill, 16 Aug 1993, Wilhelm & Dritz 21579 (MOR); Coloma, sand soil, 3 Jul 1993, Dritz 1115 (MOR); roadside between Coloma and Watervliet, a 1989, Swink & Waren 8600 (MOR), near a large sandpit between Coloma and Watervliet, 30 May 1989, Swink & Wets a Ook ; y. {Hud Co.]: Hoboken, i allast, 25 Oct 1879, Brown s.n. (AC, NY); Hoboken, ballast élling, "Jul ee Hk n. (BH, GH, US); a ballast, 28 Oct 1879, Schrenk s.n. (NY); Hoboken, ballast, 3 Jul 1880, a ee sn. (CH 2) . Orange Co.: roadside along I-84, ca. 1.5 mi E of Port Jervis, 10 Sep 1995, Thieret 50751 (KNK). Steuben Co.: steep rocky walls aaa small cliffs overlooking Tuscarora Creek along route 417, South Addison, [no day, no month] 1994, Chapman s.n. Ed ORTH CAROLINA. Jackson Co.: field 3.9 mi NE of Cherokee, 7 Mag 1968, pee i ies ee 1453 (CHRB, FLAS, FSU, ISC, KE, LL, MBG, MIN, NCU, NC Y, NHA, NLU, NY, PAC, PUR, SIU, TEX, UNA, UTC, VDB, WIS, WVA). Yancey Co.: roadside 0.8 mi SE of ae 2p Apr 1958, AAles & Duke 39158 (NCU); Sees 1.8 mi SW of Burnsville, 7 Jun 1958, Ables G Duke 42855 CAN, FLAS, KANU, MICH, NCU RSA, SIU, VDB); roadside 2.9 mi SW of Burnsville, 7 Jun 1958, Ables & Duke 42859 NCU); ads 0.9 mi NW of Ramsaytown, 16 Jul 1958, Ahbles & Duke 46824 (FSU Puees peadide, Cane er, W of Burnsville, 9 Oct 1958, Ahbles G Duke ie (NCU); wet rock Blue ee Parkway 7.2 mi NE of Black Mountain Gap, 6 Aug 1994, Vincent & Lammers 6739 (MU, VDB). . ENNSYLVANIA oo NT =collection by R.RC. Naczi and J.W. Thieret; NN=collection by R.F.C. Naczi 1 M.A T=collection by J.W. Thieret.) Blair re pe 17 mi S of Altoona, 13 Jun 1994, we pan (KNK ). Bradford Co.: roadside, Indian yer Rocks, near Wyalusing, [no day} July 1983, Chapman s.n. (BH); roadside, Troy, 15 Jun na NE 4159 (KNK); along railroad tracks, ae 16 Jun 1994, NT 4237 (KNK). Carbon Co.: roadside just E of East Side, 18 Jun 1994, NT 4356 (KNK). Clarion Co.: roadbank 2 mi SW of Clinton, 4 Jul 1995, NN 4907 (CM, KNK, MICH, PAC, PH, VDB). Clearfield Co.: roadside, Clearfield, 19 Aug 1992, T 58141 (KNK). Clinton Co.: roadside 2 mi NNW of Loganton, 14 Jun 1994, NT 4120 (KNK). Delaware Co.: roadside 10 mi S of Norristown, 18 Jun 1994, NT 4359 pee Lackawanna Co.: roadside 2 mi SW of Mt. Cobb, 1 - as me (KNK, PH). Lehigh Co.: roadside, Allentown, 18 Jun 1994, NT 4357 (KNK). rne Co.: roadside, Hazelton, 10 Sep 1995, T 59750 (KNK); roadside 2.5 mi SE of Briggsville, - Jun 1964, eee (FSU, ISC, MASS, PAC, UC, VDB). Lycoming Co.: roadside 1 mi NE of Picture Rocks, 14 Jun 1994, NT 4123 (KNK). Monroe Co.: roadbank SE of Stroudsburg, 9 Jun 1975, Rossbach 9216a (WVA). Montgomery Co.: roadbank 5 mi NE of Norristown, 18 Jun 1994, NT 4358 (KNK). Northumberland Co.: roadside 2 mi NW eal 14 Jun 1994, NT 4122 (KNK). Philadelphia Co.: ballast, West Philadelphia, 22 Jun 1890, Mc E/wee 570 (PH). Pike Co.: roadside 3.5 mi SSW of Tafton, 1 Jul 1995, NN 4882 (CM, KNK, PH); roadside 3 mi NW of 52 Sipa 17(1) Milford, 3 Jul 1995, NN 4883 (CM, KNK, PAC, PH). Schuylkill Co.: roadside between McAdoo and Delano, LO Sep 1995, T 59757 (XNK). Sullivan Co.: roadside 5 mi SW of Laporte, 14 Jun 1994, NT 4124 (RNK). Union Co.: roadside 7 mi NW of New Columbia, 14 Jun 1994, NT 4121 (KNK). Wayne Co.: roadside 5 mi NNW of Pocono Springs, 1 Jul 1995, NN 4871 7 -M, KNK, PH). Wyoming Co.: roadside 1.5 mi SE of Meshoppen, 18 Jun 1994, NT 4353 (SNK REFERENCES AuLes, H.E. and A.E, Seen — Species new to the flora of North Carolina. J. Elisha Mitchell Sci. Soc. 75:1 ae AL-SHEHBAZ, I.A. 1984. The ae of Cruciferae (Brassicaceae) in the southeastern United States. J. Arnold jee 65:343—373 1985. The genera of Brassicea ae ( Aaa Brassicaceae) in the southeastern United Spates. iG Hae Arbor. 66:279—' BAKER, ie 1974. The evolution of weeds. aa Rev. Ecol. Sysc. 5: 1-2: Brown, A. 1879. Ballast plants in New York City and its vicinity. Bull. atorey Bot. Club eh Sener . L880. Ballast plants in and near New York City. Bull. Torrey Bot. Club Burk, I. 1877. List of plants recently collected on ships’ bi ne ast in the neighborhood of Philadelphia. Proc. Acad. Nat. Sci. Philadelphia 29:105— FERNALD, M.L. 1950. Gray’s manual of pony 8th ed. American a Company, New York. GLEASON, H.A. and A. CRON {anual of vascular plants of northeastern United States and adjacent a. > . Van eceaae New York. GLeason, H.A. and A. Crongquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York Botanical Garden, Bronx, New York. Hea, G. {1918.] Mluserierce Flora von Mittel-Europa. {1st ed.} Volume 4(1). J.E. Lehmanns Verlag, Miinchen. 1986. Illustrierte Flora von oo ae 3rd ed; edited by H.J. Conert, U. Hamann, W. Schultze-Motel, and G. Wagenitz. Volume 4. Angiospermae- Dicotyledones 2(1). Verlag Paul a Soe Karresz, J.T. and K.N. Ganpui. 1994. Nomenclatural notes for the North American flora. XIII. Bhycologia 76:441-457. KRUssMANN, G. 1984. Manual of cultivated broad-leaved trees and shrubs. Translated b M.E. Epp. Volume I, A—D. Timber Press, Beaverton, Oreg¢ Leapb.ay, E.A., and V.H. Heywoop. 1990. The biol logy and systematics of the genus Coscya Porta and Rigo ex Rouy (Cruciferae). J. Linn. Soc. Bot. 102:313-398. MARTINDALE, 1876. The introduction of foreign me ou Cis, 2:55-58. 877. More about ballast plants. Bot. Ga 2 on ds —12: MUEHLENBACH, 1979. Contributions to the syne anbropi advent flora of the rail- roads in St. Louis, Missouri, U.S.A. Ann. Missouri Bot. Gard. 66:1—108. sae A.E., H.E. AuLes, ane C.R. BELL. 1968. Mert “of the vascular flora of the rolinas. University of North Carolina Press, ee tae Till, North Carolina. Ps A.F. and W.M. Kirin Jr. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American Philosophical oe Philadelphia, Pennsylvania. Ro tins, R.C. L961. A weedy crucifer again reaches North America. Rhodora 63:345—346. 993. The C ruciferae of continental North America. Stanford University Press, Enron: California. SWINK, FE. and G. Winsteis. "1994. Plants of the Chicago region. The Morton Arboretum, Lisle, Illinois. Naczi AND THIERET, Coincya monensis in North America 533 Tutin, T.G., V.H. Heywood, N.A. Burges, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. 1964. Flora Europaea. Volume 1. Lycopodiaceae to Platanaceae. Cambridge Uni- versity Press, Cambridge, U.K. Tutin, T.G., N.A. BurGes, A.O. CHater, J.R. EDMONSON, V.H. Heywoop, D.M. Moore, D.H. Vinee S.M. Watrers, and D.A. Wess, eds. 1993. Flora Europaea. Volume 1. Psilotaceae to Platanaceae. 2nd ed. Cambridge University Press, Cambridge, U.K. Wuerry, E.T., J.M. Foe Jr., and H.A. Want. 1979. Atlas of the flora of Pennsylvania. The Morris Arboretum, Philadelphia, Pennsylvania. 54 Sipa 17(1) ANNOUNCEMENT The Council of the New England Botanical Club wishes to announce that there are no longer page charges for publication in Rhodora. The Council has appointed Janet R. Sullivan Editor-in-Chief and Margaret Bogle ee Editor, beginning with volume 98. Any new manuscripts should be submitted to: Dr. et R. Sullivan, Department of Plant Biology, University of New Hampshire, es NH 03824-3597: (603) 862-4757, e- mail: janets@christa.unh.edu. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany or conservation of the flora of North America or floristically related areas will be considered. Sipa 17(1): 54. 1996 RESEGREGATION OF BARBIERIA FROM CLITORIA (LEGUMINOSAE: PHASEOLEAE: CLITORIINAE) PAUL R. FANTZ Department of Horticultural Science, Box 7609 Carolina State University Raleigh, NC 27695-7609, U.S.A. ABSTRACT Barbieria is a monotypic genus that was transferred to subtribe Clitoriinae and included within the genus Clitoria in 1981 by Lackey. The morphology of Barbieria is compared with Clitoria and contrasted with genera Centrosema, Periandra and Clitoriopsis. Barbieria 1s found to be an abberrant member of C/#toria with characters supporting its recognition as a separate genus in the Clitoriinae. A key to genera of subtribe Clitortinae and a taxonomic treatment of Barbieria is presented. RESUMEN Barbieria es un gén tipico que fue transferido a la subtribu Clitoriinae e incluido dentro del género Clie: por Lackey en 1981. La morfologia de Barbieria se compara con Clitoria y se contrasta con los géneros Centrosema, Periandra y Clamp. Barbieri ha resultado ser un miembro aberrante de Clitoria con caracteres que corroboran su reconocimiento como un género separado en Clitoriinae. Se presenta una clave de los géneros de la subtribu Clitoriinae y el tratamiento taxonémico del género Barbieria. Barbieria DC. is a monotypic genus of questionable taxonomic affinities, having been placed historically into several tribes including its own. Lackey (1981) assigned Barbieria to subtribe Clitoriinae Benth. (Leguminosae Juss., Phaseoleae DC.), and concluded that it was a member of the genus C/ztoria L. Mabberley (1987) and Smith and Lewis (1991) recently reconfirmed Barbieria as a synonym of Clitoria. My preparation of manuscripts for the genera Clitoria and Centrosema (DC) Benth. for Douglas Stevens’ Flora de Nicaragua Project, Julian Steyermark’s Flora of Venezuelan Guayana Project, and the Flora Mesoamerica Project required examination of specimens of Barbierta throughout its range. I strongly disagree with Lackey’s assignment of Barbieria to Clitoria. Barbieria has a number of characters that make it an abberant member when placed within C/itoria. Proponents of Lackey regarded the morpho- logical differences as minor, due to the adaptation of Barbieria from bee to bird pollination. The objective of this paper is to compare the morphologi- cal characters of Barbieria with Clitoria, as well as contrast differences with other genera to support its recognition as a distinct genus within the Clitoriinae. Sipa 17(1): 55-68. 1996 56 Sipa 17(1) SUBTRIBE CLITORINAE Lackey (1981) recognized four genera in Phaseoleae subtribe Clitoriinae: Centrosema (DC.) Benth., Pertandra Benth., Clitoria L. and Clitoriopsis ilczek. He saatacietize the subtribe by the resupinate flowers, naked calyx interior, and prescence of bracteoles and hooked [micro-uncinate] trichomes. Leaves were reported to be 3- or 1-foliolate, to S—9-foliolate in a few members of Clitoria and Centrosema. These four genera also are similar in other respects. They are woody genera with members being trees, shrubs to shrublets, lianas, or suffrutescent herbs (= subshrubs of some authors). The more advanced species have an underground xylopodium from which arises one or more aerial stems an- nually during the rainy season. These stems produce leaves, flowers and fruits, then die back to near ground level during the dry season. Collectors frequently only voucher the aerial stems, leading some early and modern authors to treat these plants (and genera) as herbaceous or even as annuals. Leaves are pinnately compound with 3-foliolate leaves being prevalent, 3—7-foliolate leaves being infrequent (5 species) and 7—11-foliolate leaves being rare, occurring only in Cl/itoria lasciva Boj. ex Benth. Stipules and stipels are persistent and striate. Inflorescences are pseudoracemose, often axillary, multiflowered, bearing flowers on paired pedicels, or reduced to 1—2 apical flowers. Bracts are persistent and striate. Bracteoles persist and commonly are appressed to the calyx. Flowers are resupinate and large in size, with smaller flowers being 2.5-4 cm long. The calyx has five teeth with the upper two (dorsal) subconnate and the vexillary cooth commonly longer. Petals are clawed with the vexillum large, showy, obovate to subor- bicular, prominately veined and complicate. The vexillary stamen is free to more or less connate basally; the staminal sheath is incurved; anthers are yellow. The ovary is densely pubescent, subsessile, with a 1-2 mm gyno- phore or prominately stipitate; the style is incurved. Fruits are linear, straight to falcate, short-stipitate (subsessile) to prominately stipitate, beaked, with dehiscence by spiral twisting of the valves. Pubescence consists of macro- trichomes easily viewed at 1OX and micro-uncinate trichomes hidden be- neath and more easily viewed at 30X. The four recognized genera can be segregated on several morphological features of the flowers and fruit (Table 1). The calyx is campanulate (Centrosema, Periandra) or infundibular (C/itoria, Clitoriopsis). The lengths of the vexillum, wing and keel petals vary, but che keel petal is shortest in all genera. Anthers are uniform or rarely dimorphic (C/itoriopsis). Curva- ture of the style varies, as does the degree of dilation near the stigma and its pubescence. Fruits are ecostate or costate, flat or turgid, and if turgid, vary in degrees of depression between seeds. Fanrz, Resegregation of Barbieria Taste 1. Comparison of genera in subtribe Clitoriinae. Character Centrosema Periandra Clitoria Clitoriopsis CALYX Campanulate Campanulate Infundibular Narrow infundibular PETALS Vexillary spur Present Absent Absent Absent Present Present ese Auricled wing resent t Wing vs vexillum = Much shorter Much shorter Much shorter Slightly shorter Keel vs vexillum Much shorter Much shorter Much shorter Slightly shorter Keel vs wing Slightly shorter Slightly shorter Much shorter Subequal GYNOECIUM Ovary Subsessile subsessile Stipitate or Stipitate few subsessile Style curvature Broad U-shaped — Broad J-shaped Geniculate Weakly incurved Style pubescence Barbellate Glabrous Barbellate Glabrous basally apically ANDROECIUM nthers Uniform Uniform Uniform Dimorphic Fruit Costa 2 per valve Absent 1 per valve Absent or absent Valves Flat Flat Flat or turgid Flat Depression between seeds Absent Absent Absent Absent to present LEAVES Leaflet number (1)3 (5-7) (1)3 (1)3 (S-11) 3 THE GENUS CLITORIA Species of C/itoria fall into three distinct groups based upon morphol- ogy. Fantz (1979) reported that botanists have traditionally treated these groups as sections. However, one could argue that these groups represent three distinct genera due to morphological differences in the calyces and fruits supported by leaflet number and habit. However, all three groups are similar in floral characters that are unique within the papilionaceous le- gumes. Flowers are resupinate, large and showy, with stalked ovaries bear- ing geniculate, bearded styles, within infundibular calyces that persist in fruit. Bentham (1858) concluded that C/itoria would be rendered more natural by retaining these groups together than by segregation, and treated them as sections. Baker (1879) elevated two sections to subgenus C/itorta and subgenus Newrocarpum (Desv.) Bak. Fantz (1979) agreed with Bentham and elevated the third section to subgenus Bractearia (Mart. ex Benth.) Fantz. 58 Sipa 17(1) MORPHOLOGY OF BARBIERIA Lackey (1981) did not indicate affinities of Barbierza within the genus Clitoria. Barbieria consists of one species (a liana) with a wide distribution, occurring in the western Antilles and southern Mexico south to Peru, east to Brazil. Botanists often note Barbieria as rare to infrequently observed in the localities where the species is collected. The morphology of Barbieria will be reviewed in the following discussions with comparisons made to Clitoria and to other genera of Clitoriinae to determine potential affinities. Inflorescence.—Similarities to the Clitoriinae include axillary pseudo- racemes, persistent bracts and bracteoles, paired flowers, and micro-unci- nate trichomes. Inflorescences in Barbieria are elongate, slender, and multi- flowered. The closest placement in C/itoria would be subg. Bractearia. However, inflorescences of Bractearia are nodose, and often are borne cauliflorously in lianaceous species before leaves and axillary inflorescences are produced, characters not noted in Barbieria. Bracts of Clitoria are in three series. The inner pair is smallest, caducous to deciduous, located be- tween the pedicels; the middle pair is largest, persistent, located opposite the pedicels; and the outer pair are deciduous to semipersistent, nearly as large as the middle pair, and located between the pedicels. Barbieria ap- pears to have bracts only in two series. An inner row of bracts has not been — observed. Bracts and bracteoles (Fig. 1 F) of Barbieria are acuminate to subu- late apically, unlike any C/itorza or other genus in the Clitoriinae. Bracteoles of Bractearia are described by collectors as dark greenish-yellow to strong reddish-orange. Bracteoles of C/itoria, when noted by collectors, are de- scribed always as shades of green. Flowers. —Similarities to the Clitoriinae include the large, showy, resu- pinate flowers with clawed petals and complicated banner petal. Mature flowers of Barbieria appear dissimilar to those of Clitoria (Fig. 1A), with the banner petal spreading downward away from the conspicuously pro- truding keel. C/toria has the banner broadly U-shaped around the keel, the latter inconspicuous and somewhat hidden by the flaring apices of the wing petals. Color is commonly associated with the standard, with Barbieria described as scarlet or vivid to brilliant red, a color not associated with Clitoria, but is associated with some Periandra. Nectar guides are incon- spicuous. Flowers of C/itoria bear a standard with blue to lilaceous to viola- ceous pigmentation, or white, with numerous conspicuous veins of darker pink to violet hues commonly from a yellowish to whitish medial patch. Proponents of Lackey noted that C. javitensis (H.B.K.) Benth. has light red flowers, but the floral pigmentation is violaceous-pink, not red. Wings and keels of C/:toria are described as pale or off-white with the wings bear- ing violaceous pigmentation near the apex of the flaring blade. Fantz, Resegregation of Barbieria ao Sa Fic. 1. A comparison of some morphological features of Barbieria (left) with Clitoria a A: Flower, two views. B: Standard or banner (vexillum) petal. C: Keel henna) petal. D: Wing (alae) ac E: Calyx, intact and split-open views. F: Bracteole(s). G Gynoecium. H: Androecium. J: Legume; Clitoria subg. Newrocarpum represented. Calyx.—The calyx (Fig. LE) of Barbieria is narrowly infundibular mak- ing it similar to Clitoriopsis, not Clitoria. The calyx tube of Barbieria is elongated, appearing somewhat cylindrical with a length/width ratio of 4— 5, and is 20-veined. A prominent raised vein extends into each lobe with three smaller veins in between, one extending to the sinus, and the other 60 SIpA 17(1) pair into each adjacent lobe. Calyx lobes are deltoid-ovate basally with elongated subulate apices. The upper dorsal pair are free nearly to the base. The calyx is described as dark to vivid to blood red in color. Barbieria is unique within the Clitoriinae in bearing the upper calyx lobes free and bearing subulate apices. The calyx of C/itoria is infundibular. The calyx tube does not appear elongated, as the length/width ratio is 2-3. The veination pattern is stri- ated in subg. Bractearia and 10-veined in the other two subgenera, with a prominate vein into each lobe and into each sinus. Calyx lobes are acute to short-acuminate, not subulate apically. The upper dorsal pair is subconnate. The calyx, when noted by collectors, is described as green to purplish-tinged. Petals. —Barbieria has an elongated oblong-oblanceolate standard (Fig. 1B) with the keel length subequal the standard (as in C/itor‘opsis) and wings much shorter than the keel. The shape of the standard and the wings being much shorter than the keel are unique characters within the Clitoriinae. The standard is broadly ovate to suborbicular in all other genera of the Clitoriinae. The wings are subequal to slightly longer than the keel in the other genera, except for C/itoria, where the wings are conspicuously much longer with flaring apices. The keel blade (Fig. 1C) in Barbieria is narrow elliptic-oblong, acute, and weakly incurved. The keel blade in C/itoria is broadly oblong, broadly acute to subobtuse, and strongly falcate. The wing petals (Fig. 1D) have nearly straight, oblong blades in Barbieria. The wings of Clitoria have strongly falcate, oblong-spatulate blades. Gynoecium.—Barbieria has a sessile ovary (Fig. 1G) with a gynophore lacking, to 0.5 mm long. The style is linear, not dilated above, barbellate much of its length, and scarcely geniculate just below the stigma. This gynoecium is unique within the Clitoriinae as other genera bear a gyno- phore below the ovary and have an incurved style dilated above. Clitoria has a stalked ovary (rarely subsessile) with a style distinctly geniculate, with the style dilated and bearded above the abrupt incurved point. A sub- sessile ovary (gynophore 1—4 mm long) occurs in Centrosema and Periandra. Androecium.—The vexillary stamen in Barbieria (Fig. 1H) is free at the base with the others connate, similar to C/itoriopsis. The nine connate sta- mens bear anthers on elongated filiform filaments as in C/itoriopsis and Periandva. Barbieria is unique in bearing white anthers (yellow in other genera). The vexillary stamen in C//toria is more or less connate at the base. Nine stamens are connate with short filaments bearing yellow anthers. Legume.—The legume (Fig. lJ) of Barbieria is sessile, linear, ecostate, and enclosed at the base by the persistent calyx. The valves are strongly- transversely impressed between the seeds, turgid and convex around the seeds. The style scarcely persists as a beak. Legumes are variable in C/itoria, Fantz, Resegregation of Barbieria 61 but stipitate and bearing a beak. Legumes are flat in subgenera Bractearia and Clitoria, and other genera of subtribe Clitoriinae. Legumes of subgenus Neurocarpum are turgid, convex around the seeds, and ecostate to costate with a medial raised vein on the valve. Seeds. —Seeds of Barbieria are viscid, a character found elsewhere only in Clitoria subgenus Newrocarpum. The other subgenera of C/itorza and genera of Clitoriinae have smooth seeds. Leaves. —Leaves of Barbieria are similar to Clitoriinae by being odd-pin- nately compound, petiolate, bearing persistent stipules and stipels and micro-uncinate trichomes beneath macrotrichomes. Leaflets are commonly 15-21, oblong and obtuse. The only close match in leaflet number 1s C. lasciva Bojer ex Benth. (subg. C/itoria), a Madagascarean endemic with 7— 11-fololiate leaves. Lackey (1981) omitted reporting this higher leaflet number in the characterization of the subtribe and commented that C/ztoria subg. Clitoria {reported as sect. Ternatea} is more reminiscent of some Galegeae. CONCLUSION Clitoria isa genus with morphological variation in the habit, leaflet num- ber, calyx and fruits. However, flowers are similar in structure in all sub- genera, and unique within the papilionaceous legumes. Flowers are resupi- nate, large and showy, with stalked ovaries bearing geniculate, bearded styles, within broad infundibular calyces that persist in fruit. Barbieria becomes an aberrant member of genus C/itoria when placed in any subgenus. Clitoria subgenus Bractearia can be segregated from Barbieria by the nodose pseudoracemes, flat prominately stipitate fruits of much larger size, multistriated calyx, and crifoliolate leaves. C/itoria subgenus C/itoria can be segregated from Barbieria by the flat, short-stipitate fruits, 10-veined calyx, and suffrutescent habit. C/itoria subgenus Newrocarpum can be segre- gated from Barbieria by the trifoliolate and unifoliate leaves, 10-veined calyx, stipitate fruits and non-lianaceous habit. Should Barbieria be placed in a separate subgenus within C/storza? Not in my view. Table 2 contrasts these two genera. Barbieria can be segregated from all species of Clitoria by the scarlet color of the flowers, the oblong shape of the standard blade borne on a long claw, the length of the various petals with the wing petals being the shortest and the keel subequal the standard, the standard spreading away from the keel at maturity, the sessile ovaries and fruits with inconspicuous beaks, the free calyx lobes on a nar- rower, elongated tube (length 4—5 times longer than wide) with 20 veins, the subulate apices of bracteoles, bracts, stipules, and calyx lobes, the bracts oriented in two series, the free vexillary stamen, white anthers, elongate free filaments, a slender non-dilated style barbellate lengthwise, and higher leaflet number. — 62 Sipa 17(1) TABLE 2. Morphological comparison of Barbieria and Clitoria. Character Barbieria Clitoria Habit Liana Tree, Shrub, Liana Suffrucescent Herb LEAVES Leaflet Number (9)13—25 (1)3 (rarely 5-11) Stipule apex Subulate Acute INFLORESCENCE Bract series 2 rows 3 rows 3$ract apex Subulate Acute to obtuse Bracteole apex Subulate Acute to obtuse Bracteole color Greenish-yellow/ sreen reddish-orange Flower color ed White or blue to violaceous, rarely pink Vexillum (vs Keel) Spreading Surrounding keel CAL Tube configuration Narrow infundibular Broad infundibular Tube L/W ratio 4-5 2-3 Tube veination 20-veined L0-veined or striate Tube as Dark red Green to purplish tinged Lobe shape Broad, subulate Deltoic Fusion upper 2 lobes Nearly free Subconnate COROLLA Standard shape Oblong-oblanceolate Obovate Standard claw Long-clawed Short-clawed Wing shape Oblong Falcate-oblong Keel (vs wings) Longer Shorter Keel (vs standard) Subequal Much shorter GYNOECIUM vat Sessile Stalked Style curvature Weakly incurved Geniculate Style oe Barbellate length Barbellate apically Style 2 Not enlarged Dilated ANDROECIUM Vexillary stamen Free to base Connate near base Free Filaments Elongate, filiform Short Anther. White Yellow LEGUME Atrachment Sessile Stipitate Surface Turgid around seeds Flat or turgid around seeds Beak Inconspicuous Prominent Shape Subreniform Lenticular or subreniform Surface Viscid Smooth or viscid Barbieria has several morphological traits that are unique within the Clitoriinae. These include the imparipinnate leaves with a high leaflet num- Fantz, Resegregation of Barbieria 63 ber, subulate-acuminate apices of the stipules, bracts, bracteoles, and calyx lobes, the calyx with the upper two teeth free to near the base, the wings shorter than the other petals, the oblong-oblanceolate standard, and the sessile ovaries and fruits. These characteristics make Barbieria an aberrant member also from other genera of the Clitoriinae, and support its recogni- tion as a separate genus. Barbieria does have a lot of similarities with members of the Clitoriinae, including persistent stipules and stipels, appressed bracteoles, pseudo- racemes with flowers paired at nodes, resupinate flowers with clawed pet- als, naked calyx interior, and the constant prescence of micro-uncinate tri- chomes underlaying macrotrichomes. It would appear that the placement of Barbieria within the Clitoriinae by Lackey (1981) has merit. KEY TO GENERA AND SUBGENERA IN THE CLITORIINAE 1. Calyx with dorsal lobes free to near base; standard elongate-oblong; wing petals shorter than the keel; bracts, bracteoles, stipules and calyx lobes subu- late-acuminate; anthers white; leaflets 15-21; fruits turgid, strongly de- pressed between the seeds and subsessile; seeds vis Barbieria 1. Calyx with dorsal lobes connate below; standard broaally ovate to subor- bicular; wing petals subequal to longer than the keel petals; bracts, bracteoles, stipules and calyx lobes acute to obtuse, non subulate-acuminate, anthers yellow; leaflets 3 or 1, rarely 5—11; fruits flat or prominately stipitate when depressed between the seeds; Pere smooth or occasionally viscid (Clitoria subg. Neurocarpum). 2. Keel and wing petals subequal, each slightly shorter than the vexillum; anthers dimorphic, five aie others basifixed on longer filaments; style weakly incurved from the the base; calyx narrow infundibular (length ca 4—5 times the width a Clitoriopsis . Keel shorter than wing petals, each much shorter than the vexillum, an- thers uniform; style strongly incurved, broadly U-s ee or J-L-shaped near the aes pac broadly infundibular to paces 3. Cal N panulate; keel petals slightly shorter than wing petals; style strongly, medualle eunved inward, broadly’ U-shaped or J-shaped; fruit and ovary subsessile. 4. Style glabrous; legume valves ecostate; vexillary spur absent. ......... Periandra 4. Style pubescent, barbellate basally, less so above; legume valve co- tate, with a prominent nerve or wing near the suture; vexillum spurred. Centrosema 3. Calyx broadly infundibul ar (length ca 2—3 times width); keel petals much shorter than the wing petals; style geniculate (L-shaped), abruptly incurved, fruit a es ovary stipitate, rarely subsessile. Clitoria 5. Calyx subcoriaceous, striated; seeds large (7-16 mm diam.), lenti- cular; stipe (12—33 mm), staminal sheath (25-40 mm), ovary (10-20 mm), and petiolule (4-10 mm) elongated; inflorescence lignose, nodose, several to multi-flowered; leaflets larger (8—20 long Subg. Bractearia 64 Sipa 1701) 5. Calyx cartilaginous to sub eel 10-veined; seeds small (4 x 8 mm), subreniform to ovoid: se (1-14 mm), staminal sheath setiolule (1—5 mm, rarely 6— ~~ fa ao (1¢ 0-23 mm), ovary (S—9 mm y flore scen ublignose, usually | —2- flowered, 8 mm) shortened; it rare rely few- aL flowered; lanes smaller (1-8 long x 1-4 ¢ e). 6. cannes flac, pbsrestle (stipe 1-4 mm), ecostate; ee aces gy uby “rmin 1 fea shrinking in fri coca absent Tesfler —11, rarely 3 ar Le ease anaes ae Clitoria 6. Legume turgid, convex ce seeds, stipitate (stipe 4-14 mm), costate or ecostate; seeds viscid, germination hypogeal; calyx per- sistent in size in fruit, subcarcilaginous; cleistogamy present or absent; leaflets 3, rarely 3 and 1, or unifoliate. ....... Subg. Neurocarpum TAXONOMIC TREATMENT OF BARBIERIA BARBIERIA DC.,, Prodr. 2:234. 1825. Type species: B. pinnata (Pers.) Baill. Shrub, erect or apically scandent or trailing. Leaves alternate, impari- pinnate, petiolate, stipulate, stipellate; leaflets numerous, opposite, entire; petiolules quadrate; stipules persistent, striate, subulate-acuminate; stipels persistent, 1-nerved, linear-subulate. Inflorescence axillary or terminal, pseudoracemose, few-flowered, long-pedunculate; pedicels paired at nodes. Bracts 4, striate, lance-subulate, pubescent adaxially; the inner pair often persistant, opposite and appressed to the pedicel; the outer pair narrower, between the pedicels, deciduous. Bracteoles paired at calyx base, lanceolate, subulate-acuminate, striate, pubescent adaxially. Flowers papilionaceous, resupinate, showy, red. Calyx narrowly infundibular, persistent in fruit, 5- lobed, lobes free, deltate to rapidly subulate above, subequal with ventral lobe slightly longer. Vexillum complicate, long-clawed, spurless, oblong- oblanceolate; wings oblong, long-clawed, much shorter than the keel; keel long-clawed, elliptic-oblong, subequal to the vexillum. Stamens diadelp- hous, often persistent in fruit; vexillary stamen free nearly to base. Pistil enclosed within staminal sheath; ovary sessile, style linear, densely barbate, weakly incurved, weakly geniculate beneath stigma. Legume sessile, lin- ear, straight, ecostate, enclosed at base by persistent calyx; valves strongly transverse-impressed between the seeds, spirally twisting upon dehiscence. Seeds smooth, transverse-oblong. (x= 10). A monotypic genus Sideeecad in the neotropics, but apparently rare locally since collections in herbaria are comparatively few. Be pinnata (Pers.) Baill., Hise. Pl. 2:263. 1870. Sepa ca _ PL. 2:302. 1807. Clrtaria ie (Pers.) R H. Smith & G.P. Lewis, Kew Bull. Hee 991. Type: PUERTO RICO, M. Ledr son. (HOLOTYPE: L: ISOTYPE: < oo oh dla Poir., Lam. Encycl. Suppl. 2:300. 1811. Barbieria ae (Poir.) m. Légum. 242. 1825. Type: PUERTO RICO, M. Ledra s.n FANtz, Resegregation of Barbieria 65 Barbieria maynensis Poepp. & Endl., Nov. Gen. & Spec. 3:58, t.245. 1845. Type: Peru (orientalis). Crescit in sylvis primaevis ad flumen Huallaga versus Tocache et Yurimaguas, Julio ad Decembrem floret, sve coll. & no. {[Poeppig coll.?] Shrub 1—2.5 m, or a woody vine scandent 9-12 m or trailing; branches striate-terete, 1-6 mm diam., densely hirsute, trichomes to 2 mm long, reddish-brown becoming whitish with age; pubescence of micro-uncinate (hooked) trichomes common (vidi 20-30) beneath macrotrichomes; pith continuous. Leaves imparipinnate; leaflets (9)13-—21(25), oblong to ellip- tic-oblong to ovate-oblong, broadly acute to obtuse, mucronate (to | mm long), (1.5)2.5-6 cm long, 1-2.5 cm wide, upper surface bright to dark green, micro-uncinate pubescent (beneath sparse strigose macrotrichomes when young), lower surface pale, whitish to bluish-green, moderately to densely appressed-pilose; petioles 1-4 cm long, densely rufo-pilose; rachis quadrangular-terete, sulcate above, 5—16 cm long, internodes 0.5—2.5 cm long, micro-uncinate beneath rufo-pilose trichomes; petiolules rugose, dark- colored, rufo-pilose, 2 mm long; stipules narrowly deltate-lanceolate, 5—8- striate, subulate-acuminate, 7—12 mm long, 1.2—2 mm wide, pubescence micro-uncinate; stipels linear-subulate, 1-striate, (1.5)3—5 mm long, ca (.2—-0.8 mm wide, pubescence micro-uncinate. Inflorescence 4—16(24) cm long, flowering toward apex; peduncle 4-11 cm long, rufo-pilose; rachis internodes 1—3.5 cm long; pedicels 3-5 mm long in flower, 5-7 mm long in fruit. Bracts deltate-ovate, acute, 3-8 mm long, 0.5—1 mm wide, pubes- cence micro-uncinate. Bracteoles lanceolate, subulate-acuminate, dark green- ish-yellow becoming strong reddish-orange, 7-11 mm long, 2—3 mm wide, pubescence ciliate and micro-uncinate. Flowers 4.5—6.0 cm long, red. Ca- lyx dark, vivid red; tube 16-22 mm long, 3—5 mm wide at base expanding to 6-8 mm wide at throat, venation 20-striate, one vein extending into each lobe to its apex, one vein extending to each sinus, and three less con- spicuous veins between these two; lobes deltate-ovate, subulate-acuminate, dorsal and lateral lobes 8-11 mm long, to 2 mm wide basally, ventral lobe 10-12 mm long. Standard brilliant to blood red, paler toward spotted cen- ter, blade oblong-oblanceolate, 25—33 mm long, 14-17 mm wide; claw 21-24 mm long, 3—5 mm wide; wings red, blade oblong, flaring apically, 9-13 mm long, 3.5—G mm wide; claw 26-33 mm; keel red, blade elliptic- oblong, weakly falcate, 17-23 mm long, 2.5—5 mm wide; claw 26-34 mm long. Staminal sheath white, 36-44 mm long, free filaments filiform, (4)6— 8 mm long; anthers white, lanceolate, 0.9—1.2 mm long. Gynophore lack- ing or to 0.5 mm long; ovary linear, 8-11 mm long, 1-1.2 mm wide, pubescence dense, trichomes white, to 2 mm long, ascending-appressed; style (29)38-43 mm long, exserted beyond stamens, flattened, bearded lengthwise, geniculate 5-6 mm from distal end; stigma capitate. Legume 66 Sipa L7(1) pale green becoming brown, subsessile, linear, straight, ecostate, enclosed at base by persistent calyx; stipe 0.5—1 mm long; valves puberulent-hir- sute, strongly transverse-impressed between the seeds, 36-55 mm long, 4—6 mm wide, spirally twisting upon dehiscence 0.5—1.5 turns; beak in- conspicuous, 1-2 mm long. oe i per pod, dull dark brown to black, viscid, transverse-oblong, 2.2— 4-6 X 0.6-1.5 mm. (x = 10). Iustrations.—De Candolle ( 8 oe tab. 39), Poeppig (1845, plate 264), and White (1980, fig. 6) provided excellent line drawings of vegetative and reproductive structures. Vernacular names. Peru: puspo poroto huasca (Vigo 8307), pusporota huasca (Schunke 12412). Economic uses.—"“One of the best ground cover crops” (Schipp 1108). Ex- cellent potential as a climbing ornamental. Phenology.—Plant vouchers have been collected with flowers (March) and fruits from April chrough December. Habitat —Moist soils in secondary growth, roadsides, riverine forests, forests edges or open areas with abundant sun. Elevation 390-1000 m. Distribution.—Southern Mexico south to Peru and west to Venezuela and Brazil, and in the Caribbean. Collectors often noted that the plant rarely was observed in the area sampled. Specimens examined [“g-"= gift specimen from institution cited}: CENTRAL AMERICA. BELIZE (BRITISH HONDURAS). Rio Grande, Schipp 1108 (F,MICH,MO,NY). sTaNN CREEK: 17 mi section, Stann Creek alley. Gentle 9255 (ER.LL,MEXU, MICH,NY). COSTA RICA. Larkowski 1261 (FP). PUNTARENAS: Helechales, Potrero Gr ande, Buenos Aires, Gomez- pele 10839 (BF); Cabugra, Buenos Aires, Ocampo CR ~ 2424 (CR); vic. Buenos Aires, Pittier 4714 (P); Terruba, Buenos Aires, Poveda 869 (EF): Buenos — de Puntarenas, Zamora et a Hee F); Buenas Aires de Puntarenas, Zamora LI86(C . SAN JOSE: Helechales de San Isidro del General, Led 1025 (CR); vic. El Gen- eral, ml oe (MO, NY); Rio Convento near Panamerican Hwy, El General Valley, Willrams et al. 28704 (CR,ENY),; Valle de El General, Yer/ding 246 (F); San Isidro de El General, El Pilar de Cajén, Zamora et al. 1520 (CR). GUATEMALA. Atta Veraraz: be- tween Samanzana & C cua via Seboquil, Steyermark 45101 (EKNY);, Cubilquitz, Tuerckhemm 7845 (F,M,MO,NY,P). MEXICO. Rio Piedras, Hroram s.r. (P). Nayarit: Sierra San Pedro Nolasco, Jurgensen 780) (MO). Oaxaca: Galeott7 3431 (P); Comaltepec, Liebmann 54 (F). CARIBBEAN. CUBA. i cigdes aie > (MO). IsLe OF PINES: vic. ae Pedro, Britton et al. 15105 (ER MO). DOMINICAN REPUBLIC. Wright et a/. 89 (B). La VeGa: vic. Piedra Blanca, Allard 13483 (MO); vic. aa Blanca, Allard 17062 o Cordillera Central, Jarabacoa toward Buenavista, Ekman 14189 (MO). Monte Cristi: Leonor, dist. Sabaneta, Valeur 76 (F-2 sheets, MO). SAN CrisTosaL. near El] Cacao, San Cristébal, Liogier 17767 (F). SANTO DomINGO: vic. Ciudad Trujillo, A/ard 13338 (US); Cordillera Central, Vil se Ekman 11179(P). HAITI. Artiponite: vic. Kalacroix, section Dessalines. Leonard OO (PF). Norpb: trail to Citadel in vic. Dondon, Leonard 8642 (MO). PUERTO RICO. ioe ada ad Piedra Blanca, Sitenis 5710 (FM GUADILLA. La Juanita near Las Marfas, Britton et al. 3939 (EMO,US); vic. Maricao, Britton & Cowell 4092 (F,US). HumMAcAo. PR 966 near km 4.7, E of El Verde, Sierra de Luquillo, Burch 3203 (MO,TENN); Lares ad i=) FANTZ, Resegregation of Barbieria 67 Palma Llanos, Sintenis 5942 (F.MO). MayaGtiez. near Mayagiiez, Holm 27 (MO). SAN JUAN. Rio Piedras, Otero 325 (MO). S H AMERICA. BOLIVIA. Guanai, Tipuani, Bang 1363 (F.MO-2 sheets,US); Rio Junsay (?), Kuntze s.n. (FUS); Guanai, Rusby 2356 (F,.MO-2 sheets). BRAZIL. AMazonas: Sao Paulo de Olivenga, Ducke 565 (F.MQO); Rio SolimGes, beside Estrada Bom Fim, Sao Paulo de Olivenga, Lleras et al. P17342 (MO); Lago de Tefé, opposite Tefé, Léeras et al. P17490 (MO); Mun. Humaitd, estrada Humaitd-Pé6rto Velho km 38, Tezxera et al. 270 (MO,US). Goids: ca. 17 km S of Goids Velho, 6 km NE of Mossamedes, Serra Dourada, Anderson a eens Marto Grosso: Santa Anna de Chapada (= dos Guimaraes), Regnel/ 3316a (F). Parad: Mun. Itaituba, estrada Santarém-Cuiabaé, BR-163 km 842, Serra do Cachimbo, ie 250 (MO). Ronponia: Mun. de Colorado de One. BR 364, Pérto Velho- aes km 20, Cid et al. 4367 (MO,US). COLOMBIA. Antioquia: Mpio. San Rafael, 3.1 of San Rafael along Guatope-San Ratael Rd, Brant & Roldan 1514 (g-MO). Boyaca: i oA. 130 m NW of Bogota, Lawrence 488 (F.MO,US). Meta: Los Llanos, Villavicencio, ee 2457 (F). Vaupks: Cano Grande y San José del Guaviare, Cuatrecasas 7378 (F); Mita & vic., Urania, Zarucchi 2249 (MO). PERU. Amazonas: Mirana, Woytkowski 5648 (MO). Cuzco: Paucartambo, Pileopata, Izguierda, Var. on 11642 (MO); Casnipala, Weberbauer 6951 (F,F-frag.). HUANUCO: Monzén, Woytkowski 5029 (MO). JUNIN: La Merced, Killip & Smith 2378 (P); Rio Paucartambo Valley near Perené a Killip & Smith 25367 (F). Loreto: ie Coronel Portillo, Bosque Nac. de van Humboldt, Ucayali, km 86 Pucallpa-Tingo Maria rd, Gentry 6 Siluz zar 29440 (MO); enmens Rio Huallaga, Killip & Smith 27941 (F); pe ee Klug 2978 (F,MO); Balsapuerto, K/wg 3095 (F,MO); alto Amazonas, La- gunas, trail to Argentina, McDaniel G Rimachi 16433 (F,MO); alto Amazonas, Yurimaguas, Carretera del Caserio de Munich, Rimachi 3013 (E, MO,TENN); 3 km S of Yurimaguas, Straw 2416 (US). Pasco: Oxapampa prov., Serrania de San Matias E of Loma Linda, Gentry et al. 42000 (g-MO); prov. Oxapampa, Iscozacin, Smith 2080 (DUKE,MO). San Martin: W of Huicte, Schunke 6524 (F). SAN MARTIN (MARISCAL CACERES): Dtto Tocache Nuevo, Pucayacu nr Tocache Nuevo, Plowman & Schunke 7497 (F),; Dtto. Tocache Nuevo, Quebrada de Ishichimi (Fundo Retiro), Schunke 3913 (F.MO),; Quebrada de Huaquisha (Rio Huallaga), ‘igo 7155 (MO); Almendras, camino a Pueblo Viejo, Vigo 8186 (MO); Dtto. Tocache Nuevo, Fundo Jeroglifico, del Sr. Luis Ludena (Quebrada Ishichimi), Vigo 8307 (MQ); dtto. Tocache Nuevo, Quebrada Ishichimi, cerca al Fundo del San Luis Ludena, Schunke 12412 (F). VEN EZUELA. Barinas: 0.5 km NW of Barinitas, Steyermark & Wiehler 106605 (ER.MO). Botivar: Disc. Roscio, “El Abismo,” Rio El Samay, affluente Icabaru, Holst & Liesner 2365 (F.MO,NY); km 316 S of El Dorado, 2 km S of Santa Elena de Uairén, Steyermark 111328 (KUS-2,VEN). Sucre. Dist. Benitez, entre Rio Frié y Cafio Yaguaima, Ben/tez 2908 (MO), Dist. Benitez, Serranfa de la Paloma, 28 km SE of Ajies rd to Guariquen, Steyermark et al. 121405 (MO). EXSICCATAE Allard, H.A. 13338, sane, 17062 Ekman, E.L. 11179, 14189 Anderson, W.R. 10141 Galeotti, M. 3431 Bang, A.M. 1363 Gentle, PH. 9255 Benitez de Rojas, C.E. 2908 Gentry, A. & A. Salazar 29440 Brant, A.E. & FJ. Roldan 1514 Gentry, A. et al. 42000 Britton, N.L. et al. 3939, 15105 Goémez- ee oe 10839 Britton, N.L. & Cowell 4092 Haught, . Burch, D. 3203 Holm Cid, C.A. et al. 4367 Ficls, B. & = as 2365 Cuatrecasas, J. 7378 Jurgensen, C. 78 D ; 6 Killip, E.P. & ce Smith 23758, 25367, 68 27941 Klug, G. 2978, 3095 Larkowski, C. 1261 ae a 488 Leon, J. Leonard, . . 7900, 8642 Liebmann 4654 ee A.H. 17767 , E. et al. PL7342, P17490 Mil S.&M. Rim: achi Y. 16433 Ocampos, a 2424 tero, J.I Pictier, H. re 4 Plowman, T. & J. Schunke V. 7497 Regnell I:3316a Rimachi Y., M. 3013 Rusby, H.H. 2356 ls W.A., oie Schunke V., J. 3913, 12412 (= J. Schunke Silva, M ) 5 Sintenis, P. 5710, 5942 Sipa 17(1) SUCH, A, 2208 Smith, D. 2080 Steyermark, J.A. 45101, 111328 Steyermark, J.A. & H. es 106605 Steyermark, J. et al. Straw, R.M. 24 Teixeira et al. 2 Tuerckheim, ie von 7845 : 24, 7155, 8186, 8307 (= J. Stee Vigo?) We ae A. 6951 Willia Woytkowsl .O. et al. 28704 7 5029, 5648 Zamora, a et al Zanora V., N. et ‘i an 1186 See 224¢ ACKNOWLEDGMENTS Thanks and appreciation are extended to the curators of the following herbaria who provided the specimens cited in this paper: CR, DUKE, F, LL, M, MEXU, MICH, MO, NY, P, US, VEN. REFERENCES BAKER, J.G. India. L.Reeve & Co., L BentHamM, G.1858. Synopsis of the genus C/itoria. J. Linn. Soc., Bot. 2:33—4: Ds Canbo_ir, A.P. 1826. Mémoires sur la famille des ieeaniecns A. San 244, tab. 39. FANTZ, PR. 1979. (Leguminosae). a 8:90-94 Lackey, J 198 301-32 MABBERLY, 7 J. Porppic, E. | 59, plate 264 SMITH, R.H. auf G.P. Lewis. sere Papilionoideae). Kew Bull. 46:3 White, PS. Family 83. Leguminosae sObRumnaly Papilionoideae (conclusion). Gard. 67:553—555. 1879. Leguminosae, no. 76:C/itoria. In: J.D. Hooker, ed. Flora of British on => ore S241 aay notes and new sections of C/itoria subgenus Bractearia . Tribe 10. ede ae DC. (1825). In: R.M. Polhill and P.H. Raven, eds. Avance in legume systematics. Royal Botanic G Gardens, Kew, England. Part I 1987. The plant book. Cambridge Univ. Press, Cambridge. 1845. Nova genera ac species plantarum. Friderici Hofmeister, Leipzig. 3:58- A new combination in C/itoria (Leguminosae - 1980. Barbieria. In: J. D dwyer and collaborators. Flora of Panama, part V. Ann. Missouri Bot. ~ FUERTESIMALVA, A NEW GENUS OF NEOTROPICAL MALVACEAE PAUL A. FRYXELL Department of Botany University of Texas Austin, TX 78713, U.S.A. ABSTRACT Urocarpidium sect. Urocarpidium is shown to be synonymous with the genus Tarasa, and istinct. The latter is redescribed in generic rank Urocarpidinm sect. Anurum is generically c and given the name Fwertesimalva. Fourteen new combinations are made. RESUMEN Urocarpidium sect. Urocarpidinm es un sinénimo del género Tarasa, y iets sect. ribe en el rango Anurum es considerado como un género distinto. Este Ultimo se redes génerico con el nombre de Fuertesimalt ‘a. Se proponen catorce ae nuevas. The genus Urocarpidium was established by Ulbrich (1916) with the single species U. albiflorum Ulbrich. It was subsequently enlarged to include ten other species (Krapovickas 1954a), and ultimately a total of 14 species (Krapo- vickas 1970). The same number of species is recognized in the following enumeration, although with some nomenclatural changes (e.g., Fryxell & Krapovickas 1990; Krapovickas & Fryxell 1993) and the addition of more recently described species. Krapovickas (1954a) divided the genus into two sections, sect. Urocarpidium containing only the type species, and sect. Anurum Krapov., including the remaining species. This interpretation has been followed by most subsequent authors (e.g., Hutchinson 1967). The etymology of the generic name Urocarpidium is considered problem- atical inasmuch as Ulbrich did not state his intention in proposing it. A reconsideration of its derivation led to a reconsideration of its taxonomy. Fryxell (1988) cited the roots of the name as the German “ur (primitive) and the Latin “carpidium” from the Greek “karpos” (fruit) with a diminu- tive ending, thus meaning primitive mericarp. This is a plausible deriva- tion, if one considers the relatively unornamented fruits that are typical of most of the species. However, an alternative derivation (Fuertes 1989) 1s from the Greek “ouro” (tail) plus “carpidium,” meaning tailed (or aristate) mericarp. This interpretation was earlier rejected (Fryxell 1988) because the mericarps typical of these plants are not tailed or aristate. However, the question concerns the nature of the mericarps of the type species, U. albiflorum, the only species that Ulbrich knew when he described and named Sipa 17(1): 69-76. 1996 70 Sipa 17(1) the genus. The mericarp of this species was described by Ulbrich (1916) as “apice cauda longissima ad fere 8 mm longa ciliata munita” and was illus- trated by Krapovickas (1954a) from an isotype specimen (PERU, moun- tains near Chosica, A. Weberbauer 5326, isoryre: GH). A reconsideration of the mericarp morphology of U. a/biflorum makes it clear that this species has aristate, pubescent mericarps and thus belongs in the genus Tarasa Phil. (and incidentally that the derivation given by Fuertes is correct). In fact, the mericarps of U. albiflorum (cf. Krapovickas 1954a, fig. 2A) con- form closely to those of Tarasa operculata (Cav.) Krapov. (cf. Krapovickas 1954b, fig. 2A), and the former should perhaps be reduced to a synonym of the latter. In any case, Urocarpidium is to be treated as a synonym of Tarasa, and the remaining species, which are usually placed in Urocarpidium, form a natural genus that is lacking a name. It is the purpose of this note to describe the new genus and to supply a new name and make the necessary new combinations. Fuertesimalva Fryxell, gen. nov. Type: Fwertesimalva limensis (L.) Fryxell (= Malva limensis L.) sail Ulbr. sect. Anurum Krapov. Darwiniana 10:614. 1954. Lecrotryee, here esignated: Urocarpidinm limense (L.) Krapov. (= Matlva limensis L.). Herbae stellato-pubescentes, cymis axillaribus scorpioideis et corollis purpurascentibus et fructibus glabris, mericarpiis singularis plus minusve hippocrepiformibus, necnon muticis atque irregulariter transversim rugatis, rugis paucis. Annual or perennial herbs, ascending or erect, with stellate pubescence. Leaves petiolate, the blades ovate or orbicular, usually palmately lobed or parted, crenate or dentate, the upper surface sometimes with appressed simple hairs. Flowers sometimes solitary, usually in axillary scorpioid cymes; involucel of 2 or 3 filiform bractlets; calyx stellate-pubescent, (4—) 5-lobed; corolla purplish (sometimes white), shorter than to slightly longer than the calyx; androecium included, the column glabrous or pubescent, filamentiferous at apex, the anthers sometimes purple, few (sometimes only 5); styles 5-16, the stigmas capitate. Fruits schizocarpic, oblate, glabrous; mericarps 5—16, indehiscent, horseshoe-shaped, with irregular transverse jacent eon ridges (these sometimes interlocking in the fruit between ac mericarps), sometimes with a small endoglossum; seeds solitary, glabrous. Fuertestmalva is an Andean genus (Argentina and Chile to Colombia and Venezuela), with two species (F. jacens and F. limensis) found disjunctly in Mexico, usually from relatively high elevations (generally 1000-3800 m), but from much lower elevations (100-600 m) for F. sanambrosiana and F. peruviana. Keys to the species are found in Krapovickas (1954a, 1970), and a revised key is presented below. Chromosome numbers have been reported for several species (summarized by Krapovickas 1967, Tables 1 and 2, pp. 29, 34), with counts of 2” = 10, 20, and 30. FRYXELL, Fuertesimalva, a new genus 71 Fuertesimalva pertains to the Sphaeralcea alliance, i.e. those genera char- acterized by a base chromosome number of x = 5. Within this group it is one of the few genera characterized by axillary scorpioid inflorescences, which include Tarasa and, to a more limited extent, the genera Montezroa and Sphaeralcea, the inflorescences of which conform to this pattern only imperfectly. The genera Fuertesimalva and Tarasa are very similar in leaf morphology and general aspect (as is evident from the plates published by Krapovickas 1954a, pl. 1-8), but are strikingly different in fruit morphol- ogy. The mericarps of Fvertesimalva are horseshoe-shaped, glabrous, and with characteristic transverse ridges or excrescences dorsally, in a pattern unlike that found in any other malvaceous genus. The mericarps of Tarasa, by contrast, are apically aristate, notably pubescent, and laterally reticu- late, showing a superficial similarity to mericarps of Sphaeralcea. In addi- tion, the two genera differ in pubescence characters, especially clearly pre- sented in the case of calyx pubescence. Fuvertesimalva pubescence 1s stellate (or sometimes with simple hairs) whereas that of Tarasa is commonly stipi- tate, with darkly pigmented stipes. The results of cpDNA analysis by La Duke & Doebley (1995) show that Urocarpidium and Tarasa are not closely allied, in spite of sharing a com- mon base chromosome number and a similar general aspect. KEY TO THE SPECIES OF FUERTESIMALVA —_ . Anthers 5(—7); flowers oo or in few-flowered axillary inflorescences. 2. Flowers aes solit 3. Carpels 5, the mericarps 1.5 x 1 mm; calyx and corolla pentamerous (Argentina). F. pentacocca 3. Carpels 7-10, the mericarps 1.3 1.3 mm; petals often 3, the calyx usually 4-lobed {cf. Eliasson 1970} (Galapagos). . insularis i) . Flowers solitary or in few-flowered axillary inflorescences; carpels ca. 8 the mericarps 5 x 1.5 mm or larger (Argentina). F. pentandra — . Anthers 10 or more; flowers generally in axillary scorpioid cymes. 4. Stipules S—10(-14) mm long, broadly ovate or falcate, ca. 5 mm wide; ie iage eatae carpels 12-15 aa ers sessile; calyx hirsute (Peru, Chile) F. chilensis z ee pedicellate, the pedicels 2—3(—8) mm long; calyx with 1 or 2 etae (1.5 mm long) at tip of each lobe, otherwise glabrous (Peru). ... F. sttpulata 4. Stipules 2-6 mm long, ee lanceolate, 0.5—2.5 mm wide; foliage gen- erally pubescent; carpels 7-16. 6. Petals 10 mm long; ca = 7-8 mm long, deeply divided, the lobes 6 x 2mm; — 8-10; leaves basally truncate, unlobed (Peru). ......... F. leptocalyx 6. Petals 2—4(—7) mm long; calyx 4—6 mm long, ca. half-divided, the lobes shorter and broader; ak 7-16; leaves often basally cordate, often palmately lobed. 7. Mericarps 3 x 3mm, ca. 10 (Peru). F. pennellii 7. Mericarps usually 2.5 x 2.5 mm or smaller (or if larger, then only 8), 8-1 Sipa 17(1) 8. Carpels 7-10; inflorescences 2—6-flowered. 9. Inflorescences 2—3-flowered; mericarps 4 acute (Peru). x 2.8 mm, apically F. corniculata 9. Inflorescences 2—6-flowered; mericarps 2.5 x 2.5 mm or smaller. 10. Mericarps 1.5 x 1.5 mm (Colombia, Ecuador). ...........00..... F. killipii 10. Mericarps 2 x 2 mm. 1. Inflorescences subequal to the subtending leaf (Peru, Bolivia, Argentina). F. echinata i Inflorescences generally shorcer than the subtending pet orca Carpels more el Or1 ‘ences often with LO flowers or more, often equalling or a the nde leaf 12. Mericarps lacking an endoglossum; plants of “etively phigh elevation (1400-3200 m) (Mexico, Venezuela, Colombia, Ec- uador, Per F. jacens nn = Loan| po) . liamensis 12. Mericarps we a small internal projection (endoglossum); plants of relatively low elevation (100—G600 m). 13. Mericz Ups black at mz LCUTILY, 1.5 —2 mm long (Peru, Chile). F, peruviana 13. Mericarps pale brown at maturity, 1.2—1.5 mm long (Chile: Islas Desaventuradas). F. sanambrosiana 1. Fuertesimalva chilensis (A. Braun & Bouché) se comb. nov. Malva chilensis A. Braun & Bouché, Ind. Sem. Hort. Berol. |. 1857. U) pidii hilense A. Braun & Bouché) Krapov., Darwiniana 10:619. 1954. Type: CHILE. eX = hore Berolinensis (B as photo F-93 22). Malva scorpioides Vurcz. Bull. Soc. ae Naturalistes Moscou 36:562. 1863. Ma/vastrum peruvianum Var. scorpioides (Turcz.) E. G. Baker, J. Bot. 29: i 1891. Type: PERU. Lima: valley of Lima, A. Matthews 1006 (worn K, OXF Matvastrum hinkleyorum I. M. Johnst., Contr. Gray F -n.s. ae 1924. Type: PERU. year Arequipa, 2100 m, Mar 1920, PoE, ae ei Irs, ie erates GH, IsoTYPE: US). — Distribution.—Peru and Chile; 2100-2800 m elevation (0O—3000 m fide Brako & Zarucchi 1993), [/lustrations.—Krapovickas (1954a, fig. 2F, pl. 3). 2. Fuertesimalva corniculata (Krapov.) Fryxell, comb. nov. Ty, td tt 2 Krapov., ee andia 3:67. 1970. Type: PERU. Lima: Matucana (Val lle ae ae = 100 Om, 3 Apr 1967, M. Chanco 25 (HoLorypr: CTES) Distribution.—Peru (Lima); 1000 m elevation (SO0—3000 m fide Brako & Zarucchi 1993). I/lustrations. —Krapovickas (1970, fig. 3A-B), 3. Fuertesimalva echinata (C. Presl) Fryxell, comb. nov. Matva echinata C. Presl, Relig. Haenk. 2:122. 1835. Urocarpidinm echinatin (GC. Pres)) Krapov. & Fryxell, Bonplandia 7:58. 1993. Typr: PERU. in Cordilleris Peruviae, T. Haenke sn. (LECTOTYPE, designated by Krapovickas & Fryxell, loc. cit.: PR: ISOTYPE: PR). 73 FRrYXELL, Fuertesimalva, a new genus Malvastrum shepardae 1. M. Johnst. Contr. Gray Herb. n.s. 70:75. 1924. Urocarpidinm shepardae (1. M. Johnst.) Krapov. Darwiniana 10:621. 1954. Type: PERU. Puno: vi- cinity of Lake Titicaca, Dec. 1919, R. S. Shepard 123 (HoLoTYPE: GH). Distribution.—Peru, Bolivia, Argentina (Salta, Tucuman); 2000-3800 m elevation (O—4500 m fide Brako & Zarucchi 1993) Illustrations. —Krapovickas (1954a, fig. 2E, pl. 4). 4, Fuertesimalva insularis (Kearney) Fryxell, comb. nov. eed insulare Kearney, Leafl. W. Bot. 6:167. 1952. Urocarpidium v. Darwiniana 10:631. 1954. Type: ECUADOR. GaLapaGos ne ee ianfe nae Island, summit of Tagus Cove Mountain, 26 May 1932, J. T. Howell 9570 (HOLOTYPE: CAS). Distribution.—Ecuador (Islas Galapagos); 1500-1600 m elevation (cf. Eliasson 1970). Ilustrations. —Bates (197 1, fig. 190), Eliasson (1970, fig. 2), Krapovickas (1954a, fig. 2L). 5. Fuertesimalva jacens (S. Watson) Fryxell, comb. nov. Malvastrum jacens 8. Watson, Proc. Amer. Acad. Arts 21:417. 1886. Urocarpidium gacens Watson) Krapov. Darwiniana 10:267. 1954. Type: MEXICO. CHIHUAHUA: Norogachic, 150 mi N of Batopilas, Aug—Nov 1885, E. Palmer 430 (Lectotype, des- ignated by Hill 1982: GH; isorypes: K, US). Malvastrum jacens var. palmatifidum Hochr., Annuaire Conserv. Jard. Bot. Genéve 6:3 1902. Urocarpidinm gad ate (Hoc i Krapov. Darwiniana 10:623. 1954. a PE: MEXICO. Cuimuanua: in Sierra Madre, 2 Oct 1988, C. G. Pringle 1574 (HOLOTYPE: G; 1soTypes: BM, BR, G, HO. Distribution. —Mexico (Chihuahua to Chiapas; see map in Fryxell 1988, fig. 112); 2300-3000 m elevation. [ustrations:—Fryxell (1988, fig. (1954a, fig. 2G-H) 6. Fuertesimalva killipii (Krapov.) Fryxell, comb. nov. Urocarpidium killipii Krapov., Darwiniana 10:626. 1954. Type: COLOMBIA. SANTANDER: vicinity of ie 1600-1800 m, 4-10 Jan 1927, E. P. Killip & A. C. Smith 16757 (HOLOTYPE: US; IsoTyPrs: G Y). Distribution. et (Santander) and Ecuador (Chimborazo, Imbabura, Pichincha, Tungurahua); 1600-2200 m elevation. Illustrations. —Krapovickas (1954a, fig. 2K, fig. 3G-I, pl. 6). 13; 1992a, fig. 25), Krapovickas an 7. Fuertesimalva leptocalyx (Krapov.) Fryxell, comb. nov. Urocarpidium leptocalyx Krapov., Kurtziana 2:118. 1965. Type: PERU. ANCAsH: prov. Santa Ana, 1700 m, 10 May 1960, A. A/za 11 (HOLOTYPE: Distribution. —Peru (Ancash, Cajamarca); 1700-2100 m elevation. IMustration.—Krapovickas (1965, fig. 3D-G). S 74 Sipa 17(1) / 8. Fuertesimalva limensis (L.) es comb. nov. Matva limensis L., Cent. Pl. 2:27. 1756; Amoen. Acad. 4:325. 1759. Mal/vastrum limense L.) Ball, J. a. Soc., Bot. 22:32. 1885. Malvastrum pernviannum var. limense (L.) E. G. Baker, J. Bot. 29:168. 1891. Urocarpidinm limense (L.) Krapov., Kurtziana 4:33. 1967. _ {t — Tyee: “limensis” (HOLOTYPE: N-870.6). Matlva costata C. Presl, Relig. Haenk. 2:123. 1835. Type: PERU. in collibus Cordillerum Peruviae, 7. Haenke s.n. (HOLOTYPE: PR). Distribution.—Mexico (Durango to Morelos and Veracruz; see map in Fryxell 1988, fig. 112), Venezuela, Colombia, Ecuador and Peru; 1400— 3500 m elevation (cf. Brako & Zarucchi 1993). [/lustrations. —Fryxell (1992a, fig. 16; 1993, p. 166), Fuertes (1989, pls 20, 29%. 50). 9. Fuertesimalva pennellit (Ulbrich) Fryxell, comb. nov. Malvastrum pennellti Ulbrich, Notizbl. Bot. es Mus. Belin Dahlem 11:524. 1932. Type: PERU. ARreEQuipa: Arequipa, 2600-2700 —16 Apr 1925, F. W. Pennell 13206 (HoLotyeeE: F). Hill (1982, p. 3 on notes ie the onan is missing at FE No isotypes are known, but one should be sought ac PH, where Pennell’s original her- barium is preservec Urocarpidium macrocarpum Krapov., Darwiniana 10:624, 1954, Type: PERU. Lima: Canta, 2800-2900 m, 11-19 Jun 1925, F. W. Pennell 14589 (Hotorypr: US; isotypes: GH, S). Distribution.—Peru (Arequipa, Lima); 2500-2900 m elevation (O—3500 m fide Brako & Zarucchi 1993). [ustrations. —Krapovickas (1954a, fig. 2J, fig. 3D-F, pl. 5). 10. Fuertesimalva pentacocca (Krapov.) Fryxell, comb. nov. Urocarpidium pentacoccum Krapoy., Bonplandia 3:69. 1970. Type: ARGENTINA CATAMARCA. dep. Ambato, Sierra de sea desde E] Rodeo he icia el Cerro Manchi do. 2900-3100 m, 23—25 Feb 1967, A. ziker 19223 (HOLOTYPE: CTES). Distribution.—Argentina eae 2900—3100 m elevation. I/ustrations. —Krapovickas (1970, fig. 3C-E). 11. Fuertesimalva pentandra (Schumann) Fryxell, comb. nov. Matlvastrum pentandrum Schumann, Martius Fl. Bras. 12(3):273. 1892. Urocarpidinm pentandrum (Schumann) Krapov. Darwiniana 10:362. 1954. Typr: ARGENTINA CaTaAMARCA: Cuesta de Muschaca, F. Schickendantz 305 (tecroryer, chosen by Krapovickas 1954a: B as photo F- 93 21; ISOLECTOTYPE: CORD), pg catamarcense 1.M. Johnst., Contr. Gray Herb. 70:75. 1924. Type: ARGEN- INA. CaTAMAarca: dep. ee Igala, El Candado, 2 Sep 1916, P. ee iisen 1388 (HOo- LoTYPE: GH; tsoryers: MO, Distribution.—Argentina (Catamarca). Ilustrations. —Krapovickas (1954a, fig. 21, pl. 8). 12. Fuertesimalva peruviana (L.) Fryxell, comb. nov. FRYXELL, Fuertesimalva, a new genus re. — ile ee L., Sp. Pl. 688. 1753. Malvastrum peruvianum (L.) A. Gray, Bot. U.S. Exp. 1:146. 1854. Mal/veopsis peruviana (L.) O. Kuntze, Revis. Gen. Pl. 3(2):21. i. npn ashe (L.) Krapov., Darwiniana 10:269. 1954. Type: Hortus Upsaliensis, s ssieu (HOLOTYPE: LINN-870.5; IsoryPe: P-JU). Malva uaaglegs Te. Bull. So. Imp. Naturalistes Moscou 36:563. 1863. cee mathewsit (Turcz.) Krapov., Darwiniana 10:616. 1954. Type: PERU. Lima: Valley o Lima, A. ia (ISOTYPE: K). Distribution. —Coastal deserts of Peru and Chile; 100-600 m elevation (O—3000 m fide Brako & Zarucchi 1993). [/lustrations. —Diaz (1969, fig. 5), Krapovickas (1954a, fig. 2B-D, fig. SAL, ple. 2.7). 13. Fuertesimalva sanambrosiana (D.M. Bates) Fryxell, comb. nov. — tapananianiiza D.M. Bates, Gentes Herb. 9:383. 1965. Type: ae : TURADAS: San Ambrosio, above Punta Potalas, 250 m, 5 Dec 1960, Is SLA POCAVE Sie 7 (HOLOTYPE: K). Distribution. —Chile (Islas Desaventuradas); I/lustration.—Bates (1965, fig. 280). “above 250 m” elevation. 14. Fuertesimalva stipulata (Fryxell) Fryxell, comb. nov. Urocarpidium stipulatum Fryxell, Contr. Univ. Michigan Herb. 17:168. 1990. Typs: PERU. CaJAMARCA: Las — above San Benito (NE of Trujillo), 1400 m, 2 Apr 1987, C.L. Burandt, DJ. Keil & A. Sagdsteguit 2333 (HOLOTYPE: F; tsorype: NY). Distribution. —Peru (Cajamarca); 1400 m elevation. Ilustration. —Fryxell (1990, fig. 4). ACKNOWLEDGMENTS I am grateful to Javier Fuertes for insightful conversations on both the taxonomy and the etymology of Urocarpidium. Since these conversations led to the reduction of the name to synonymy, I feel it is appropriate to propose the new name Fvertesimalva in his honor, compounded from his surname plus the root of the family name Malvaceae, meaning “Fuertes’ mallow,” in recognition of his continuing contributions to our knowledg of this family. I also thank L.J. Dorr and an anonymous reviewer for comments that materially improved this manuscript. Support from NSF Grant DEB 9420233 to John La Duke and Paul Fryxell is greatly appreciated. REFERENCES Bates, D.M. 1965. Notes on Urocarpidium Ulbrich and Tarasa Philippi. Gentes Herb _ «971. Malvaceae. In: LLL. Wiggins and D.M. Porter. Flora of the Galapagos Islands. Stanford University Press. Pp. 666-69 A, 76 Sipa 17(1) Brako, L. and J.L. Zaruccui. 1993. Catalogue of the flowering plants and gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45:663-664. [U — ,A. 1969 Diaz C. . Estudio de species de la familia Malvaceas ideradas malezas en iis oar de la costa peruana. Raaaondiena® :149-18 Eviasson, U. 1970. Studies in as apagos plants IX. New aie and distributional records. Notis. 123:34 57. ae XELL, R.A. 1988. eee Of Mexico. Syst. Bot. Monogr. 25:1—522 «dO, Be species of Malvaceae from South America. Contr. Univ. Michi- gan Herb. 17: 163-17 .1992a. een In: Flora de Veracruz, fasc. 68, pp. 1-255. 1992b. Malvaceae (118). In: Flora of Ecuador, no. 44 , pp. 1-14 —_________. 1993. Malvaceae. In: Flora del Bajfo y de regiones ren Ose 16, pp.1-175. and A. Krapovickas. 1990. The Malvaceae published by Turczaninow. Contr. ea Herb. 173-182 RTES, J. 1989, veer In: Blore del Real Expedicién Botanica del Nuevo Reyno de Granada hn 1816) promovida y oo sor José Celestino Mutis. Ediciones de Cultura Hispanica: Madrid. 25(2):1—8 Hint, S.R. 1982. A eecius of a. os Matvastrum A. Gray (Malvaceae: Malveae). Rhetlor 84:1-83, 159-264, 317- HurTcuinson, J. 1967, The genera of ee plants. Oxford University Press. {Urocarpidium, vol. 2, p. 558 1954 5) KRAPOVICKAS, A, 4a. Estudio de las especies de ae nueva seccion del género U ‘enema Ulbr. (Malvaceae). Darwiniana 10:606 . 1954b. Sinopsis del género Tarasa (Mi - on Bol. Soc. Argent. Bot. 5: 1965. Notas sobre Malvaceae. III. Kurtziana 2:113—126 1967. Notas citotaxonémicas sobre Malveae. Kurtziana 4:29—37. 1970, Malvaceas nuevas sudamericanas. Bonplandia 3:63—7 2 —_________ and P. A. Frvxet. 1993. Algunos tipos de Malvdceas descriptos por C. B. Presl. Bonplandia 7:57—62. La DUKE yJGvand J. ee BLEY. 1995. A chloroplast DNA based phylogeny of the Malvaceae. . Bor. 20:259-2 Un BRICH, E. 1916. aa vaceae age novae vel criticae imprimis Weberbauerianae II. Bot. Jahrb. Sysc. 54 (Beibl. 117):48— ROBINSONECIO (ASTERACEAE: SENECIONEAE) A NEW GENUS FROM MEXICO AND GUATEMALA THEODORE M. BARKLEY and JOHN P. JANOVEC Herbarium, Division of Biology Kansas State University Manhattan, KS 66506-4901, U.S.A. ABSTRACT Robinsonecio, gen. nov., is a distinctive genus of two species from high elevations in Mexico and Guatemala. The plants are subscapose herbs with ovate to lanceolate or spatu- late aa eaves and tussilaginoid microcharacters. Two new combinations are proposed: Sch.Bip.) T.M. Barkley & J.P. Janovec, and Robinsonecto porphyresthes (T.M. Bakley) T T. M. Barkley oy P. Janovec. RESUMEN Robinsonecio, gen. nov., es un género con dos especies que viven en lugares elevados de éxico y Guatemala. as plantas son herbaceas, subescaposas, con hojas basales de ovadas a lanceoladas o espatuladas y microcaracteres tussilaginoides. Se proponen dos nuevas bing es: Robinsonecio gerberifolius (Sch.Bip. IT M. Barkley & J.P. Janovec, y Robinsonecto pea (T.M. Barkley) T.M. Barkley & J.P. Janovec. A combination of new information and revised taxonomic theory has led to the notions that the traditional, encompassing concept of the super- genus Senecio is no longer tenable and that instead, numerous segregate genera are justified. The relevant thinking was presented in a series of pa- pers by C. Jeffrey, B. Nordenstam, H. Robinson, and others, and it 1s sum- marized in the monumental review of the Asteraceae by Bremer (1994, q.v. for references). A catalog and justification of the segregate genera for Mexico and Central America are presented by Barkley et al. (in press). Our studies have shown that the Mesoamerican Senecio gerberifolins and S. porphyresthes clearly are related to each other but that they do not belong to any of the currently recognized segregate genera. Therefore, the purpose of this paper is to erect a new genus, Robinsonecio, to accomodate these two species. The tribe Sevecioneae includes two phyletic lineages that are recognized formally as the subtribes Senectoninae Dumortier and the Tussilagininae Dumortier, or the “senecionoids” and “tussilaginoids.” The latter lineage was known in the older literature as the “tephroseroids” or the “cacalioids.” Species that were referred to Senecio, s.1., in the traditional, super- genus sense are found in each of these phyletic lineages, 1.e., “Senecio, s.l.” 1s clearly polyphyletic (Barkley et al., in press; Bremer 1994). A third subtribe, Sipa 17(1): 77-81. 1996 58 Sipa 17(1) the distinctive Blennospermatinae Rydb., includes no species referable to Sene- cio, S.1., and its taxonomy is not relevant to this paper. The senecionoid lineage is characterized by having style branches with separate stigmatic lines, the upper ends of the stamen filaments with swol- len collars (“balusterform”), the anthers often with distinctive thickenings in the lateral walls of the endothecial cells (“radial” thickenings), and chro- mosome numbers based on x = 10 or 20 or numbers derived therefrom (in- cluding 22 or 23). In the tussilaginoid lineage, the stigmatic surface is entire or nearly so across the inner face of the style branch, the filaments are cylin- drical and not typically swollen, the anthers often have distinctive thicken- ings in the transverse (end) walls of the endothecial cells (‘polarized” thicken- ings), and the chromosome numbers are based mostly on x = 30 or numbers derived therefrom, including numbers down to » = 24 (Bremer 1994), The species that we call Robinsonecio gerberifolins was among the herba- ceous senecios of the informal assemblage “Lygentes” in the scheme of Barkley (1985), which recognized Senecio, s.1., as a single, encompassing genus. Its inclusion there merely reflected superficial similarity, bue the description of the segregate Senecio (Robinsonecio) porphyresthes noted that the two species were different from the other members of the “Lugentes” assemblage (Barkley 1989). Specimens were borrowed for study (cited with the species descriptions below) and slides made to observe the microcharacters, using the tech- niques developed by Dr. H. Robinson of the Smithsonian Institution (pers. comm.). Slides were made for R. gerberifolius from the collections Beaman 2535 (GH) and Morales A. 164 (WIS) and for R. porphyresthes from Stanford et al. 079 (NY). The permanent slides are deposited in herb. KSC The slides indicate that both Robinsonecio gerberifolus and R. porphyresthes have the entire stigmatic areas and cylindrical stamen filaments of the tussilaginoid lineage. In addition, R. gerberifolins has a chromosome num- ber of v = 30, vouchered by: Beaman 1948 (Stoutamire & Beaman 1960) ” thickenings in the endothecial cells of the anther. The chromosome number for R. porphyresthes is unknown, and it seems to and it has “polarized have radial thickenings in the endothecial cells, but in our experience, this character 1s rather variable. The achenes of R. gerberifolius have a pronounced and perhaps a distinctive carpopodium,; the achenes in-hand for R. porphyresthes are immature, and so the nature of the carpodium is unknown. The two species of Robinsonecio resemble each other in gross aspect and habitat and are unlike any other species of the tussilaginoid lineage. The species of the “Lugentes” group of Senecio are typically senecionoid in mucrocharacters and chromosome numbers and are comfortably similar to each other in gross aspect. Robinsonecio is distinctive in that it is the only tussilaginoid segregate of BARKLEY AND JANOVEC, Robinsonecio, a new genus 79 Senecio, s.1., that consists of subscapose herbs with leaf blades ovate to spatu- late and tapering to a winged petiole, and which has radiate heads with yellow corollas. Robinsonecio is keyed and its presumed taxonomic place- ment is noted among its Mexican and Central American relatives in Barkley et al. (in press). The geographically restricted, subscapose Senecio cuchumatanes Williams & Molina (Phytologia 31:431. 1975) of Guatemala has a gross aspect similar to that of Robinsonecio and it may prove to belong there, but the taxonomic disposition of this distinctive species must must await further information. Robinsonecio T.M. Barkley & J.P. Janovec, gen. nov. Tyre species: Robinsonecio gerbertfolius (Sch.Bip.) T.M. Barkley & J.P. Janovec. Herbae perennes subscaposae e rhizomate radices fibrosas emittenti ortae. Folia majora in caespitem conferta, caulina bracteiforma. Capitula 1-7; phyllaria (plus/minus) aut 13 aut 16-20, inter se aequilonga sed in series exteriorem at interiorem ordinata; flosculorum micro- structura cum tribu Senecioneis subtribu Tassi/agininis conformis, superficeibis stigmatis trans styli ramulorum faciem interiorem continuis staminumque filimentis cylindricis. 7 = 30. Subscapose herbs 10—30(+) cm tall, herbage variously tomentose to arach- noid or lanate, unevenly glabrate in age. Stems single or rarely 2, arising from a thick, fibrous-rooted rhizome. Basal leaves in a prominent rosette, blades ovate to lanceolate or spatulate and tapering to a weakly winged petiole, margin revolute or flat and with callose denticles; cauline leaves few and reduced. Capitula 1—7(+); principal phyllaries ca. 13 or ca. 16-20, about equal in length but in an inner and an outer series; ray florets ca. | 2— 13(+), pistillate, and apparently fercile; disk florets bisexual and fertile; stigmatic surfaces continuous across the inner faces of the style branches; upper stamen filaments cylindrical and without expanded collars. Achenes glabrous or with appressed hyaline hairs; pappus of both ray and disk florets of abundant, white, minutely barbellate bristles that are about as long as the disk corollas. 7 = 30. The genus name is derived from Robinson and Senecio to honor Dr. Harold Robinson, an intrepid explorer of the highways and byways of systematic botany. KEY TO SPECIES 1. Capitula (1—)3—7(+); principal phyllaries greenish or yellowish to lightly anthocyanic; calyculate bracts linear and nearly as long as the principal phyl- laries; achenes glabrou 1. Robinsonecio gerberifolius — eccie 1(—2); principal phyllaries with margins and apex pinkish-purple — » magenta; calyculate bracts few and reduced; ac me apparently with ee hyaline hairs. Robinsonecio porphyresthes 1. Robinsonecio gerberifolius (Sch.Bip. in Hemsley) T.M. Barkley & J.P anovec, comb. nov. BasionyM: Sevecio gerberifolius Sch.Bip. in Hemsley, Biol. 80 Sipa 17(1) Centr. Amer. Bot. 2:240. 1881. Type: MEXICO. Veracruz: pic d’Orizaba, 1838, J. Linden 487 (isotypes: GH!, K, KSC-photo of K!, MICH!). Subscapose perennial herb 1.5—2.5(-3+) dm tall; herbage densely to loosely floccose-tomentose to arachnoid or lanate, unevenly glabrate in age, especially on the upper sides of the leaves. Stems I(—2), arising from a thick, fibrous-rooted creeping rootstock. Basal leaves in a prominent ro- sette, blades ovate to ovate-lanceolate or spatulate, tapering to a weakly winged petiole, 4-18 cm long overall and 1—3(—4) cm wide, coriaceous, margin minutely toothed, weakly revolute; cauline leaves few, reduced to mere bracts. Capitulescence a corymbiform cyme of 4— 7(+) heads; princi- pal phyllaries ca. 13, broadly ovate and of equal length, in an inner and an outer series, (8—)10—-15 mm long, the tips greenish or yellowish to but lightly anthocyanic; calyculate bracts linear, 4—7 and nearly as long as the phyllaries; ray florets ca. (8?—)13, the ligule 12-16 mm long. Achenes gla- brous, 3.5—4 mm long. »# = 30. Alpine and subalpline grasslands and open woodlands at high elevations in selected sites in the Trans Mexican Volcanic Region of Mexico and in the Sierra de los Cuchumatanes in Guatemala. Specimens examined: MEXICO. DISTRITO FEDERAL: Telapon, Sep 1937, E. Lyonnet 1653 (NY). MEXICO (Edo.): Ixcaccihuatl, 30 Jul 1958, J.H. Beaman 1948 (GH, MEX, MO, NY, WIS, UC); Tlaloc, 22 Aug 1958, aan Beaman 2303 (GH); Telapé6n, 20 Nov 1965, W. Boege 23 (CAS); Contrafuertes de la Joya, 13 Aug 1955, R.T. Clausen s SM. a cerca Paso de Cortes, 10 Sep 1966 R. Cruz ne oe (WIS); Ixtaccihuatl, 23 Oct 1976, J. Garcia P. 171 (CAS, F, MICH, NY); Ixtaccihuatl, 27 Jul 198 4, J. Garcia P. & M. Seis L. 1810 (CAS, KSC); Ixtaccihuatl, 16 Aug 1960, H. I/tis, F tis, & R. Koeppen 989 (MICH, WIS); Telapé6n, Dec 1928, E. Lyonnet 395 (GH, MO, NY); Ixtaccihuatl, 13 Sep 1953, E Matuda 29036 (NY); La Joya de Alcalican, 14 Nov 1976, R.A. Morales A. 164 (WIS); Ixtaccihuacl, Oct 1905, C.A. Purpus 1516 (F, GH, MO, NY); Ixtaccihuatl, 30 Dec 1965, J. Rzedowski 22773 (MICH), Ixtaccihuatl, Dec 1968, R. Weber, s.n. (CAS). PUEBLA: apc 2 Sep 1958, J.-H. Beaman 2535 (MICH). VERACRUZ: Orizaba, 16 Aug 8, JH. ae 2290 (GH), Cofre de Perote, 27 Sep 1973, C. Delgadilla & J. Dorantes 3085 (F); Cofre . gue 27 Jun 1982, G. Diges, M. Nee, & G. Schatz 2594 (F); Cofre de Perote, 28 Aug 1971, J. Dorantes 324 (GH, MO), Cofre de Perote, 20 Aug 1972, J. Dorantes etal, O1560 Aon Cofre de Perote, 2 Oct 1984, J. Garcia et al, 1901 (CAS, KSC); peak of Orizaba, Aug 1840, H. Galeotti 2170 (K, photo-KSC!, paratype); Pic Orizaba, 1867, Liebman 140 UX, photo-KSC!, Se Citlaltepetl (Pico de Orizaba), 30 Jun 1949, L.W. Swan, sm. (CAS). GUATEMALA. HUEHUETANANGO: cumbre de la Sierra de los Cuchumatanes, 28 1940, B.C. Standley 81489 (F); between Tojquia and Caxin, Sierra de los Cuchumatanes, 6 Aug 1942, J. A.Steyermark 50178 (F, MO). 2. Robinsonecio porphyresthes (T.M. Barkley) T.M. Barkley & J.P. Vee: oP nov. aos M: Senecto porphyresthes 'T.M. Barkley, Phytologia 6 7:243. 1989. Typr: MEXICO. Tamautipas: on mountain top 7 km SW of Mina in forest of large i ses floor of low vegetation, elev 3430 m °40°'N 99°45'W), 5 Aug 1941, L.R. Stanford, L. itl & R.D. Northcraft i (HOLOTYPE: NY!; isorypes: G nee MO!,NY!, W BARKLEY AND JANOVEC, Robinsonecio, a new genus 81 Subscapose herb 1.0—1.5 dm tall, herbage closely lanate- tomentose, but loosely tomentose toward the oe with long, arachnoid hairs, unevenly glabrate in age, upper side of leaves glabrescent, lower side persistently felted-lanate. Stem 1, arising in a cluster of basal leaves at the end o rhizome. Rhizome creeping, simple, nearly 1 cm in diameter, covered by exfoliating bases of old leaves and producing abundant fibrous branching roots. Basal leaves with blades narrowly oblanceolate to subspatulate, ta- pering toa winged petiole, 4-10(—12) cm long overall and (0.8—)1—1.5 cm wide, subcoriaceous, margin denticultate with callose denticles, weakly revolute or flat; cauline leaves 2—5, reduced to mere linear bracts to 15 mm long. Capitula 1(—2), principal phyllaries ca. 16-20, linear-lanceolate and of equal length, in an inner and an outer series, 12-14 mm long, the mar- gins and tip permanently and prominently pinkish-purple or magenta; calyculate bracts few and reduced; ray florets 12(+?), the ligule 10+ mm long. Achenes with flat, hyaline hairs that presumably persist, 2 mm long (ammature). A distinctive entity known to us only from the type collection, cited above. If it has been collected again in the intervening years, the specimens have escaped our attention. ACKNOWLEDGMENTS It is a pleasure to thank the curators of the above noted herbaria for lending specimens. Jose Luis Villasenor of the Herbario Nacional (MEXU), Instituto de Biologia, Universidad Nacional Autonoma de Mexico, gladly shared his superb knowledge of the Mexican Compositae with us. Rupert Barneby of the New York Botanical Garden kindly prepared the Latin di- agnosis. Relevant type specimens were located by D.J.N. Hind in the her- barium of the Royal Botanic Gardens, Kew (K), and photocopies were gra- ciously supplied. This paper is Contribution No. 96-397-] from the Kansas Agricultural Experiment Station, Manhattan. REFERENCES BARKLEY, T.M. 1985. Infrageneric groups in Senecio, s.l., and Cacalia, s.1. (Asteraceae: Senecioneae) in Mexico and Central America. Brittonia 37:211—218 _.S«d'OS. New taxa and oo combinations in ne in Mexico and the United States. Phycologia 67: 253 BARKLEY, T.M., B.L. Clark, a A. M. . NSTON (in press). The se ae genera of Senecio, s.l., and Cacalia, 5.1. (Asteraceae: Senecioneae) in Mexico and Central America. In: D.J.N.Hind et al., eds. Proceedings of the international Cee conference, Kew, 1994. he Botanic Gardens, Kew. BREMER, 1994. Asteraceae: cladistics & classification. Timber Press, Portland. ea W.P. and J.H. BEaAMAN. 1960, Chromosome studies of Mexican alpine plants. ricconia 12:226-230 Sipa 17(1) BOOK NOTICE Brako, Lois, AMy Y. RossMAN, and Davib EF. Farr. | Common names of 7,000 Vascular Plants in the United States. (Contr.U.S. Nat. Fungus Coll. No. 7). (SBN 0-89054-171-X, pbk.) American Phytopathological Society Press, 3340 Pilot Knob Road St. Paul, Minnesota, 55121, 612/454-7250. $29.00. 304 pp. 1995. Scientific and Designed primarily with the plant pathologists and agriculturists in mind, this book is a useful resource for anyone oe in the common names for plants native to and cultivated in the United States, including Hawaii and southern California, Texas, and Florida. The book is well designed to facilitate its use with four cross-referenced alpha- betical sections: 1) Scientific names - entry by genus followed by the family name and generic common name, if any; species listed under each genus followed by one or more 2) Common names - multiword vernacular names; authorities given for all listed taxa. ere name(s) listed under names entered separately under each word of the name; the comp the indexed word; and the Latin name placed after the vernacular. 3) Synonyms - scientific = names by which a species may be known are listed, followed by the correct latin name; appears to be somewhat incomplete. 4) Families and Genera - the included genera listed by sues The nomic ee Only those taxa are giv consulted sources. No attempt has been made to manufacture “standard” common names the coverage of genera is rather exaustive (e.g cluded taxa were selected from standard phytopathological, floristic, and agro- en for which common names are published in the for other species of a genus. Hence, Bombacaceae with f ive listed genera, Bignoniaceae with 12, and Myrtaceae with 16), whereas tural) sis much less complete (e.g. only 14 listed species non IOTI-dyVer ULCUL at ICs the coverage o of Verbena {including ee ee eae six species of Tradescantia s. ae and six of Penstemon,: however, 25 species of Trifolinm).—Roger Sanders, Research Associate, Botanical Research Insti- tute of ‘Texas. Stipa 17(1): 82. 1996 A NEW VARIETY OF SWERTIA RADIATA (GENTIANACEAE) JAMES HENRICKSON Department of Biology California State University Los Angeles, CA 90032, U.S.A. ABSTRACT _ Plants of the wide-ranging Swertia radiata from the Sierra de la Madera near Cuatro negas, Coahuila, in México differ from all other ee pulations in having corollas that an et y open and remain globose at anthesis. As this feature is consistent through- . radiata Vat. out the Sierra de la Madera, the plants are ate as a new variety, maderensis. RESUMEN Las plantas de Swertia radiata, una especie de distribucién amplia, procedentes de la Sierra de la Madera, cerca de Cuatro Ciénegas (Coahuila, México), se diferencian del resto mente y permanecen globosas — de las poblaciones por tener corolas que se abren solo parcia durante la antesis. Dado que este cardcter es consistente a lo largo de toda la Sierra de La Madera, estas plantas son reconocidas aqui como una nueva variedad: S. radiata var. maderensis. Studies in connection with the Chihuahuan Desert Flora project have provided specimens of Swertia radiata (Sellogg) Kuntze from the Sierra de la Madera near Cuatro Ciénegas in central Coahuila, México. These speci- mens differ from all other collections of this wide-ranging species in that their petals remain erect at anthesis forming a globose corolla. In all other populations of the species the petals are distinctly spreading at anthesis. Based on this character, which is consistent throughout the Sierra de la Madera, the plants are recognized as a new variety. Swertia radiata var. maderensis Henrickson var. nov. (Fig. 1) Differt a var. radiata petalis erectis corollam cupulatam facienti (petalis non reflexis), staminibus erectis per anthesin non patulis. Perennial, rarely biennial, monocarpic? herbs 3-18 dm tall from a thick- ened, black-barked taproot; stems 0.7—1.5 cm wide at the base, glaucous, glabrous. Basal rosulate leaves lanceolate, oblong-lanceolate to oblong-oblan- ceolate, (10—)15—30 cm long, 2—4 cm wide, tapering to a winged petiole at base, present as rosettes throughout the growing season; cauline leaves whorled, (2—) 3—4(-6) per node, of similar outline to the basal leaves, S—15 cm long, (1—)2—3 cm wide, reduced to leafy bracts in the inflorescences, all Sipa 17(1): 83-87. 1996 84 Sipa 17(1) Fic. 1. Swertia radiata var. maderensis. A. Growth habit showing basal rosettes and an erect flowering plant, which develops directly from the central rosette. B. Portion of Howering stem showing subtending whorled leaves, flower distribution, spreading sepals and glo- bose corolla. C. Flower with characteristic globose corolla; note outer impression of the paired basal petal foveae. D. Petal abaxial view showing basal fringed corona, the paired fovaea with the marginal fringes. E. Same in side view. F. ovary. G. Immature fruit. All from Henrickson & Wendt 11887. leaves acute to + obtuse at tip, broadly to narrowly cuneate at the sessile, sheathing base, entire, fleshy, glaucous, glabrous. Flowers borne in much- branched dichasia, or cymose, leafy-bracted panicles (4—)5—9 cm long, these often continuous from the lower half or base of the plant to the tip; pedicels 25-65 mm long; sepals 4, lanceolate, lance-ovate, 9-15 mm long, green with white, entire to finely erose margins, spreading at anthesis; corolla tubes 1-2 mm long, the corolla lobes obovate to oblong-ovate, 9-18 mm long, 6-13 mm wide, obtuse to apiculate at the tips, the margins incurved HENRICKSON, A new variety of Swertia radiata 85 distally, pale dull green, more yellowish upon drying, spotted with purple inside, incurved and erect at anthesis forming a cup-like corolla, each co- rolla see with a pair of parallel foveae (glands) on lower inside surface, the glands 2.5-4.5 mm long, 0.8-1.7 mm wide, surrounded by a strongly fringed membrane 2—4 mm high, the corolla-tube base with a fringe of united fimbriae 2.5—8 mm long forming a corona; stamens erect at anthe- sis, laments 5-10 mm long; anthers 2.5—4 mm long, inverted (the anther tip pointing downward), slightly purplish to light yellow-white, visible at the opening of the corolla; ovary 8-10 mm long; style 2-4 mm long. Fruit 2—3 cm long, 5—7 mm wide, the style persistent; seeds 2.5—4 mm long, 2— 3 mm wide, pitted, rusty-brown, winged along the angles. Type: MEXICO: Coanutta. ca. 40 km WNW of Cuatro Ciénegas in limestone Sierra de la Madera, in mid canyon just west of Cafién de la Hacienda, near 27°02'N, 102°51'W, 7800 ft, 28 Sep 1980, Henrickson 18659 (HOLOTYPE: TEX!; isoryPes: ARIZ, TEX, MEXU). Additional collections: MEXICO. Coanuita: Sierra de la Madera, high crest of main ridge about 2 km E me de mee 8600-8800 ft, 13 Sep 1941, 1.M. Johnston 9035 (LL); ca. 35 air mi Sof C o Ciénegas in Cafién de la Hacienda on limestone, Sierra de la Madera, 27°03'N, aes 7300— ane ft, 5 Aug 1973, Henrickson & Wendt 11887 (TEX). The Sierra de la Madera is an isolated east-west oriented limestone range near the eastern edge of the Chihuahuan Desert in central Coahuila that rises to 3032 m elevation. The Sierra’s massiveness and orientation relative to prevailing air movements from the Gulf of Mexico result in develop- ment of heavy cloud covers during much of the summer rainy season. The upland forests contain stands of Pinus strobiformis Engelm., Abies durangensis Martinez var. coahuilensis (1.M. Johns.) Martinez, Psewdotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco, and Cupressus arizonica Greene along with species of Quercus, Arbutus, Arctostaphylos, and Garrya, and in the mossy understory the new taxon is very common. When first observed in flower (Henrickson & Wendt 11887), the plants were distinctive in their globose, dull greenish corollas. Subsequent observations of the species in California, Nuevo Leén, Arizona, and in various wildflower books, as well as in her- baria (ARIZ, RSA-POM, TEX,LL) have shown that in this feature, the population from the Sierra de la Madera is unique. Swertia radiata is a wide-ranging species with collections known from southern Washington, southwestern Montana, Idaho, Wyoming, the Black Hills of South Dakota, eastern California, Nevada, Utah, Colorado, Ari- zona, New Mexico, west Texas (Guadalupe Mountains) and the high moun- tains of central Nuevo Leén, México (Sierra Coahuilon, Sierra la Marta, Cerro Pefia Nevada, Cerro del Potosi) and in central Coahuila (Sierra de la Madera). In the material I have seen of populations outside of Coahuila, the petals are dull light green to light yellowish, typically maculate with purple, 86 Sipa 17(1) sometimes with a larger purple spot near the petal gland or near the tip; but in all cases the cupped corolla lobes and stamens are strongly spreading at anthesis. In contrast the corollas from the Sierra de la Madera popula- tions remain erect forming a distinct globose cup, and the petals are dull light green or yellow-green with sparse purplish maculations and with a purple marking at the very base below the glands. The anthers are also on slightly purplish in color, though they dry a light yellow color. There is disagreement as to the generic placement of the species. It has been included in Swertia as Swertia radiata (Kellogg) Kuntze [Higgins (1987), Pringle (1993)]} and in Frasera as Frasera spectosa Dougl. ex Griseb. in Hook. {Hitchcock (1959), Munz (1959), Holmgren (1984)}. Holmgren (1984) neatly distinguished the two genera—Swertia having 5-merous flowers with short, thick styles, alternate to subopposite leaves, occurring in wet-moist habitats, and an Old World distribution, with our one New World species Swertia perennis L. being circumboreal. In contrast his Frasera has 4-merous flowers with slender styles, opposite or whorled leaves, and occurs in drier habitats in the New World. Pringle (1979, 1990), however, has concluded that recognition of two genera is not warranted when con- sidering the wide range of variation present in Old World Swertia. He notes that some species of Swertia have distinct styles [see also St. John (194 1)}, and others have 4-merous flowers and that the generic distinctions are not clearly supported by the various cytological, palynological, or phytochemical studies undertaken [see references in Pringle (1979, 1990)}. Thus the spe- cies 18 recognized as Swertia in this study. Throughout the range of the species many variants have been named, all associated with differences in plant size and vestiture. Frasera speciosa var. scabra M.E. Jones (1893), of central Arizona, was based on strongly sca- brous plants with small leaves and large flowers. Three new taxa were pro- posed by Greene (1900): (1) Frasera venosa Greene of southwest New Mexico, with narrow, strongly veined leaves; (2) Frasera ampla Greene of northern Arizona with minutely hirtellous-scabrellous herbage, large but not nota- bly veined basal leaves, and shorter sepals; and (3) Frasera macrophylla Greene of southern Colorado, robust plants with large, glabrous leaves, long se- pals, and purple-tipped corolla lobes {this taxon was retained as a glabrous variety by St. John (1941)}. Lastly, PA. Rydberg recognized two varieties of Frasera speciosa: (1) var. stenosepala Rydb. with numerous flowers, narrow, long sepals, long petals in Colorado and Wyoming, and (2) var. angustifolia Rydb., low plants with short basal leaves, short, narrow cauline leaves, and small flowers from Montana to Colorado. None of these taxa, based on growth form and vestiture differences, have been recognized in modern floras except for taxon macrophylla as noted above. It is certainly realized that the taxon proposed in this paper is of a minor nature, but the charac- HENRICKSON, A new variety of Swertia radiata 87 ters upon which it is based are consistent throughout the large population in the Sierra de la Madera and in this feature it stands apart from other populations of the species. One could suspect that its differences may be associated with a change in pollinator. The flora of the Sierra de la Madera and adjacent Cuatro Ciénegas basin has been enumerated by Pinkava (1984). The new variety joins a long list of taxa endemic to the Sierra de la Madera including Clematis coahuilensis Keil, Rosa woodsii var. maderensis Henrickson, Sedum parvum subsp. diminutum Clausen, Euphorbia pinkavana M. C. Johnston, Choisya katherinae C. H Muller, Poliomintha maderensis Henrickson, Satureja maderensis Henrickson, Penstemon henricksonii Straw, Agave scabra subsp. maderensis Gentry, and Festuca coahuilana Gonzalez & Koch. ACKNOWLEDGMENTS I thank Tom Wendt (LSU) and the reviewers for comments on the manu- script, Paul Fryxell (TEX) for the Latin translation, F. Javier Fuertes-Aguilar (TEX) for translation of the abstract, Bobbi Angel (NY) for the illustra- tion, and the Plant Resources Center at the University of Texas, Austin, for use of facilities. REFERENCES Greene, E.L. 1900. A decade of new Gentianaceae. Pittonia 4:180—-186. Hicains L.C. 1987. Gentianaceae. In: $.L. Welsh, N.D. Rae L.C. Higgins, S. Goodrich, eds. A Urah flora. Great Basin Naturalist Men 9:3 Hircucock, C.L. 1959 Gentianaceae. Gentian Family. i a : Hitchcock, A. Cronquist, M. Ownbey, J.W. st Vascular plants of the Pacific Northwest. Univ. Was Publ. Biol. 17:57—76, 8 Hoimcren, N. 1984. a paaeh In: A. Cronquist, A-H. Holmgren, N.H. Holmgren, Reveal, and P.K. Holmgren eds. Intermountain flora: Vascular plants of the Inter- mountain West, U.S.A. 4:4—23. Jones, M.E. 1893. Contributions to Western Botany No. 5. Zoe 4:254—282. Munz, P.A. 1959. A California flora. Univ. Calif. Press. Berkeley. Pringle, J.S. 1979 New combinations in Swertia (Gentianaceae). Phytologia 41:139-1+ 43. 1990 Taxonomic notes on western American Gentianaceae. Sida 14:179- 187. oe 2 DS, ees In: Hickman, J. ed. The Jepson manual: Higher plants of Caiiorut a. Ae 666— Rep G, P.A. 190 aie on the Rocky Mountain flora XIII. Bull. Torrey Bot. Club. 1-631 —655 Si: cae H. 19. 41. Revision of the genus Swertia eo of the Americas and the reduction of Frasera. Amer. Midl. Naturalist 26:1— 88 Sipa 17(1) BOOK NOTICES BreeDEN, JAMES O. (ed. & intro.) 1994. A Long Ride in Texas: The Ex- plorations of John Leonard Riddell. (ISBN 0-89096-582-X, hbk.) Texas A & M University Press, Drawer C, College Station, TX 77843- 4, (A409) 845-1436. $24.50. frontisp. + vviii + 144 pp., 3 figs, | m ap : Recently discovered in the archives of Tulane University, the travel diary of J.L. Riddell’s explorations of central Texas forms the core (62pp.) of Breeden’s presentation of Riddell’s contribution to Texas natural history. Although Riddell attempted several times to pub- lish his journal, he failed to find an interested publisher and set it aside. Years after his death, a re tele donated it to Tulane University, where he had been a professor for 29 years. The journal is of considerable interest because it documents his observations of the geology, botany, and zoology of the Texas Hill Country in 1839. This antedates by almost 10 years the exploration of Ferdinand Roemer, who is generally thought to be the first natural historian to the region. To round out Riddell’s contribution to Texas, Breeden includes an extensive preface (Spp.), a brief but insightful biography of Riddell (34pp.), and an unaltered version (9pp.) of Riddell’s major published article on Texas (“Observations on the Geology of the Trinity rome Texas, made during an excursion there in April and May, 1839.” Amer. J. Sci. ts. 37:211—17. 1839.). In the case of the preface, biography, and diary, Breeden provides copious end notes to explain outdated plant names, geographic locations, customs, and obscure references made by Riddell. These notes add a dimension to the readability of this volume.—Roger Sanders, Research Associate, Botanical Research Institute of Texas. LAWERENCE, ELEANOR. 1995. Henderson’s Dictionary of Biological Terms: Eleventh Edition. ISBN 0-470-23507-1, hbk.) John Wiley & Sons, New York, New York. (1-800-225-5945) $ $39.95. 693 pp The updated version of this standard reference dictionary incorporates terms related to recent advances in molecular biology and genetics. The main dictionary portion now in- cludes about 23,000 entries. It will serve botanists primarily as an information source for disciplines wick which they are least familiar; classical botanical terms are often omitted (e.g. salverform, hypocrateriform, nexine, tricolpate, fractiflex, medifixed, prolepsis, and brochidodromous lacking; cataphyll inadequately defined). Other helpful brane and appendices include: oe units/conversions, decimal prefixes, the Greek alpha- bet, common Latin and Greek plural endings, structures of 80 common eee -lassification ourlines« f organisms and viruses (modern evolutionary, non-cladistic schemes), and Greek and Latin word roots. The promotional literature purports “Figures that help clarify complicated terms.” In reality this amounts co 12 rather crude diagrams of such items as parts of the awiee trophic level pyramid, and meiosis.—Roger Sanders, Research Associate, Botanical Research Institute of Texas. Sipa 17(1): 88. 1996 NOTES ON SPIGELIA (LOGANIACEAE) JAMES HENRICKSON Department of Biology California State University Los Angeles, CA 90032, U.S.A. ABSTRACT A treatment is provided for the — of Spigelia eee in the genus Coe/ostyl7s that lack elongated secund inflorescences o taxa are recognized: Spigelia loganioides (includ- ing S. texana), and S. ee ae S. lindheimeri, a S. coulteriana). RESUMEN Se presenta un tratamiento taxonémico para las especies de Sprge/7a anteriormente incluidas en el género Coe/ostyl1s, 0 sea las especies sin inflorescencias ee y unilaterales. conocen dee § taxa: Shipelea Vege (incluyendo S. texana), y S. hedyotidea (que incluye S. lindbeimeri y S. coulteriana). INTRODUCTION This paper deals with the species of Spigelia placed by Torrey and Gray in their genus Coe/osty/is, namely Spigelia loganioides (Torr. & A.Gray in Endl.) A , of Florida, Spigelia texana (Torr. & A.Gray) A.DC., of Texas and Spigelia lindheimeri A.Gray of Texas and north central Mexico. North American Spigelia were previously studied by Homer Hurley (1968), but his taxonomic conclusions were never published. This treat- ment differs from that of Hurley. He intended to accept S. /indheimeri as a variety of S$. fexana, and he considered S. /oganioides synonymous with S. texana. However, S. loganioides is an older name than S. texana, and also S. lindheimert differs from S. loganioides and S. texana in several characteristics including growth habit, leaf size, leaf placement, vestiture, and corolla throat width that support its recognition at the species level. Studies of the type S. hedyotidea show it is referrable to S. /indheimeri, and S. hedyotidea is older than S. /indheimerr, studies of the type of S. cowlteriana, reveal that it has been misapplied and belongs within the Coe/osty/is group. TAXONOMY Spigelia loganivides, 8. texana and S. lindheimert have been considered by Torrey and Gray (1839, 1841) and Small (1903) to constitute the genus Coelostylis. Coelostylis was merged into Spigelia by A. deCandole (1845) and later accepted as such by Gray (1878). Torrey and Gray’s Coe/ostylis difters from the other species of Spgelia primarily in the basic structure of the Stipa 17(1): 89-103. 1996 90 Sipa 17(1) inflorescence (Fig. 1). Both groups have terminal, highly modified, deter- minate, dichasial inflorescences and the plants often exhibit a dichotomous branching pattern effected by the presence of terminal inflorescences. In the Coe/ostylis group the upper nodes produce two flowers, a terminal flower ona short, ebracteolate pedicel that is the first to open, and a second flower that develops laterally, opposite a stipule, produced upon a 3-bracted pe- duncle-pedicel that is longer than pedicel of the first flower (Fig. | b—d). Each node also produces two shoots from the leaf axils that terminate in a node with two leaves, 2 flowers, and two more lateral shoots in a dichoto- mous branching pattern that is again repeated. Some plants produce only a single lateral shoot at the upper nodes and exhibit unilateral branching producing leafy monochasia. This pattern is consistent in Coe/osty/7s with one exception, a specimen of S. /oganioides that produced two flowers on the longer stalk, each separated by a three bractlets (Ginzbarg & Davis 914, TEX). But the specimen still produced the single terminal, ebracteate flower characteristic of the group. In the true Spigelias, the inforescences are again terminal and structur- ally determinate, and bordered by lateral leaves that may produce lateral branches, but they appear as indeterminate, elongated, one-sided spikes or racemes. However, structurally they are determinate, laterally branched, helicoid cymes derived from a dichasia through strong one-sided lateral branching (Fig. le-g). The flowers may be sessile or slightly pedicellate along the inflorescence axis, and each flower is subtended by two bracts, one lateral to the flower, the other either above or below the flower on the inflorescence axis (Fig.le), and the bracts occur on alternate sides in con- — secutive flowers, which indicates that the inflorescence may actually be a scorpoid cyme. The inflorescences alone has served to separate the two groups at the generic level, however, some species of Sp/gel7a produce flowers that are essentially identical to those of Coe/osti/is and the two groups have been combined within Sprgelia. Coelostylis is not formally recognized as a genus — or section or subgenus of Sp/ge/ia in this paper, as I defer classification to Katherine Gould, who ts presently studying the genus at the University of Texas, Austin. While Coe/osty/7s is no longer recognized as a genus, the comprising taxa form a closely related group that share many characteristics. They are per- ennial herbs with few to several stems arising from underground fibrous yave a pink color, and the plants —" root-bearing rootstocks. The roots often are thus known as pink-root or worm-grass (Godfrey & Wooten 1981). The stems are erect to ascending, sometimes basally decumbent, few to several branched from the base. Flowering stimulates lateral branching of- ten in a dichotomous pattern, but one or three lateral shoots are produced HENRICKSON, Notes on Spigelia 91 SS \Y| Le GE DE Fic. 1. Comparative diagrams and drawings of the Coe/osty/is and Spigelia inflorescences. A. Diagram of dichasium; a determinate inflorescence with opposite lateral branching. B. Diagram of flower clusters in Coe/osty/is group as modified from the basic dichasium struc- ture; retaining one terminal flower on an ebracteolate pedicel, and a lateral flower on a peduncle-pedicel subtended by three bracts. C. Drawing of nodal flower cluster in Coelostylis. D. Drawing of inflorescence of Coe/ostylis; each node has one terminal flower, one lateral flower produced opposite a stipule — rs shown as circles), two shoots produced from the leaf axils that repeat the pactern. E. Diagram of possible origin of inflorescence of Spigelia group as modifie he dichasial structure. F. Terminal helical cymose inflorescence of Spigelia humboldtiana showing one-sided orientation of flowers, continued branching occurs through axial shoots. G. Enlargement of f, showing the presence of two bracts at nodes of inflorescence. The bracts occur on alternate sides of consecutive Howers. at the flowering nodes. The internodes are shortest near the base, longest medially. The nodes each have two leaves, the margins of which extend as decurrent wings down the subtending internodes, and two, broadly deltate, acute to rounded, whitish, membranous, erect stipules lie between the paired leaf bases. The stems may be glabrous, or more commonly, they have dis- tinct papillae at the nodes, on the stipules, and on the margins of the de- current wings. Leaves are opposite, but in S$. /oganioides are often produced in a con- spicuous whorl of four large leaves below the inflorescence caused by the supression of an internode. In most plants the lower-stem leaves are more oblong-ovate, oblong-elliptical, sometimes ovate to obovate, more obtuse 92 Sipa 17(1) to rounded at the tip, and subpetiolate to petiolate. The upper-stem leaves may be similar to the lower leaves in shape or they may be considerably larger, more lanceolate, elliptic to narrowly ovate, acute, sessile, and thin- ner in texture. The blades are bicolored and range from thin and membra- nous, to firm and sub-coriaceous; the midvein is impressed above, and the midvein and secondary arcuate veins are raised and sometimes yellowish beneath. The leaf surfaces may be smooth or scabrous with hardened papil- lae, particularly along the basal margins of the upper surface. Flowers, as noted above, are terminal at the nodes, with each node pro- ducing 2 leaves and 2 flowers, one terminal with an ebracteolate pedicel 0.5—3 mm long, the second produced opposite a stipule with a longer pe- duncular-pedicellar stalk S—10(-14) mm long that bears 2—3 subulate brac- teoles 1—3(—6.5) mm long. In addition each upper node produces 2 (some- times | or 3) leafy shoots, from the leaf axils, and these shoots again terminate —_— in leaves and flowers. Calyces are deeply 5 parted, with linear to narrowly lanceolate, acute to obtuse, green, scarious-margined, sometimes papillate-margined sepals. The corollas are salverform to funnelform, thickish, white or tinged or verti- cally striped with pink or lavender, 10-20 mm long. The tubes are angled to cylindrical, slightly ampliate. The 5 lobes are oblong-ovate, obtuse, val- vate, and their margins flare outwards in bud. The stamens are epipetalous, borne in the mid or distal corolla tubes, with free filaments only 1-1.5 mm long; the linear, sagittate, introrse anthers surround the style. The styles are included to slightly exserted, cylindrical and is densely pilose-setose medially—the hairs serve to remove the conspicuous yellow pollen from the anthers as the style elongates. The stigmas are capitate, slightly 2- lipped, and the terminal 70 percent of the style is deciduous leaving a hardened style base tipped with a crown-like structure on the fruit. Fruit consist of 2 (rarely 3) indurate, sclerified, obovoid to nearly spher- oid, yellowish cocci fused for 70-80 percent of their common surface. The outer surface may be initially papillate, but later is smooth and marked with stronger yellow veins. The fruit cocci dehisce along the common ver- tical septum with the two halves springing opening and falling from the subtending nectary disk. The basal disk is spongy but strongly indurated, plate-like, stramineous, oblong-ellpitical in outline and obtuse at each end as seen from above. Each cocci contains 6, (rarely 10—12) seeds that develop around a central placenta. The seeds are obpyramidal, with the outer sur- face uneven, spongy-pitted; the interior 3 surfaces are more coarsely cuber- culate-ridged. The inner-most point has a prominent hollow pit marking the point of vascular attachment. The Coelostylis and Spigelia taxa and the two taxa comprising taxa Coelostylis can be distinguished by the following key: HENRICKSON, Notes on Spigelia 93 ro Hows 2 : node, the first-to-flower terminal with an ebracteolate pedicel ong, the lateral flower with a three-bracteate peduncle-pedicel 5— 14 mm ee he ee nodes producing (1—)2(—3) lateral aa shoc Th mere group that repeat the patter B. Plants 1.5—3.5 leaves mostly oblong-lanceolate, 3-6 cm long, (sometimes smaller and more ovate os Florida) drying thin membranous, the plants usually with dm =i with 1-few stems from the base, the Herpes oO. a whorl of 4 leaves subtending the inflorescence; the internode wings Se aaa ony for a short distance below the node; corollas broadly ampliate, 4-5 mm wide below the lobes; plants of west-central Florida igelia loganioides and east- vena Tex B. Plants 0.5—1.2(—2) de tall, usually bushy with many stems from the base; oe leaves mostly narrowly lanceolate (sometimes more ovate), 1.2—3(—3.5) cm long, typically rather chick with the upper sur- face eee pay sillate and wrinkled in dried specimens; the plants rarely forming a whorl of 4 leaves below the inflorescence; the internode papil- late all along the internode wings a also on the internode between the wings; corollas narrowly ampliate, 2—3.5 mm wide below the = plant of east-central Texas, north- cent Mexicc pigelia Pena: A.Flowers produced in terminal, elongated, one-sided. spike-like or like inflorescences, the inflorescences actually of determinate ae cymes The true Spigelias e- with indeterminate, unilateral branching. Spigelia loganioides (Torr. & A.Gray in Endl.) A.DC., Prod. 9:4. 1845 Coelostylis loganioides Torr. & A.Gray in Endl. Nov. stirp. dec. (5):33, 1839; Iconogr. 9):c. 101, 1841 (illust.). Type: U.S.A. Floripa. Marion Co.: Prope Fort King, Ee Ale sm. (HOLOTYPE: NY!, lsoTyPE: NY!). sae texana (Torr. & A.Gray) A.DC., Prod. 9:5. 1845. Coelostylis texana Torr. & Gray . Amer. 2(1):44. 18: a TYPE: U.S.A. Texas: (exsiccata fascicle IIT) Drammond oe ne NY!; tsoryprs: G!, GH!, GH!) Erect, 1-few stemmed herbs (10—)20—30(—50) cm tall; internodes (1-) 2—6(—8.5) cm long, glabrous except where papillate along the margins of the distal-most decurrent internode wings, on the stipules, and on the lower leaf margins. Leaves opposite, usually in a whorl of four below the in- florescences; basal-stem leaf blades often ovate, oblong-ovate, to 2 cm long, acute to rounded at the tip, tapered to a winged petiole at the base, the mid- and upper-stem leaf blades narrowly ovate, lanceolate, oblong-ellip- tical, elliptical to rhombic, (1.4—)3—6.5 cm long, (0.5—)1-1.7(—2.2) cm wide, acute, obtuse, mostly minutely apiculate at tip, the bases cuneate, sessile or tapering to a winged petiole-like base to 3 mm long, the margins entire, often revolute or enrolled when dry, fleshy, the blades drying thin membranous, glabrous except where sometimes scabrously papillate along the upper-surface margins above the petiole, glabrous, slightly more glau- cous beneath. Terminal-flower pedicel 1—1.5(—2.5) mm long, ebracteolate; lateral-flower peduncle-pedicel 4—7(—14) mm long, with subulate bractlets 1—4.5 mm long, 0.2—0.4 mm wide; sepals linear-lanceolate, (2.5—)3.5—5.5 94 Stipa 17(1) mm long (slightly accrescent in fruit), 0.4—1 mm wide at base, acute, gla- brous or weakly scabrous along the margins; corollas white throughout or with vertical, pale-lavender lines extending up the tubes and bordering the lobes, funnelform, 12—14.5(—20.5) mm long, the tubes ampliate, 8—9(— 15) mm long, to 4-5 mm wide at aes throat (pressed), the lobes oblong- ovate, spreading, 3.5—5.5 mm long, 2.2—3.5 mm wide, acute, entire, thick- ish; stamens borne in the mid-corolla Pine filaments 1-1.5 mm long; anthers 1.1—1.3 mm long; styles 6—7.5 mm long, Pes, the stigma borne in the distal tube, the persistent style base 1.7—2.3 mm long. Fruit cocci 2, nearly spheroidal, slightly divergent, cee the pair 3-4.5 mm high, 5—5.5 mm wide, the subtending thickened disk (3.2—)4—5 mm long; seeds 6-12 per cocci, 1.5—1.8 mm long, 1.3—1.5 mm wide. (Figs. 2, 3a—c, 4). As recognized here Spigelia loganivides occurs in both Florida and Texas. It is distinguished from S. hedyotidea by: (1) its taller growth habit; (2) its arger, usually more membranous leaves; (3) a tendency to be glabrous — throughout except at the nodes where stiff papillae usually occur on the basal portion of the upper leaf margins and continue down the petiole and the discal portion of the decurrent internode wings, and on the margin and outer surface of the stipules; (4) its tendency to produce a whorl of four leaves below the leafy inflorescences, and; (5) the slightly longer, more con- spicuously ampliate corollas. Specimens in Texas generally conform well to these features as do most specimens from Florida, however, some Floridian collections, including the type of S. /oganioides, differ vegetatively. While the type locality was designated in the original description as ly is from near Fort King ante being near Tampa Bay, the type collection actua in Marion County (Fort King now lies within the City of Ocala, Florida). Plants from this region differ from those of most other localities in having small, mostly broadly ovate, sometimes lanceolate, obtuse to acute-tipped, opposite leaves that do not form a whorl below the inflorescence; the plants are also relatively short in stature (Fig. 2a). The type collection, was made by Burrows and Alden near Fort King, (E. Silas Burrows was an army phy- sician stationed for time at Fort King during the second Seminole War— 1835-1842, Wunderlin pers. comm.) and the type specimen, illustrated in Endlicher (1841), is redrawn here as Fig. 2a. It consists of a curved stem 25 cm long, with ovate to broadly ovate, obtuse-tipped leaves 14—20 mm long and 6-8 mm wide in the distal 10 cm of the stem. The leaves are all opposite and do not form whorls below the flowers. A second old collection from the type region, fin pinetis, prope Fort King, Florida, Jul 1848, F Ragel 327, (GH-2 sheets, NY, FLAS)} also con- sists of moderately short plants 1-2 dm tall. Most (21) of the 24 stems present on these four sheets have ovate, sometimes broadly ovate, obtuse- tipped leaves similar to those present on the Burrows and Alden collection Henrickson, Notes on Spigelia ye Fic. 2. Variation in Spregelia logantoides in Florida. A-B. From the Fort King population in Marion Co., Florida, showing plants with more ovate, shorter leaves and both axillary and terminal flowers (Redrawn from the figure in Endlicher (1840) from the Burrows and Alden holotype). C. Typical specimen from Levy Co., Florida, showing obovate to ovate basal leaves and characteristic quartet of larger elliptical leaves and characteristic branch- ing pattern. These plants are identical to plants from eastern Texas [From Wood, Wilson & Cooley 9169 (A)}. Magnifications as indicated. 96 Sipa 17(1) with mid-stem leaves measuring 14-23 mm long, 9-11 mm wide. How- ever, one GH and the NY sheet contain stems with larger, thinner, more elliptical, acute-tipped, mid-stem leaves, 22—25 mm long, 7-10 mm wide; these elliptical leaves are similar to those occurring in other specimens in western Florida. Other collections from the Ocala region (Sma//, Mosier & DeWinkeler 10810) and from Levy County (Holland & Mears s.n., Murril s.n., Garber s.n.) are also small, to 2.5 dm tall, but have all opposite, more lanceolate to elliptical, acute leaves 2—3.5 cm. Still other collections (Woods, Wilson & Cooley 9169, Curtis s.n.) are taller, have ovate, obtuse- round-tipped leaves at the base, but more lanceolate leaves above and the leaves subtend- ing the inflorescences may or may not forma single whorl of 4 leaves—this character varying from plant to plant. Through these and other specimens one sees that there is continuum from small plants with all ovate, obtuse- tipped, opposite leaves, to plants that have smaller, broadly ovate, obtuse- tipped leaves at the lower nodes, and larger more lanceolate, acute-tipped leaves at the mid and upper nodes that are all opposite in some specimens but form single whorls of leaves below the inflorescences in others. Sell other plants have large lanceolate, elliptical-lanceolate leaves and well de- veloped whorls of leaves below the inflorescences. All these plants have om similar inflorescences and flowers. The small, broadly ovate, obtuse-tipped leaves characteristic of the type collection (Fig. 2a) initially stand in strong contrast to the larger, elliptic, acute-tipped leaves found on other spigelias in Florida (Fig. 2c). However, the larger plants with lanceolate, acute-tipped leaves that form a whorl of leaves below the inflorescence typically produce short-ovate leaves on the lower stems, although sometimes they will have fallen from the plants by the time of flowering. It seems reasonable to consider the shorter, blunt- tipped leaves as characteristic juvenile foliage of the species. In the collec- tions from the type locality, it appears that flowering occurs on specimens with only juvenile foliage. This can be considered a form of paedomorpho- sis, Le. the retention of juvenile characteristics on mature plants. The ten- dency to flower while having only juvenile leaves is probably fixed geneti- cally. There is also the possibility that if the plant develops late in the season and flowering triggered earlier, the plants may have only produced the shorter, basal leaves by the time of flowering. In this study, both the ovate-leaved and elliptical-leaved specimens are considered to be the same taxon. One could argue that ovate-leaved and elliptical-leaved specimens represented distinct taxa, but this would not be supported by the presence of plants with both leaf types and the overall continuum of variation from the extreme, represented by the type speci- men (Fig. 2a), and other larger specimens from Florida (Fig. 2c) as well as ja Texas (Fig. 54), Henrickson, Notes on Spigelia 97 Fic. 3. Spigelia loganioides and S. hedyotidea from Texas and Mexico. A-C. S. loganioides from Texas. A. Stem showing paired large cauline leaves, quartet of leaves subtending first lower of dichasia, leaves, fruits, and flowers in upper branches of dichasia. B. Node showing deltate stipule, leaf bases, and distinct decurrent ridges extending down glabrous intern- odes (A-B from Killip 43289, TEX). C. Flower showing bractlets, calyx and corolla (Corre// & Johnston 17503, LL). D-G. S. hedyotidea from Texas. D. Stem showing smaller opposite leaves, dichotomously branched inflorescences, flowers, and fruit. E. Node showing deltate stipule, leaf bases, and decurrently ridged internodes with distinctive scabrous papillae on cae and lower leaf margins (D—E from Smith 507, TEX). E Open flower showing bractlets, calyx, corolla; note broader sepals. g. Fruit showing ver cocci, remnants of calyx and nectary at base and persisting style base at fruit tip (F—G from C ‘sprall Rollins & C ‘hambers 16055, LL). Magnifications as indicated, scale with A holds for all stems; scale with F holds oe all fowers and fruit. 98 Sipa 17(1) \ i oe _* } x r ; i f Hi ee an te | ie : ee Niece oa bs es \ : \ 4 7 | a — ‘ 1 — “t are : S. loganioides o S. hedyotidea e¢ 500 miles Fic. 4. Documented distribution of Spige/ta loganioides in Florida and Texas and S. hedyotidea in Texas and Mexico. The plants from Texas known as Spigelia texana are identical to those of Florida and are considered conspecific with the older S. /oganivides. The Texas specimens are more consistently larger with a quartet of leaves below the inflorescences. The species has a disjunct distribution (Fig. 4); it is known from lime- stone areas in Levy, Marion and Sumter counties in west-central Florida where it occurs in wet woodlands, hydric hammocks, and floodplain swamps. It also occurs in Brazoria, Colorado, De Witt, Gonzales, eon Polk, Victoria, Waller and Washington counties in eastern Texas in black-clay soils, thickets, woods along creeks, and riparian forests. Soils in both of these areas are largely derived from limestone and chalk parent material. The taxon does not occur in the broad region between Florida and Texas as limestone-chalky parent material is not present in this region. There are a number of other taxa that have similar disjunct Florida-Texas distributions including Dyschoriste, Cladium, and Samolus. Representative egies U.S.A. Floripa. Levy Co.: ca 1 mi SW of Gulf Hammock, T14S, R1GE, sec. 29, 3 May 1959, Wood, Wilson & Cooley 9169 (A); a a a 0? May 19¢ Marrill n.(F TAS) Gulf pee just W of Griffin Creek, T ,RI4E, sec. 8& 17, 22 May 1980, Judd et al. 2660 (FLAS); 6 mi W of Otter Creek, ‘ ee Hwy ys 5 Jul 1979, Dunevitz 29 (FLAS); 6 mi W of. ene Creek, between Ortrer Creek and Rosewood, 29 May 1976, Godfrey 76520 (GH, NY, FLAS-phoro); Spee Jun 1876, Garber s.n. (FLAS, NY-2, US-2); C.R. 326 at Wekiva Run, E of US 16 Jul 1989, Holland & Mears sm. (USF). ae on Co.: Oklawaha River, E of Ocala, ‘1 Apr 1923, Small, Mosier & DeWinkeler 10810 (FLAS, GH, NY, US). Sumter Co.: near Sumterville, Jun, Curtiss 2258 HENRICKSON, Notes on Spigelia 99 (GH, NY-2, US). Texas. Brazoria Co.: woods along Cedar Lake Creek, San Bernard Natl. Wildlife Ref., 8 May 1970, Fleetwood 9753 (TEX); Crowell Ranch, 31 May 1970, Fleetw' - 9789 (TEX); Lake Jackson, woods along Oyster Creek, 2 May 1953, Killip 43289 (SM TEX). Colorado Co.: Columbia, 5 Oct 1900, Bush 1288 (NY, US). De Witt Co.: 7 mi - of Cuero, 6 Jul 1957, Correll & Johnston 17503 (LL, TEX). Gonzales Co.: Ottine Swamp, 8 Aug 1935, Parks 14603 (GH); 4 mi ESE of RR 532 near US 90A, Kokernot Ranch on Peach Creek, 12 May 1991, Ginsbarg 941 (TEX). Polk Co.: Menard Creek, near Trinity, 28 Apr 1981, Watson 3250 (SMU). Victoria Co.: Guadalupe River bottom, 29 May 1932, Tharp s.n. (TEX). Waller Co.: along Brazos River, Stephan F. Austin State Park, near Sealy, 25 a 1957, Correll 16444 (LL). Washington Co.: Burton, 26 May 1872, Hall 288 (NY-3, US). Spigelia hedyotidea A.DC., Prod. 9:7. 1845. Type: MEXICO: without loca- tion, date, Sessé & Mociio, plate No. 813 (HoLorype: illustration 813 (deCandole’s number) at G (microfische!), photo of holotype F!, US! Described by A. deCandole from a copy of a Sessé and Mocifio drawing, which remains at G, with photographs at F and US. No original drawing 1s ae at the Hunt Institute for Botanical Documentation (Kiger, pers. comm.; see Kiger 1981; McVaugh 1982). Two ae ments of authentic Sessé and Mecmo material labeled Spigelia axillaris n., No. 58 and No. 392 were found in F, these fragments were presumably removed Gn speci- mens borrowed from MA (McVaugh 1977, 1980). The “axillaris” combination was not published in either Flora Nuevo Hispanica (1878-1891) or the Flora Mexicana (1891-1897). These specimens exactly agree with the illustration upon which S. hedyotidea is based and are comparable to S. /indheimeri A. Gray; their source from éxico is discussed below. Spigelia lindheimeri A. Syn. fl. N. as 2(1):108. 1878. Coelostylis page Fl. s.e. U.S. 922. 1903. Type: U.S.A. Texas. Comal Co Braunfels, rarely on ee silt soil, damp prairies, Jun 1846, F Lindheimer 172 fee ToTYPE, here designated, GH!). Gray’s (1878) citation of the locality as “Prairies of . Texas, Lindheimer, Wright” made available many annotated mee at GH; the designated lectotype has an attached note from Lindheimer stating “distinct from S. texana?, fls. apparently half as large.” The specimen has narrow to broad, relatively large ipneeolare leaves, mature fruit, and ae Spigelia coulteriana Benth., J. Linn. Soc., Bot. 1:90. 1857. Type: MEXICO. Hipaico: Zimapan, Coulter 962. (HOLOTYPE: KI}, ae of holotype: NY!). Long considered a species having pseudospicate inflorescences, close examination of the type reveals that the nodes have 2 flowers and belong within f oelostylts. Multistemmed, often decumbent, bushy herbs 5—15(—19) cm tall; stems usually scabrous with erect, stiff, papillae 0.05—0.07 mm long throughout the decurrent internodal wings and usually on the internode surfaces be- tween the wings, on the stipules and upper leaf surfaces or margins; inter- nodes (0.6—)1.5—3.5 cm long. Leaves all opposite, very rarely in whorls of 4 below the inflorescence, the lowermost leaves usually ovate, obtuse to rounded at the tips, to 15 mm long, petiolate, the lower mid stem leaves often oblanceolate, obtuse, the mid- and upper-stem leaf blades lanceolate, narrowly lanceolate, lance-elliptic, (1.2—)1.5—3(—3.5) cm long, 0.3—-1(-1.3) cm wide, acute, apiculate at the tip, (sometimes all leaves oblong-ovate, LOO Sipa 17(1) and blunt tipped) the blades narrowly cuneate, tapering to short petiole- like base 1—3(—4.5) mm long, the margins entire to undulate, often revo- lute or enrolled when dry, the blades often stiff and coriaceous, the upper surface strongly punctate with cystoliths and with stiff-walled papillae particularly along the margins, typically appearing wrinkled when dried, the lower surface glabrous, gray-green (in shaded plants the leaves often more membranous). Flowers produced in terminal, leafy dichasia with two flowers per node, the shorter flower with an ebracteolate pedicel 1—3 mm long, the taller lateral flower with a bracteate peduncle-pedicel 6—10(—14) mm long, the bractlets subulate-linear, 2.5—7 mm long, 0.4—0.6 mm wide; sepals linear, linear-lanceolate, 3.4—8.5 mm long, 0.5—1(—1.5) mm wide at base, acute at tip, often scabrously papillate along the margins and some- times along the midveins; corollas white, suffused or lined with pink exter- nally, funnelform, 10-13.5 mm long, the tubes cylindrical to slightly ampliate, (S—)7—9 mm long, to 2—3.5 mm wide at the throat, the lobes obliquely oblong-lanceolate, 3—4.8 mm long, 1.5—1.9 mm wide; stamens borne in the mid-upper corolla tube, the filaments 0.9-1.2 mm long, the anthers 1.0—-1.2 mm long; styles 6-7 mm long, the persisting style base 1.2—2 mm long. Fruit cocci 2, nearly spheroid, very slightly divergent, glabrous, the pair 2.5—3.7 mm high, 5-6 mm wide; subtending thickened isk 3.2—3.5 mm wide, 3.5—5 mm long; seeds 6-7 per cocci, 1.5—2 mm long, 1.2-1.6 mm wide (Figs. 3d—g, 4). Spigelia hedyotidea can be distinguished from S. /oganioides in that S. hedyotidea has: (1) a smaller, bushy growth habit with many stems develop- ing from the basal rootstock; (2) a much smaller, mostly narrowly lan- ceolate, often thicker leaves that are usually papillate and rather strongly wrinkled on the upper blade surface when dry; (3) a more strongly papil- late epidermis with papillae occurring throughout the internode wings (not just below the nodes), and usually over much of the internode surfaces, on the upper-leaf surfaces and stipules; (4) consistent opposite leaves; it only rarely forms a whorl of four leaves below the inflorescences; and (5) narrower, less ampliate corolla tubes. However, S. hedyotidea is quite variable and there many exceptions to the above-noted characteristics. While open-grown plants typically are short and bushy with rather firm, coriaceous leaves, plants collected in shady habitats tend to have longer internodes, less coriaceous, and often larger leaves that may press flat and appear quite smooth and membranous in specimens. Some specimens have short, broadly ovate, more strongly peti- olate leaves throughout the stem (reminescent of the condition in Florida with the type of S. /oganioides). The plants from San Luis Potosi and Hidalgo, and some from Texas, are quite glabrous with papillae occurring only along Henrickson, Notes on Spigelia 101 the lower margins of the leaves, on the stipules, but they typically occur all along the internode wings. The small, more glabrous specimens from San Luis Potosi appear very much like specimens of S. cou/teriana, a species long considered to range from Hidalgo to Guatalmala (Gibson, 1969). However, the specimens from southern Mexico and Central America have elongate pseudospicate inflorescences characteristic of the true Spigelias. Careful examination of the type of S. cou/teriana from Zimapéan, Hidalgo, revealed an inflorescence type characteristic of the Coe/ostylis group and thus S. coulteriana is here considered synonyous with S. hedyotidea. While most specimens of S. hedyotidea are strongly papillate, the type of S. cowlteriana is very weakly papillate. It is not possible to use this characteristic to recognize two taxa here as many other Mexican collections are also weakly papillate, as are some collections from Texas. The oldest name for this taxon turns out to be Spigelia hedoyotidea A.DC described in the Prodromus in 1845 from a drawing from Sessé and Mocino in possession of deCandole. The holotype of the taxon remains the drawing of the species at G. Original Sessé and Mocifio herbarium collections of this taxon presumably exist at MA, and fragments at F. The collected speci- mens are very similar to the type specimens of S. /indbermerz. It is not known exactly where the collected material was obtained for neither Sessé or Mocifio visited areas where this species grows (McVaugh 1977). McVaugh (1977, p. 110), however, notes that at least two other Sessé and Mocifio species were obtained from the Texano-Mexican frontier region from Ignacio Leén, a pharmacist, who sent seeds and plants to Sessé from his station in Valle de Santa Rosa, (Santa Rosa de Mtizquiz) now known as Muzquiz, Coahuila. This is well within the range of the taxon and probably is the source of the material in the Sessé and Mocifio herbarium and the source of the illustra- tion, a copy of which was used by deCandolle in his original description. The taxon is known from open gravelly, sandy clay loams, dark-soiled praires, limestone slopes, limestone bluffs on generally dry soils in chapar- ral thickets, mesquite thickets, mixed desert scrub, shaded woodlands, river banks, and rocky creek beds, from central Texas to Chihuahua and Coahuila, San Luis Potosi and northern Hidalgo in México. Specimen data indicates that the flowers are closed in the morning (McVaugh 8342) and open in the afternoon (Jones 537). Representative specimens: U.S.A. TEx ander : Hill Senate: State Natural Area, 29°38’48"N, 99°12’45"W, 4 ee oo eee ae (TEX). Bell Co.: near Little River, 13 May 1930, Wolf 2110 (US). Bexar Co.: mesquite thickets, San Antonio, Apr 1884, Havard 13 (GH). Comal Co.: New Braunfels, May—Jun 1847, oe 43 (GH). Guadalupe Co.: Seguin, 28 Apr 1942, Parks 39529 (TEX). Hayes Co.: San Marcos, 1 Jun 102 Sipa 17(1) 1917, Palmer 12122 (NY, US). Karnes Co.: 5.5 mi SW of Karnes City, 19 Apr 1957, Correll, Rollins & Chambers 16055 (LL). Kendall Co.: shady woodland below Edge Falls, 3 ay 1947, Tharp, Mericle, & Barkley 17T139 (GH, TEX). Kerr Co.: Kerrville, 7-14 May 1894, Heller 1719 (NY, US). McLennan Co.: Middle Bosque, Ist crossing, north bank, 2 May 1947, Smith 507 (TEX). Palo Pinto Co.: hills above Possom Kingdom State Park, 17 May 1947, McVaugh 8342 (SMU, TEX). San Patricio Co.: 7.5 mi S of Taft, 29 May 1951, Jones 537 (SMU). Terrell Co.: Creek bed, Independence Creek, 1 mi above its junction with Pecos River, 22 Jun 1949, Webster 392 (TEX). Tom Greene Co?: Middle fork of Rio Concho, Apr 1886, Reverchon s.n. (GH). Travis Co.: Austin, 9 May 1936, Tharp s.n. (TEX- 2). Uvalde Co.: Uvalde, 11 May 1918, Pa/mer 13565 (US). Val Verde Co.: bank of Devil’s River at Fawcett Lodge, 20-30 mi up river, 3 Apr 1953, Warnock 11345 (LL, SMU). MEXICO. Chihuahua: Santa Eulaia (sic.) Mts., | May—6 Jun 1885, Pringle 25 (GH); Sierra del Roque, NNE of Julimes above Mina Las Playas, 28°39-40'N, 105°18-19'W, 19 Jun 1973, Johnston, Wendt, & Chiang 11389A (LL). Coahuila: Sierra de la Madera, Cafién de la Madera, 1890 m, 27°07’N, 102°37'W, 29 May 1975, Wendt & Lott 842A (LL); Sierra de la Gloria, Cafién El Cono, a side canyon of Cafién Chilpitin, 1160 m, 26°49’N, 101°17°W, 6 Sep 1976, Wendt & Riskind 1054 (TEX). San Luis Potosi: San Dieguito, 13-16 1904, Palmer 86 (NY, US-2); Mcpio. Valles, Rio Valles at El Banito, 7 mi S. Valles, 26 Jun 1940, Leavenworth 185 (BP). ACKNOWLEDGMENTS I thank Richard Hilsenbeck (Florida Nature Conservancy), Richard Wunderlin (USF), and Laurence Dorr (US) for information concerning Florida populations of Spigelia loganioides and historical collectors, Robert Kiger (Hunt Institute for Botanical Documentation) for searching through the Sessé and Mocifio collections for original illustration of Spigelia hed yotidea, Katherine Gould (TEX), who is now monographing Spigelia, for comments on the manuscript, Bobbi Angell (NY) for the plant illustrations, Roberta Wilson (TEX) for the inflorescence illustrations, Thomas Wendt (LSU) for the Spanish translation of the abstract, and A, FLAS, GH, NY, SMU, TEX- LL, US for specimen loans, and the Plant Resources Center at the Univer- sity of Texas, Austin for use of facilities. REFERENCES De Cannot, A. 1845. Ordo CXXXI, Loganiaceae. Prodromus systematis naturalis regni vegetabilis. 9:1—37 ENDLICHER, S.L. 1841. esawsephia generum plantarum. (9): t sae - 1878. Order LXX XIX. Loganiaceae. Synoptical flora oe h America 2(1):106— a. y, R.K. and J.W. Wooren. 1981. cy and wetland plants of soucheastern United States. University of Georgia Press, Athen Hurtey, H. 1968. A taxonomic revision oe genus Spigelza (Loganaceae). unpubl. dissert., George Washington University, Washington, Kicer, R.W. 1981. Long-lost drawings from Sessé ad Mocifio expedition acquired by Hunt Institute. Syst. Bot. 6:189— McVaucu, R. 1977. Botanical result of the Sessé and Mocifio expedition (1787-1803). I. Summary of excursions and travels. Contr. Univ. Michigan Herb. 11:97—195 HENRICKSON, Notes on Spigelia 103 McVauGu, R. 1980. Botanical result of the Sessé and Mocifi pedition (1787-1803). II. The icones florae Mexicanae. Contr. Univ. Michigan Herb. 14:99-140. McVauGu, R. 1982. The ee paintings of the Sessé & Mocifio expedition: a newly avail- able resource. Taxon 31:619-692. Torrey, J. and A. Gray. 1839. In: S.L. Endlicher. Novarum stirpium decades 5:33— Torrey, J. i A. Gray. 1841. Suborder IH. Loganieae R. Br. Flora of North Aare 2(1):43-4 WUNDERLIN, R.P. 1982. Guide to the vascular plants of Central Florida, University Presses of Florida, Tampa. SMALL, J.K. 1903. Flora of the southeastern United States. Privately published. New York. 104 Sipa 17(1) BOOK NOTICES Z a : cg Lapp, Douc. 1995. Tallgrass Prairie Wildflowers. ISBN 1-56044-299- 9, pbk). Falcon Press Publishing Company, P.O. Box 1718, Helena, MT 59624. $19.95. 262 pp This guide covers the more common and characteristic plants of tallgrass prairies from northeastern Oklahoma north into Canada and eastward through Ohio. Included ... are {about 320} photographs {by Frank Oberle} ... of 295 tallgrass prairie | plants, including a few examples of very rare plants that epitomize the tallgrass prairie.” The seven main sections of the book, which present the wildflowers arranged by flower color, follow the standard introductory material (e.g., description of area covered and plant morphology) and precede the glossary, a useful “Tallgrass Prairie Directory, “Selected Further Reading,” and an index. Accompanying each photograph is text describing the plant, flowering time, and habircat/range. The photos (one or two per page) are, almost without expection, su- perb; in conjunction with the brief descriptions, they will make identification of the target plants an easy task. This book should be required equipment for anyone visiting the callgrass prairie. Even elsewhere, though, it is a pleasure to read through.—John W. Thieret. Dunmire, WittiAM W. and Gait D. Tierney. 1995. Wild Plants of the Pueblo Province. Exploring Ancient and Enduring Uses. (ISBN 0-890 13-272-0, pbk.). eau of New Mexico Press, P.O. Box 2087, Santa Fe, NM 87504. $19.95. 290 pp. “A region of enormous diversity,” “The ancestral Puebloans,” “Contact, C olonization, and a oe “Modern descendants,” “Indicator plants as living artifacts,” “Plants and planccraft,” ‘Ethnobotany 1 in New Mexico,” “Recent modifications in the ie and “Other places to visit,” these chapter titles indicate the wide and full coverage of this fine production, an exemplary historico-ethnobotanical volume. Abundantly illustrated, the work is well produced and will indexed. Coverage of relevant literature is good (1 oe ” references in the “Bibliography”) “Plants and plantcraft,” the book’s longest section (1 pages), considers 55 of the most important species, oes with its own text, color oe and line drawing. Data on these species plus about 240 others are conveniently summa- rized in a 12-page table. The authors and the Museum of New Mexico Press have done a superb job.—John W. Thieretr. RICHARDSON, ALFRED. 1995. Plants of the Rio Grande Delta. (ISBN 0- 292-77070-7, pbk). University of Texas Press, RO. Boc 7819, Austin, TX 78713-7819. Price not indicated. 332 pp. This volume “expands and updates” Dr. Richardson's Plants of Southermost Texas (1990), an identification guide to the Hora (excluding the grasses) of Cameron, Hidalgo, and Willacy counties (see notice in Sida 15:170. 1992). Most at the text is a repeat of that in the earlier work, but new and welcome indeed is the array of 94 beautifully reproduced color plates bound together at the end of the book. They greatly increase the book’s value for its stated purpose.—John W. Thieret Sipa 17(1): 104. 1996 A NEW COMBINATION IN ERAGROSTIS (POACEAE: ERAGROSTIDEAE) PAUL M. PETERSON Department of Botany National Museum of Natural History Smithsonian Institution Washington, DC 20560, U.S.A. ABSTRACT The new combination Eragrostis pectinacea vat. tracy? is proposed as a more appropriate ae means of recognizing the slight morphological difference of possessing longer anthers. A revised key to the three varieties of E. pectinacea, a diagnosis of var. ¢racy7, and specimens examined are provided. RESUMEN Se propone la nueva combinacién Eragrostis pectinacea vat. tracyi como mas apropiada para reconocer las ligeras diferencias morfolégicas de tener anteras mds largas. Se ofrece una clave revisada de las tres variedades de E. pectinacea, una diagnosis de la var. tracy7, y los especimenes examinados. The genus Eragrostis consists of approximately 350 species worldwide and, of these, about 120 species are represented in the western hemisphere (Peterson et al., in press). The present biogeographical pattern supports the hypothesis that the genus arose in the Old World, probably in southern Africa where it is most speciose. The best combination of characters used to discriminate Eragrostis from other Eragrostideae is: disarticulation of the lemma and palea occurring separately; longitudinally bowed-out paleas with ciliolate keels; and 3-nerved, unawned lemmas. Within Eragrostis the spe- cies limits are often overlapping. Few agrostologists have attempted to work out a suitable phylogenetic scheme. Based on 442 morphological and anatomical characters, and using a worldwide sample of 53 species, Van den Borre and Watson (1994) recognized two subgenera, Eragrostis and Caesiae. The new combination appears to reside in the former. While preparing the Eragrostis treatment for the new Manual of North American Grasses, Edited by M. Barkworth, (Peterson & Harvey, in press) I became aware of the similarities between E. tracys Hitchc. and the common purple lovegrass [E. pectincea (Michx.) Nees}. The following paper provides a rationale for the new combination of E. pectinacea var. tracyt (Hitchc.) P.M. Peterson. Hitchcock (1934) first described E. tracy from specimens collected by S. Sipa 17(1): 105-107. 1996 106 Sipa 17(1) M. Tracy on Sanibel Island. In his paper he wrote, “apparently perennial” and in his Latin description he indicated “Perennis (?).” Koch (1972) re- evaluated E. tracys and found it to be an annual with a hexaploid chromo- some number of 7 = 30, x = 10. Koch (1972, 1978) concluded that E. tracyl was Closely related to E. /utescens Scribn., E. pectinaea, and E. tephosanthos Schult. Koch (1972) found that E. tracy could be distinguished from these other species by its larger anthers. Eragrostis pectincea and E. tephrosanthos have previously been determined to be hexaploids at 7 = 30, x = 10 (Koch 1974), More recently, Reeder (1986) recognized two varieties in E. pectinacea, var. pectinacea and var. miserrima (E. Fourn.) Reeder. The latter variety in- cludes E. tephrosanthos as a synonym. I agree with Reeder’s treatment of placing E. tephrosanthos as a synonym of E. pectinacea. Eragrostis pectinacea is a highly variable taxon as described by Koch (1974). Anther length is the only morphological characteristic that can be used to differentiate E. tracy: from E. pectinacea (0.5—0.7 mm long in the former and 0.2—0.4 mm long in the latter). To remain consistent with the new broader delimitation of E. pectinacea and since a single morphological trait separates these two taxa, I feel that E. tracyi should be reduced to the varital rank of E. pectinacea. A key to the three varieties of E. pectinacea, a diagnosis of var. tracyi, and specimens examined in the U.S. Herbarium are given below. KEY TO THE VARIETIES OF ERAGROSTIS PECTINACEA Li a 0.5—0.7 mm long var. tracyi 2. Anthers 0.2—0.4 mm long. 2. Pedicels appressed or rarely diverging up to 20° from the branches .... var. pectinacea 2. Pedicels widely spreading var. miserrima Eragrostis aaa ae (Michx.) Nees var. tracyi (Hitchc.) P.M. Peterson, ‘comb. nov. Eragrostis tracy? Hitche., Amer. J. Bot. 21:130. 1934. Type: U.S.A. Fioripa. Lee on Sanibel Island, 19 May 1901, Tracy 7168 (HOLOTYPE: US!; isoryPE: US!). Panicles with spreading branches 20—80° from the culm axis. Anthers 0.5—0.7 mm long Ecology and Distribution.—Sandy soils, along shell beaches, and roadsides: associated with Eragrostis ciliaris (L.) R. Br., Agave, Juniperus, Sabal, and Yucca; 0-30 m elevation; Lee, Manatee, Pinellas, and Sarasota cos., Florida: flowering March through May and August through December. Common Name.—Sanibel lovegrass. Representative specimens (all at US). U.S.A. Florida. Lee Co.: Sanibel Island, Jul-Aug 1900, Hitchcock 5.n.; 28 Dec 1953, Cooley 2495; 13 Apr 1954, Cooley 2608; 11 Mar 1971, Koch 7123, Western ee , Brumbach 7721, 3 Nov 1971, Eastern Sanibel, Brumbach 7767, 13 Dec 1971. Mantee Co.: ne a Maria Key, 24 Sep 1968, Harvey 8174; Longboat Key, 27 Apr 1900, Tracy 705: 24 Sep 1968, Harvey 8177; 9 Mar 1971, Koch 7113; 23 Aug 1971, PETERSON, Eragrostis 107 Godfrey 70891. Sarasota Co.: Longboat Key, 6 Oct 1964, Lakela 27566; 21 Nov 1964, Godfrey 65243; Venice City, 10 Mar 1971, Koch 7116. ACKNOWLEDGMENTS John F. Pruski and Harold Robinson are thanked for reviewing the manu- script. REFERENCES Hrrcucock, A.S. 1934. New species, and changes in nomenclature, of grasses of the United States. Amer. J. Bot. 21:127-139. Kocu, S.D. 1972. A re-evaluation of the life cycle of Eragrostis tracyi war ea Eragrostoideae) and its taxonomic implications. J. Elisha Mitchell Sci. Soc. 88:21 Kocu, 8.D. 1974. The Eragrostis adaed -pilosa complex in as and Central America (Gramineae: Eragrostoideae). Ilinois Biol. Monogr. 48:1— Kocnu, $.D. 1978. Notes on the genus a eee tenn in othe southeastern United States. Rhodora 80:390—403. PETERSON, P.M. and L.H. Harvey. In press. Eragrostis. In: M.E. Barkworth, ed. Manual of North American Grasses. 83 manuscript pp PETERSON, P.M., R.D. Wesster, and J. VaLpes-ReyNa. In press. Genera of New World Eragrostideae (Poaceae: eae Soar bsonien Contr. Bot. 215 manuscript pp. REEDER, a R. 1986. Another look at Eragrostzs tef (Gramineae). Phytologia 60:15 3-— 154 VAN DEN Borre, A. and L. Watson. 1994. The infrageneric classification of Eragrostis (Poaceae). Taxon 43:383—422 108 Sipa 17(1) BOOK NOTICES SWANSON, Ropert E. 1994. A Field Guide to the Trees and Shrubs of the Southern Appalachians. (ISBN 0-8018-4556-4, pbk.). The Johns Hopkins ean! Press, 2715 North Charles Street, Balti- more, MD 21218-4319. $18.95 (pbk); $55.00 (hbk). 399 pp. The area covered by this work is “the mountains and higher foothills of western North Carolina, upper South Carolina, Northeast Georgia, and Great Smoky Mountains National Park.” Well accounted for in this book are the area's 180 genera and 280 species of woody plants. An introductory section covers morphological features useful in identification. Carefully constructed keys are provided for both summer and winter identification. Ar- ranged by families, the main part of the book is devoted to clear illustrations (black-and- white line drawings by Frances R. Swanson) and detailed descriptions of the various ae cies. The volume closes with a glossary, a list of references (10 of them), and an index. recommend the work to anyone planning a visit there. It worked well for chose 20 species I chose as test cases. Kudos to the author.—John W. Thieret. Daumer, Frep. 1995. Caddo was ... A short History of Caddo Lake. (ISBN 0-292-72479-9, pbk.). University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819. $8.95 80pp Caddo Lake, the largest natural freshwater lake in Texas, covers ca. 26,800 acres and straddles the Texas/Louisiana border. This book on various aspects of the lake’s history, was first published as a series of essays in the 1980s, then as a book in 1988, and now as a “revised” edition in 1995. It includes but lictle botany, nor claiming to be a source of data on the lake’s biota. Some of the photographs (all black and white) are of magnificent, Spanish-moss-draped bald-cypresses; and a short chapter has a few data on duckweeds, waterlillies, goldenclub, spatterdock, and American lotus. Having been many times to Caddo Lake, I enjoyed reading Caddo was.—John W. Thieret ~ Sipa 17(1): LO8. 1996 A NEW COMBINATION IN BOUTELOUA (POACEAE) JOSEPH K. WIPFF and STANLEY D. JONES Herbarium (BRCH) Botanical Research Center PO; Boxe ost 7 Bryan, TX 77805-6717, U.S.A. wipftf@bihs. net or sdjones @ bihs. net ABSTRACT The new combination Bowteloua hirsuta Lag. subsp phan (H. Featherly) J. Wipff & S.D. Jones is proposed. A key to the two subspecies is provide RESUMEN e propone la nueva combinacién Bonteloua hirsuta Lag. subsp. pectinata (H. Featherly) J. S Wipff & S.D. Jones. Se ofrece una clave de las dos subespecies. Bouteloua hirsuta M. Lagasca y — subsp. pectinata (H. oe Wipff & S.D. Jones, comb. et stat. nov. Basionym: Bouteloua pectinata H Featherly, Bot. Gaz. 91:103. 1931. ee hirsuta var. pectinata (H. Featherly) V. Cory, Rhodora 38:405. 1936. Type: UNITED STATES. OKLAHOMA. Comanche Co.: near Fort Sill, 17 Aug 1929, B. English 71 (HoLoTYPE: US Bouteloua pectinata, closely related to B. hirsuta, has been recognized at various taxonomic ranks: Roy and Gould (1971) and Gould (1975, 1979) recognized this taxon as a distinct species; Cory (1936) recognized it as a variety of B. hirsuta; and Hitchcock (1935) and Chase (1951) treated it as a synonym of B. hirsvta without any infraspecific rank. The two taxa can be distinguished by the following characters (Roy & Gould 1971): L. Tuft of trichomes present at the base of the lowermost rudimentary floret; anthers ¢ long; culms 37-75 cm tall, strictly erect, unbranched, usually with three nodes; the inflorescence axis 25—40 cm long (above up- subsp. pectinata permost culm lea . Tuft of trichomes absent at the base of the lowermost rudimentary floret; anthers 2—2.5 mm long; culms 15—40 cm tall, decumbent at base, usually branched, with 4—6 nodes; the inflorescence axis 10-30 cm long (above — subsp. hirsuta uppermost culm A biosystematic study of B. pectinata and B. hirsuta by Roy and Gould (1971) resulted in the following conclusions: 1) B. pectinata is restricted to well-drained, relatively undisturbed calcareous soils and is most frequent on thin-soiled limestone outcrops, occuring from Pontotoc and Comanche counties, Oklahoma, south to Uvalde County, Texas; 2) Boutelouwa hirsuta, Sipa 17(1): 109-110. 1996 110 SIDA 17(1) found in a wide variety of habitats, is widely distributed and occurs from Wisconsin and Illinois to North Dakota, south to Louisiana, Texas, New Mexico, Arizona, southern California, and northern México; 3) the mor- phological uniformity of B. pectinata, 2n = 20 populations contrasts strik- ingly to the morphological variability observed in the diploid plants 2” = 20, of B. hirsuta. Usually associated with diploid B. Airswta are plants with widely oe chromosome numbers, e.g. 27 = 22, 24, 26, 28, 30, 32, 34, 36, 40, 42, 43, 44, 45, 46, 48, 50, 52, 53, 54, 56, 58, and 60 (Roy 1968); 4) where populaenns of the two taxa are sympatric, swarms of putative hybrid plants are found that are intermediate in some or all of the critical morphological characteristics; and 5) for the most part, these putative hy- orid swarms are found only along the margins of typical B. pectinata sites (when populations of B. a are also present). Sites in Kendall County, Texas appear to be representative of the ecological distribution and separa- tion of the two taxa. Roy and Gould (1971) stated, “Throughout this area the two species, B. hirsuta and B. pectinata, were intermingled in slightly dis- turbed road right of way sites. Bowteloua hirsuta predominated at the base of the slopes and in the gulleys [s/c] and ditches, and B. pectinata was most abundant on the ledges and hilltops. Intermediate plants were abundant.” ay ‘rom these data, B. pectinata is restricted in its geographic distribution, occupies a specific ecological site, and is generally ecologically allopatric and morphologically distinct from B. hirsuta. However, there are putative hybrid swarms at the boundary where the two taxa come in contact. These swarms show an integration of morphological characters between the two taxa indicating a lack of complete reproductive isolation. These data war- rant the recognition of B. pectinata at the subspecific rank. We are grateful to W.E. Fox, HI (TAES), Stephan L. Hatch (TAES), Gretchen D. Jones (USDA, AWPMRU), and Clifford W. Morden for re- viewing this manuscript. REFERENCES Criase, A. 2 1. A revision of A.S. Hitchcock’s, manual of the grasses of the United States, 2nd ed. S.D.A. Miscellaneous Publication 200. U.S. Government Printing Office, een Cory, V. 1936. New names and new combinations for Texas plants. Rhodora 38:404—408. GouLb, F.W. 1975. The grasses of Texas. Texas A&M Uni versity Press, College Station. GouLb, F WW. 1979. The genus Bowtelowa (Poaceae). Ann. Missouri oi G: 7 66:348—416. Hircucock, A.S. 1935. Manual of the grasses of the United States. U.S.1 2.A. Miscella- neous cone ation 200. U.S. Government Printing Office, Washington, eel GP. 1968: A ae study of the Bowteloua birsuta- Boutelona pine complex. 1.D. eee ition, Texas A&M University, College Stat ,G.P. and RW. Goutp. ee . Biosystematic investigations .) Mez, a la sinonimia de Myrsine peregrina (Mez) Pipoly. INTRODUCTION As part of the Flora of the Philippines Project, jointly undertaken by BRIT ae the Philippine National Herbarium, a revision of the Philippine species of Myrs/ne, including those taxa formerly assigned to Rapanea, was undertaken. The genus is now defined as pantropical, with approximately 300 species, of which nearly 1/4 remain undescribed. Myrsine is the type genus of the tribe Myrs/neae, defined by its uniseriate ovules and imbricate or valvate perianth aestivation. Recent studies (Pipoly 1992b, 1994) have shown that the principal technical characters defining the tribe are found rather at random in the tribe Ardsszeae, and biseriate ovules are most com- mon in the genus Parathesis Hooker f., and Sty/ogyne A. DC. Therefore, a worldwide generic monograph is needed before long-term nomenclatural stability is established. Pipoly (199 1a, 1992a, 1992b) and Pipoly and Chen (1995) have shown that characters traditionally used to separate Rapanea from Myrsinve grade and therefore cannot be used with confidence to define either group unequivocally, and have begun the process of transferring all species of Rapanea into Myrsine. NOMENCLATURAL HISTORY AND SYSTEMATIC POSITION OF MYRSINE The genus Myrsve Linnaeus, as here defined, includes Rapanea Aublet and Swttonia A. Richard, among other genera rarely recognized today (see Taxonomic Treatment). While no modern author has considered maintain- ing Swttonia (except Rock 1974), the discussion of the systematic position of species heretofore assigned to Rapanea has varied considerably from one author and world region to another. The first monographer of the family, Alphonse de Candolle, proposed uniting Rapanea with Myrszne in his generic level reviews of the family (1834a, 1834b), which he consistently followed in later “memoirs,” or analy- ses of the generic and tribal taxonomy of the family (184la, 1841b), as well as in his contribution to the DC Prodromus (1844). Koorders and Valenton (1900) followed the De Candolle philosophy in their treatment of the family for Java. Mez (1901, 1902) separated Myrsine from Rapanea based on the obvious style on the pistil and pistillode with obvious fila- ments of the stamens and staminodes in the former, and an apparently Pipoty, Synopsis of Myrsine in the Philippines Lag obsolete style with sessile anthers or antherodes (sterile anthers or ones that produce non-viable pollen) in the latter. Subsequently, in the most com- prehensive revision of the family covering Asian species since Mez, Walker (1940) maintained Rapanea as a genus distinct from Myrsine, by essentially paraphrasing Mez, a view concurring with Standley (1938) and Degener (1939). Oliver (1951) followed Degener as did Bathie (1953). However, Walker’s treatment of the group for Taiwan (1959) concluded that the two genera were in fact, synonymous, following the argument proposed by Hosaka (1940). Later, Li (1963, 1978) Meyer and Walker (1965), Wilbur (1965) and Backer and Bakhuizen van de Brink (1965) also followed Hosaka. Lundell (1966, 1971) in his treatments of the family for Guatemala and Panama, followed Mez’s arguments, but did not provide specific details to refute the transfers made by Stearn (1969) who was the first modern author treating tropical American taxa to synonymize Rapanea within Myrsine. Liogier (1971, 1989) who had earlier recognized Rapanea in his treatment of the Myrsinaceae for the Flora de Cuba (1959), also followed Stearn. In- deed, Lundell (1981) recognized Rapanea as synonymous with Myrszne, “to follow current practice, not out of any conviction as to the justification for this disposition of the taxa !” Subsequently, Lundell (1984) described two new neotropical species in Myrszne. The most cogent argument presented in the recent literature for main- taining Rapanea as distinct from Myrsine is that of A. C. Smith (1973, 1981), who based his decision on the fact that the Rapanea species he was familiar with did not have the “flange,” or apically free staminal tube, present in Myrsine africana L., the type species of Myrszne, but rather, the staminal tube was developmentally fused with the corolla tube such that the anthers appeared epipetalous. Smith’s philosophy was largely that of Degener (1939) and Oliver (1951) but neither of those authors had so eloquently stated their rationale. Sleumer (1986) followed A. C. Smith’s concept in his treat- ment of the taxa for New Guinea, as did Fournet (1978) for Guadeloupe and Martinique, Taton (1980) for Central Africa, Kupicha (1983) for the Zambesiaca Region, Halliday (1984) for Tropical East Africa, Little et al. (1988) for Puerto Rico and the Virgin Islands, and Green (1986, 1990) for the Norfolk and Lord Howe Islands’ taxa. On the other hand, Fosberg and Sachet (1975, 1980), and Sachet (1975) discussed their philosophy for including Rapanea within Myrsine, largely because they considered the staminal tube difference the only character separating the two groups. They proposed that the Rapanea group sensu stricto be recognized as a section within Myrsine, but did not formalize that position nomenclaturally. Correll and Correll (1982), Liogier (1989), Howard (1989), Nicholson (1991), and Guzman-Teare (1992) all followed 118 Sipa 17(1) Stearn (1969) and the Fosberg and Sachet papers. Wagner et al. (1990) presented essentially the same argument as Fosberg and Sachet (1975), but allowed for the possibility that future, geographically broad studies, might reveal that a small staminal tube difference may in fact, be important. Based on the study of at least representative taxa for the genus world- wide, Pipoly (199 1a, 1992a, 1992b), Pipoly and Chen (1995), Harvey and Pipoly (1995), and Chen and Pipoly (in press), along with my unpublished work, have shown that the species previously placed in Rapanea, including the type species, R. gwyanensis Aublet, have two vascular traces in the lower corolla tube, such that the removal of an anther from the corolla results in the tearing of the adaxial surface of the corolla tube without detaching the corolla lobe. As Pipoly (1981, 1983a, 1983b, 1987, 1988, 1991b, 199 1c, 1992c, 1992d, 1993, 1994) has demonstrated in the neotropical genus Cybianthus, a transitional series from a staminal tube free from the corolla tube to one where the staminal tube is morphogenetically fused in its en- tirety is easily seen within one myrsinaceous genus. In addition, new Ven- ezuelan, Colombian, and Peruvian species of Myrszne recently described by Pipoly (1991a, 1992a, 1992b) clearly show that some taxa possess fila- ments totally free from each other and also from the corolla tube, others have filaments briefly connate basally, but free from the adaxial corolla tube (thus, a “flange”), others again have filaments partially connate but wholly adnate to the adaxial corolla tube and lobe surfaces, and finally, still others have lower portions of the filaments forming a tube morphogenetically fused to the corolla tube, as well as having the upper portions of the same filaments morphogenetically fused to the corolla lobes. Clearly, these vari- able morphotypic gradations, perhaps subject to the influences of sex ex- pression, and a range of hormonal activity at times associated with reitera- tion phenomena, cannot be adequately used to characterize the taxa at a supraspecific level. Likewise, stigmatic and stylar characters have also been shown to vary in similar fashion, (Pipoly 1991a, 1992b) with one notable exception. To date, no ligulate stigmas have been found in neotropical taxa, although some taxa have been described with short styles (Pipoly 1992b). Therefore, I see no reason to recognize Rapanea as a distinct entity from Myrsine, even though some nomenclatural adjustments will have to be made to accomplish its total synonymy. In phylogenetic terms as well, it is more parsimonious to assume that the small, umbellate or fasciculate inflorescence structure, concomitant with the acropetally accumulating floral bracts along the perennating peduncle forming the “short shoot,” would have arisen only once. However, owing to the sheer size of the group and dearth of knowledge about its taxa in many of its most species-rich geographic areas, a full phylogenetic analysis must await the accumulation of further data from the taxa on a comprehensive, worldwide scale. Pipo.y, Synopsis of Myrsine in the Philippines LT NOTES ON MORPHOLGICAL TERMINOLOGY AND SPECIMEN CITATION Morphological terminology follows Pipoly (1987, 1991a, 1992b, 1992c) and Chen and Pipoly (in press). Specimens are cited according to island, north to south (e.g., Luzon, Mindoro, Panay, Palawan, Mindanao), then alphabetically by province within each island. Traditional divisions into regions, such as that used by Merrill (1923) is problematic, and is not used here, because some islands have since been reassigned from one region to another. When known, the collector, his personal collection number, fol- lowed by the herbarium number, is cited. In some cases, such as that of the Ramos and Edafio collections, specimens deposited at A, BISH, GH, K, L, NY, PNH, or US may have only the collector’s personal number, only the BS (Philippine Bureau of Science), the FB (Forestry Bureau) or the PNH (Philippine National Herbarium) number, or may have the personal and one of the institutional numbers. The institutional numbering has resulted in much confusion for the curator desiring to update determinations. In order to remedy this, I include both number types whenever I have been able to match them. TAXONOMIC TREATMENT Myrsine L., Sp. Pl. 1:196. 1753, Gen. Pl. ed 5:90. 1754; R. Brown, Prodr. 1:533. 1810; Ree ater sehulees, Syst. Veg. 503. 1819; ies oe Veg. 1:664. 1825. A. DC., Trans. Linn. Soc., London, Bot. 17: 104 4, Ann. Sci. Nat., Ser. 2, 9:292. 1834, Ann. Sc. Nat., Ser. 2 2, 16:65-97, reueee t. ve : a 1841; ‘A. DC in DC, Prodr. 8:92. 1844; Miquel in Martius, Fl. Bras. 10:306. 1856, Fl. Ned. Ind. 2:1014. 1856. Scheffer, Myrs. Arch. Ind. 46. 1867. Hooker in Bentham & Hooker, Gen PI. 2:642. 1876. Clarke in Hook. f. Fl. Brit. India 3:511. 1882., Pax in Engler & en Pflanzenfam. IV. 1:92. 1889; Koorders & Valeton, Bijdr. Boomsoort. Java 5:2 1900; Mez in Engler, Pflanzenr. 9(1V. 236):338. 1902; Nae ee of 73:1940, Bot. Mag. Tokyo 67: A 1954, Bull. Agri. Home Ec. v. Ryuku 2:76. 1955, Quart. J. ae Mus. 164. 1959; F. ase & E. Wa = EL Japan 713. 1965; Stearn, Bull. Brit. ae na Hist. ), Bot. 4:174. 1969; Fosberg & Sachet, Smithsonian Contr. — 21:3-11. 1975; H-L. Li, Fl. Taiwan 48. 1978; re, Phytologia 48:137. 1981, Phytologia 56 a 1984; pore Fl. Tr. E 6. 1984; A. Nicholson in D. Nicholson, Fl. Domin. 2:160. 1991; Pipoly, a 1:204. 1991, Caldasia 17:1. 1992, Novon a 1992; Guzman-Teare, Cat. Fl. Pl. & Gymno. Peru 732. 1993; Pipoly & Chen, Novon 5:360. 1995; Harvey & Pipoly, Fl. Pico das Almas 487. 1995; Chen & Pipoly, Fl. China 15:34—38. Un press). TyPE SPECIES (by monotypy): Myrszne africana L. — Aublet, Hist. Pl Guiane [:121. t. 46. 1775; A. Jussieau. Gene Phi 238 oo! uel in Martius, Fl. Bras. oe 1856; Mez in Urban, Symb. ae eee on Mez in Engler, Pllanzenr. 90V. 236):342. 1902; Pitan, in LeCom Indoch. 3(6):786. 1930: Standley, Publ. Field Mus. Nat , Bot. on ee 7 1938; Bathie, Fl. Madagascar 161:138. 1953; Lundell, sar Bot. 24(8—1):190. 1966, Fl. Panama 8:286. 1971; Fournet, Fl. Ilus. Phan. Guadeloupe et Martinique 1046. 1978; Taton, Fl. Afr. Cent. Myrs. 53. 1980; Kupicha, Fl. Zambesiaca 7:201. 120 Sipa 17(1) 1983; Halliday, Fl. Trop. E. Afr. Myrs. 2, 8. 1984; Little et al., Arbol. Puerto Rico y Islas Virg. 2:872. 1988 Samara Swartz, Prodr. 1:261 ae non io innaeus, Mant. 144, 199. 1771 Manglilla A. Jussieau, Gen. Pl. 89 Athruphyllum Loureiro, Fl. eee : 148. the Caballeria Ruiz et Pavon, Fl. Peruv. Prodr. 141. t. 30, 1794. Suttonia A. Richard, in Busnont dUrville, Déc. anit Bot. 1:349. en 2, et Ess. Fl. Nouv.-Zel. 349. pl. 1832; Léveillé, ms Spec, Nov. Reoni Veo 1 44 4.1912, Rock, Indig. Tree ne Is. 367-380. 1974 Shrubs or small trees. Leaves alternate, secu Inflorescences lateral (axillary), umbellate or fasciculate, sessile or on short, perennating peduncles girdled by persistent floral bracts (thus forming “short shoots”). Flowers 4—5(—6)-merous, bisexual or unisexual (plants then monoecious, dioecious, or polygamous); sepals nearly free or united to 1/2 length, imbricate or valvate, usually ciliate, punctate, persistent; petals nearly free or rarely united to 1/2 their length, usually ciliate, glandular-granulose at least along mar- gin and often throughout within, punctate; stamens and staminodes simi- lar, subequalling corolla length, the filaments free or connate basally to form a tube, the tube with or wien sterile appendages alternating with the filaments, and all merely adnate to the corolla tube; or developmentally fused throughout, the anthers thus appearing epipetalous, the anthers ovate or reniform, rarely sagittate, 2-celled, dehiscing by longitudinal slits, rarely by subterminal pores opening later into wide longitudinal slits. Pistil and pistillode similar; obconic, obturbinate, obnapiform, or variously subglobose; ovary globose, costate or not, glabrous or glabrescent; ovules few, uniseriate, or rarely biseriate, completely immersed in placenta or seated below apical pores in placenta or variously projecting; style obsolete to present, tapering into stigma; stigma morchelliform, liguliform, sinuate to lobate, prismatic and 3-lobed, or rarely, conical. Fruit a globose, subglobose, ellipsoid, ovoid, or sieve drupe, with somewhat fleshy exo- carp and crusty or leathery endocarp, l-seeded. Seed occupying cavity; en- dosperm horny, ruminate; embryo cylindric, transverse. Myrsine, as here defined, contains ca. 300 species, of which nearly 1/4 remain undescribed, distributed pantropically. They occupy habitats from mangroves to subalpine scrub, but always in moist, wet or pluvial habitats. In the Philippines, 14 species are known, separable by the following key. KEY TO MYRSINE OF THE PHILIPPINES 1. Flowers borne in sessile to subsessile umbels, the peduncle apparently not perennating, obsolete to 3.5 mm long. 2. Leaf blades 1.5—1.8(-1.9) cm wide. 3. Young branchlets and petioles rufous-puberulent or glandular-papil- lose; leaf blades dull to subnitid above, the midrib impressed above, prominently raised below, the punctations pellucid, inconspicuous. Prpoty, Synopsis of Myrsine in the Philippines eal 4. Leaf blades linear to ee oblanceolate; petioles marginate, not canaliculate; pedicels obconic, (1.5—)2—2.5(—3) mm long; young branchlets and petioles nae suberalene and tae ranulose. 1. M. medeciloae 4. Leaf blades elliptic to lanceolate; petioles deeply canaliculate, not marginate; pedicels cylindric, 5-7 mm long; young branchlets and petioles: rufous glandular-papillose, not puberulent or glandular- M. apoensis N= granu 3. Young branchlets and petioles glabrous; leaf blades nitid to laccate above, the midrib Lies ae raised above and below, the puncta- tions numerous, prom eC 5. Leaf blades symmetric, oblong or ailipae the apex obtuse to rounded, prominently retuse to emarginate at tip, the margins revolute; fruit . M. amorosoana jon globose. 5. Leaf blades asymmetric, narrowly oblong to narrowly wh ancec late. the acuminate but bluntly rounded at tip, the margins flat; fruit 4. M. oblongibacca oblongoid 2. Leaf blades > (1.8—)2 cm wide. Branchlets angulate; fruit depressed-globose, subglobose, or ovoid. 7. Leaf blade cartilaginous, laccate above, smooth above and below, the apex rounded to obtuse, the tip retuse to emarginate; pedicels straight, not reflexed in anthesis; fruit 5-6 mm diam. ............. 5. M. cruciata 7. Leaf blade coriaceous, dull or somewhat nitid above, sparsely to densely scrobiculate above and below, the apex short-acuminate to acutish or obtuse, not emarginate or retuse at tip; pedicels reflexed in anthesis; fruit 2.2-5 mm diam. 8. Leaf blade symmetric, apically acutish to obtuse; pedicels cylin- , 1.5-2.5 mm lon 6. M. densiflora ]; le nl 8. Leaf blade asymmetric, apically short-acuminate; p 3 long 6. Branchlets terete; fruit globose. 9. Leaf blade chartaceous, inconspicuously punctate above, the base acute to cuneate, decurrent to base of petiole; petiole 6-8 mm long; branchlets minutely rufous papillose and early glabrescent, or ie M. peregrina age at first. . Leaf blade scrobiculate above; branchlet apex minutely rufous papillose at first; calyx 0.8-1 mm long; plants of submontane forests. ‘ieiaciliail eos blade smooth above; branchlet apex glabrous; cali 1 ( 1.5 mm long; plants of mangrove swamps. ............-- 9.M = liepieees 9. Leaf blade coriaceous, prominently pellucid punctate above, the base obtuse to rounded, not decurrent on petiole to base; petiole 10-15 n long; branchlets densely rufous tomentose at first. ............ 10. M. avenis 1. Flowers borne in fascicles or umbels, on perennating peduncle-like short sa usually aad than 3.5 mm long at maturity. Flowers in fascicles; pedicels obsolete to 2.5 mm long; leaf blades sparsely Pe conspicuously, Sait not prominently punctate above 12. Floral bract margins erose-fimbriate, long white glandular-ciliolate; peduncles (3.5—)4-12.5 mm long; corolla lobes densely and promi- nently red-punctate along entire margin, densely and prominently Sipa 17(1) N bo pellucid punctate-lineate medially; anther connective red see 11. M. penibukana lineate; leaf margin flat. Floral bract margins entire, rufous glandular-ciliolate; pec ace 5 4 mm long; corolla lobes densely. and prominently black punctate medially, epunctate marginally; anther connective punctate; leaf olute. 12. M. fastigiata i) margin rev . Flowers in umbels; pedicels 2.6—6 mm long; leaf blades densely and — a aa or prominently punctate above . Branchlets 7-10 mm diam., drying anlenee: leaf blades thickly co- riaceous to cartilaginous; densely but not prominently pellucid or punctate; leaf buds covered by minute translucent or rubigi- rec nous glandular scales; petioles marginate and canaliculate, 5—10 mm long; without raised midrib; peduncle stout, (3—)3.5—4 mm diam.; pedicels 2.6—4 mm long; sepals delrate, 1.5—2 mm long. .... 13. M. aralioides 13. Branchlets 3—3.5 mm diam., loneitudiaally ridged, but not sul- cate; leaf blades coriaceous, densely and prominently pellucid punc- tate above and below; leaf buds densely appressed rufous glandular- stellate and -dendroid tomentose; petioles short-marginate, 3—5 mm ong, not canaliculate, the midrib raised to petiole base; peduncle thin, 1-2 mm diam.,; pedicels 4-6 mm long; sepals widely ovate, 14. M. glandulosa i 0.6-0.9 mm long. 1. Myrsine medeciloae Pipoly, nom. nov. (Fig. 1A, B). Rapanea ee Merrill, Philipp. - Sci. 20:429. 1922, non Myrsine a (A. DC.) D. Dietr., Synops. 1:619. 1839 [= an aoa DC. }, nec Myrsine angustifolia E. Meyer ex Schimper, Traité Pal. 2:922. 1872 ak cea ), nec Myrsine cl Heller, Minn. Bot. Stud. a . pes 1897, nec Myrsine ps (Mez) Hosaka, Occ. Pap. Bish. Mus. 16:42. 1940= hae angustifolia Mez, ie 9 (IV. 236):337. 1902.) Tyre: PHILIPPINES. Luzon Istanp. Ilocos Norte Prov.: Mt. Nagappatan, Aug 1918 (fr), M. Ramos BS 33243 (HOLOTYPE: PNH-destroyed; Lec- roTyPe, here designated: A!). Shrub or small tree, height unknown. Branchlets: essentially terete, |.5— 2mm diam., densely rufous-puberulent and glandular-granulose apically glabrescent. Leaves: buds densely rufous-puberulent and glandular-granulose, rufous glandular-ciliate along margins; blades chartaceous, symmetric, linear to narrowly linear-oblanceolate, (2.5—)5—6.5 cm long, 0.6—1.0(-1.5) cm wide, obtuse apically, retuse at apex tip, acute basally, decurrent to petiole base, smooth and nitid above, pallid below, densely and prominently black punctate below toward apex, the midrib impressed above, prominently raised below, secondary veins 12-15, slightly prominulous above and below, the marginal collecting vein ca. 0.5 mm from margin, entire and revolute; petioles marginate, not canaliculate, 4-9 mm long, rufous puberulent at first, glabrescent. Staminate inflorescence: sessile, 6-12- flowered; floral bracts coriaceous, lanceolate, | mm long, 0.3 mm wide, apically acute, densely and prominently black punctate medially, the margin entire, densely glan- dular-ciliolate; pedicels cylindric, 1.5—3 mm long. Staminate flowers: 4- merous, chartaceous, 2.5—3 mm long; calyx cotyliform,0.7—0.9 mm long, Prpoty, Synopsis of Myrsine in the Philippines Were ESN be SLvOrs Fic. 1. A-B. Myrsine medeciloae ee - piscillate branch, note sessile umbels (lectotype). B. Staminate branch (R. Meyer BS 311 C. Myrsine oblongibacca (Merrill) Pipoly, showing aay tee fruit (lectotype). D. ae eu (Elmer) fev w ipoly, pistillate branch, showing - d inflorescences and elliptic leaves (1 ype). E-G. Myrsine amorosoana Pipoly. E. Piscillate branch, showing revolute leaf margins and eben. fruits (G. Edaio 76198). F. Staminate branch, showing retuse leaf apices (G. Argent G E. Reynoso 89112). G. Pistillate ae, show! ewer-fruited inflorescences and much larger leaves (G. Edatio BS 76198). ee oruciata (Philipson) Pipoly. H. Staminate branch (J. & M. Clemens 30258). 1. eis branch, showing ovoid fruits (C. Ridsdale SMHI 225). Black scale bar equals 1 cm, smallest unit on white scales equals | mm 124 Sipa 17(1) the tube 0.1—0.2 mm long, the lobes deltate to ovate, 0.5—0.8 mm long, 0.5—0.7 mm wide, acute apically, prominently black punctate medially, along the margins opaque, irregular and apically somewhat erose, sparsely glandular-ciliolate; corolla rotate, 1.8—2.2 mm long, the tube ca. 0.5 mm long, the lobes oblong, 1.3—1.7 mm long, 0.5 mm wide, rounded apically, densely and prominently black punctate without except at margin, densely glandular-granulose on the margin and apically within, entire along mar- gin; stamens 1.4—1.6 mm long, the filaments developmentally fused to corolla tube, the anthers apparently epipetalous, sessile, widely ovate, 1— 1.2 mm long, 1—1.3 mm wide, apically obtuse, basally subcordate, densely and prominently black punctate abaxially; pistillode hollow, conic, ca. 0.5 mm long, the stigmatic area somewhat lobed. Pisti/late inflorescence: sessile, 3—6-flowered; floral bracts coriaceous, ovate, ca. | mm long, 0.8—1 mm wide, acute apically, densely and prominently black punctate medially, entire and densely glandular-ciliolate along margin; peduncle obsolete; pedicels obconic, (1.5—)2—2.5(—3) mm long. Pisti/late flowers: like stami- nate but 2.3-3.0 mm long; calyx cotyliform, 0.7—1 mm long, the tube ca. 0.2 mm long, lobes deltate, 0.5—0.8 mm long and wide, apically acute to subacuminate; corolla subrotate, 1.6—-1.9 mm long, the tube ca. 0.5 mm long, the lobes oblong, 1.1-1.4 mm long, 0.5 mm wide; staminodes re- sembling stamens but filaments obsolete, the antherodes widely ovate, l— 1.2mm long, |—1.3 mm wide, apically obtuse, basally subcordate, densely srominently black punctate abaxially, devoid of pollen; pistil conic, 1.3—1.5 mm long, the ovary 0.7—0.9 mm long, 1-1.2 mm diam, the stigma 3—5 mm long, 4-lobed, the lobes ligulate with variously lacerate margins; ovules 3—4, completely buried in the placenta. Frwit globose, 3-4 mm long and diam., densely and prominently black punctate with minute punctations in the upper 1/3 of fruit, the persistent stigma 3—4-lobed. Distribution.—Endemic to western Luzon Island, in Ilocos, Bataan and Zambales provinces, Philippines, at 1,000—1,800 m elevation. Ecology and conservation status. —Myrsine medeciloae is a rare species, occur- ring only in submontane mossy forests in the northern portion of the coun- try. Because of habitat destruction, this species is considered endangered and most populations may be extinct. Etymology.—It gives me great pleasure to dedicate this species to Melanie Medecilo, Philippine Adminstrator for the Philippine Plant Inventory Project. Ms. Medecilo is a budding systematic botanist with strong orga- nizational skills, infinite patience, and much creativity. “Aribangib” (Negrito language, Madulid 1992), and — — Common name. Specimens examined: PHILIPPINES. Luzon IstaAnb. Baatan Prov.: Mt. Mariveles, Lamao River, May 1905 (stam. fl), R. Meyer BS 3114 (NY, US). Ilocos Norte Prov.: Burgos, Jul 1918 (fr), Ml. Ramos BS 32757 (A); en route from miner's camp to Mt. Burnay Pipoty, Synopsis of Myrsine in the Philippines 12) in Mt. Sicapoa Range, 1,700-1,800 m, 3 Dec 1975 (pist. fl, fr), K. lwatswki et al. 719 (L- 2 sheets.). Zambales Prov.: Mt. Lapulao, Nov—Dec 1907 (fr), H. M. Curran & M. L. Merritt BS 8072 (US). Myrsine medeciloae is closely related to M. apoensis, but can esily be distin- guished by its linear or narrowly oblanceolate leaves, short, cylindric pedicels, and young branchlets and petioles rufous puberulent and glandu- lar-granulose. 2. Myrsine apoensis (Elmer) Pipoly, comb. nov. (Fig. 1D). Rapanea apoensis Elmer, Leafl. Philipp. Bot. 2:669. 1910. Type: PHILIPPINES. Minpanao Isianp. North Cotabato Prov.: Mt. Apo, Todaya, May 1909 (pist. fl, fr), A. E/mer 10629 (HOLOTYPE: PNH- See LECTOTYPE, here designated: A!; lsoLecrorypss: BISH}, GH-2 sheets!, K!, L!, Tree to 4 m tall. Branchlets: terete, 1.5—2.5 mm diam., densely rufous glandular-papillose, glabrescent. Leaves: buds densely rufous glandular-pap- illose, margins densely translucent-ciliate along margins, glabrescent; blades membranaceous to chartaceous, symmetric, elliptic to lanceolate, 1.7—6.5 cm long, 0.8-1.8 cm wide, apically acuminate to an obtuse tip, basally acute, narrowly decurrent about 3/4 along the petiole toward base, smooth, subnitid and inconspicuously pellucid punctate above, inconspicuously pellucid punctate below, midrib immersed above, prominently raised be- low, the secondary veins 9-13 pairs, barely prominulous above and below, the submarginal collecting vein ca. 0.25 mm from margin, entire and revo- lute along margin; petioles deeply canaliculate, not marginate, 4-9 mm long, densely rufous glandular-papillose above, glabrescent. Staminate inflorescence: unknown. Pistillate inflorescence: a subsessile to sessile umbel, 3— 6-flowered; peduncle rarely to 1 mm long, ca. 0.5 mm diam.; floral bracts chartaceous, oblong, 1.8—2.2 mm long, 1.3—1.5 mm wide, apically broadly rounded, prominently black punctate-lineate medially, slightly erose and sparsely glandular-ciliate along the margin; pedicels cylindric, 3-6 mm long. Pzstillate flowers: S-merous, chartaceous, 2—2.5 mm long; calyx cotyliform, 0.8—1 mm long, the tube ca. 0.2 mm long, the lobes subor- bicular, 0.6—0.8 mm long and wide, apically acute, medially carinate at first, chen flattening with age, densely red punctate and punctate-lineate, with a hyaline, irrregular, entire, rufous glandular-ciliolate margin; corolla rotate, 2—2.5 mm long, the tube ca. 0.2—0.4 mm long, the lobes lanceolate, 1.8—2.1 mm long, 0.2—0.4 mm wide, apically acute, densely and promi- nently red punctate-lineate medially, densely glandular-granulose toward apex within and along the maar eta, with margins densely glandular- granulose, entire; staminodes |.2—1.4 mm long, the filaments 0.2—0.4 mm long, completely fused to corolla ae the antherodes ovate, 0.9—1.1 mm long, 0.3—0.4 mm wide, apically rounded, basally cordate, devoid of pol- len, the connective darkened dorsally; pistil obnapiform, the ovary 1.5 mm 126 Sipa 17(1) long and diam., the style short, ca. 0.5 mm long, truncate terminally, the style ligulate, alate, the wings 4; ovules 3-5, partially immersed in the placenta. Frit subglobose, slightly longer chan wide, 2.6—3.5 mm long, 2.3—3 mm wide, prominently red punctate-lineate. Distribution.—Endemic to Mindanao Island, on Mts. Apo and McKinley, North Cotobato, Davao and Davao del Sur provinces, at 1,666—2,900 m elevation. Ecology and conservation status.—Myrsine apoensis is restricted to the mossy forests ocurring on the upper slopes and summits of the mountains it is known from. These forests are not commercially exploited, but are subject to de- struction due to activities related to construction and maintenance of tele- communication towers. Therefore, the species should be considered highly endangered. Etymology.—The specific epithet refers to the type locality, Mt. Apo, lo- cated at the junction of North Cotabato, Davao and Davao del Sur provinces. Common names,— Karyos” (Bagobo language, Merrill 1926); “Marintok” (Bagobo, Madulid 1992), “Tongog” (Bagobo language, Bisayan, Panay dia- lect, Madulid 1992). Specimens examined. PHILIPPINES. MiInpANAO IsLAND. Davao del Sur Prov.: Mc. ele N 7 Lake Linao, 2,100 m, 30 Oe 1946 (fr), G. Edaiio 1266 (PNH , | Nov 1946 (fr), G. Edaiio 1267 {PNH 1383] (A Seite 20 May 1974 (fr), Univ. hee cin 586 (L); ae McKinley, summit, 2,466 m, 13 Sep 1946 (fr), G. Edaito on {PNH 1048} (A, L, PNH);1,666 m 28 Aug 1946 (ster.), 2 Edatio 821 {PNH 1032] ( PNH). pea Coca eka Kidapawan Municipality, Mt. Apo Geothermal ae Site G, 06°59.5'N, = 14'E, 1,820 m, 8 Nov 1992 (fr), L. Co 3649 (A, CANB, CAHUP, K, L, PNH, PUH, TI, US _ to Myrsine apoensis appears to be closely related to M. medeciloae, but is readily distinguished by the elliptic to oblanceolate leaf blades, canaliculate peti- oles, the longer, cylindric pedicels and the rufous glandular-papillose (not puberulent or granulose) young branchlets and petioles. 3. Myrsine amorosoana een nom. Te LE-G). Rapanea retusa Merrill, Philipp. J. Sci. 2:297. 1907. Typr: PINES. Minporo Istanp. Oriental Mindoro Prov.: Mc. an con, oe 1906 i 4 Menai 3 5734 (HOLOTYPE: PNH-c stroyed; LECTOTYPE, here designated: US!; isoLecrorypes: K!, NY!), von Myrsine retusa Aiton, Hort. Kew 1:271. 1789. nee Myrsine africana var. retusa (Aiton) A. DC., Trans. Linn. Soc., Bot. 17:105. 1834. Shrub or small tree to 3(—5) m tall. Branchlets: terete, 2.5—3 mm diam.glabrous. Leaves: buds glabrous with margins sparingly rufous glan- > dular-ciliate; blades thinly coriaceous, symmetric, oblong or elliptic, (2.3—) —5(—6) cm long, (0.8—)1.5—1.9 cm wide, apically retuse to emarginate at tip, basally cuneate, smooth and nitid to laccate above, densely and promi- nently black punctate below, midrib prominently raised above and below, Pipoty, Synopsis of Myrsine in the Philippines 127 the secondary veins 6-10 pairs, not visible above, more or less prominulous below, the submarginal connecting vein ca. | mm from margin, entire and revolute along the margin; petiole marginate, 4-9(-12) mm long, with a decurrent midrib prominently raised adaxially to base, glabrous. Staminate inflorescence: subsessile, 3—9-flowered; peduncle 1.5—2 mm long, 0.5—1 mm diam.,; floral bracts coriaceous, deltate, ca. 0.5 mm long and wide, apically acute, inconspicuously punctate, irregular to erose and glandular ciliolate along the margin; pedicels cylindrical, 2-3 mm long. Staminate flowers: 4- merous, coriaceous, 1.5—1.8 mm long; calyx cupuliform, 1-1.2 mm long, the tube 0.3—0.5 mm long, the lobes oblate, 0.5—0.7 mm long, 0.6-0.8 mm wide, apically obtuse to subacute, medially prominently red punctate, irregular, erose, and sparingly glandular ciliolate along the margin; corolla rotate, 1.6—1.8 mm long, white, the tube ca. 0.2 mm long, the lobes ellip- tic, 1.4-1.6 mm long, 0.7—1 mm wide, apically obtuse, densely and promi- nently red punctate and punctate-lineate toward apex and along margins without, densely glandulat-granulose outside and along margin within; stamens 1—1.2 mm long, the filaments fused to the corolla tube 0.2 mm, the anthers widely ovate, 0.8—1 mm long, 0.6—0.8 mm wide, apically acute, basally slightly cordate, opening by large subapical introrse pores, then large longitudinal slits, the connective inconspicuously brown-punctate dorsally at apex; pistillode ellipsoid, 0.7 mm long, the ovary 0.3-0.5 mm long, 0.2—0.3 mm diam., the stigma translucent, amorphous, 0.2 mm long. Pistillate flowers: like staminate but 1.6-1.9 mm long; calyx 1-1.2 mm long, the tube ca. 0.1 mm long, the lobes 0.9-1.1 mm long, 0.6—0.8 mm wide; corolla 1.6—1.9 mm long, the tube 0.2—0.3 mm long, the lobes 1.4— 1.6 mm long, 0.2—0.3 mm wide; staminodes 0.8—1 mm long, the filament completely fused to the corolla tube in development, 0.3 mm long, the antherodes sublinear to ovate, 0.5—O.7 mm long, 0.1—0.2 mm wide, apically acute, the base not distinguishable; pistil globose, 1.3—1.5 mm long, the placenta globose, the ovules 4—G, in 2 series, the stigma 4-lobed, arrow- head-like, the lobes fin-shaped, ca. 0.5 mm long, 0.7 mm wide. Fruit glo- bose, 2—3 mm long and in diam., violet at maturity, densely and promi- nently red punctate. Distribution.—Endemic to the Philippines, from north central Luzon Is- land southward through Mindoro, Sibuyan and Negros Islands to northern Mindanao, at 1,400—2,825 m elevation. The collection from Bicol Na- tional Park (B. Hernaez & M. Cajano CAHUP 572, 57283, 57284) was undoubtedly from near the 1,900 m elevation and not the lower extreme indicated on the label. Ecology and conservation status.—Frequent in mossy forest, rag Ramayan” forest, dominated by gymnosperms and Tristaniopsis (Myrtaceae) and upper 128 Stipa 17(1) montane moist forest, particularly on ultramafic substrates. Myrsine amorosoana forms large patches of individuals along ridgetops in forests dominated by Phy/locladus hypophyllus (Podocarpaceae), with an understory dominated by Polyosma verticillata (Escallionaceae). It occurs with a density of approximately 50 individuals per hectare, a figure perhaps exceeded only by Myrsine dependens (Ruiz & Pavon) Sprengel f., of the northern Andes of South America, and M. pitrieri (Mez) Lundell of Costa Rica and Panama. It may be that some of the species currently under Rapanea and undergoing revision for the entire island of New Guinea, may have even more extensive populations. Myrsne amorosoana, by virtue of its large, but extremely local- ized populations, should be considered threatened. Etymology.—This species is dedicated to my friend and colleague, Dr. Victor Amoroso, professor of biology and curator of the herbarium of Cen- tral Mindanao University, in Musuan, Bukidnon Province, Mindanao. Dr. Amoroso is an authority on the systematics of Philippine pteridophytes, a prodigious fieldworker and an enthusiastically supportive colleague. His indefatigable efforts in conservation and monitoring of the biodiversity of Mindanao’s forests are exemplary. Common names. —"Biribenlog” (Bikol language, Ramos & Edaito BS 26577), “Maromo’ (Negrito, Madulid 1992). Sper imens pes PHILIPPINES. Luzon IsLanb. s/ve Joc. esp. 1792 (fr), TE. Haenke 440, 447, 472, (NY). Benguet Prov.: Kabayan, Mt. Pulog National Park, 10 Aug 1992 (fr), E. Barbon et ale PPI 8845 (A, BO, BRIT, K, L, PNH, US); Mc. Pulog, ae a 1925 (fr), M. Ramos & G. Edafto BS 44936 (US). Camarines Norte Prov.: Dec 1913 (fr), AM. Ramos BS 1524 (A, L, NY), (stam. fl), M. Ramos BS 1525 (A, L, NY). Camarines Sur v.: Mc. Isarog, “Dec 1928"{probably Nov] (fr), G. Edaitto BS 76198 (A, K, NY, PNH),1,370 m, 28 Nov 1928 [”Dec 1928"on lower label} G. Edafto 630 {BS 76225} (A, NY); BS 76268 pe Mt. Isarog, Naga City, 6 Apr 1992 (fr), Barbon et al. PPI st 23 (A, BO, BRIT, K, L 1); Mt. Isarog, Nov—Dec 1913 (fr), M. Ramos 22009 ee ‘ee Bicol National Park, ae ampo del Rosario, S access trail W of PLDT, 670-1 ae m, 22—27 Oct 1991 (fr), B. Hernaez & M. C chan CAHUP 57282, 57283, 57284 (C P). Mountain Prov.: Mc. Pulog, 16°36'N, 120°S4'E, 2,550—2,650 m, 30 Jan oe ms 7 Jacobs 7227 (A, K, L, L i PNE): 2,60 nen Om, | me 1968 (fr), M. ee 7274 (A, K, L, LBC PNH); 2,300—2,500 m, 5 Feb 1968 (stam. fl), M. Jacobs 7333 (A, K, L, PNH), 2,300— 2,400 m, 17 rb ne hoe M. eae 7488 (L, PNH); Lake Palas Aug 1915 (pist. fl), M. Ramos BS 23616 (A, K, PNH, US), 24—25 Oct 1965 (fr), R. Robbins s.n. (L). Nueva Vizcaya rov.: Mt. i apan, May—Jun ee (fr), M. Ramos & G. Edatto BS 45602 (NY). Quezon {Tabayas} Prov.: Mt. Binuang, May 1917 (fr), M. Ramos G G. Edaito BS 28758 (A, K, PNH, US); Mc. Dingalan, Aug—Sep 1916 (stam. fl), M. Ramos & G. Edaito BS 26577 (A, US); Mc. Mirador, summit, Antimonan Municipality, 13 Sep 1975 (fr), B. Hernaez as (CAHUP), Quezon National Forest Park, Atimonan summit, 400 m, 26 Aug 1984 (fr), Lit IL84—001 {CAHUP 40296) (CAHUP). Sorsogon Prov.: Jul-Aug 1915 (stam. fl), : Ramos BS 23512 (BISH, K, PNH, US). aes a AND. Prov. Oriental Mindoro: Mt. Halcon, near gy Dulangan River, 13°12 ay. °12'E, 1,200 m (fr), C. Ridsdale et al. 1705 (A, K, L, LBC, PNH); NE summit, 2 eo 80 m, i May 1995 (pist. A), E. aan Pipoty, Synopsis of Myrsine in the Philippines 129 Mandia 421 (CAHUP, DLSU); N coast, Mindoro, Subaan River headwaters, inland from San Teodoro, 1,200 m, 30 Apr 1986 (fr), M. Coode et al. 5635 (A, K, L, LBC, PNH); Mt. Halcon, 15—24 Jun 1906 (fl), M. Merritt BS 4449 (US), Nov 1906 (fr), E. Merrill 5735 (L); NE Mindoro, Mustning, Mt. Halcon Complex, above Paitan on Dulangan River, 1,450- 1,500 m, 11 May 1986 (fr), M. Coode et al. 5765 (A, K, L, PNH). Rizal, Augilog Mar 1906 (fr), A. Loher 6128 (K); sine loc. esp., Nov 1914 (fr), M. Ramos BS 1961 (L, NY). StiBUYAN IsLAND. Prov. Romblon: Mt. goa -guiting, Camp 3 above Magdiwang on ridge lead- o Mayos’ Peak, 1,400 m, 27 Aug 1989 (stam. fl), G. Argent G E. Reynoso 89112 i none (E, K, PNH); Magdiwang, Mt. Guiting-guiting, Mayos Peak, 12°28 IN, 122°32'E, 5 Jun 1992 (fr), B. Stone et al. PPI 6906 (A, BO, BRIT, K, L, PNH, US. NgGros IstaND. Negros Occidental Prov.: Mt. Canlaon ee 1950 m, 10 Apr 1954 (fr), G. Edatio PNH aie 8195 {PNH 22003} (A, K, L, PNH). Minpanao IsLtanp. Bukidnon Vv. ndoon, Jun—Jul 1920 (fr), M. Ramos . G. Edafio BS 38918 (A); Mt. Lipa, ai “jul ion (fr), M. Ramos & G. Edaiio BS 38528 (A, L); Lantapan Municipality, Sitio Sungco, Mt. Kinasalapi, part of Kitanglad Mt. Range, W of Alanib River, SW slopes 08°00'05"N, 124°30! 56" E; 2,090-2 360 m, 4 Dec 1984 (ster), J. Pipoly et al. PPI 16610, (BRIT, PNH) (fr) Pipoly et al. PPI] 16647.(BRIT, PNH) (fr), J. Pipoly et al. PPI 16684 (BRIT, PNH), 6 Dec 1994 (fr), Pipoly et al. PPI 16871 2,200-2,320 m, 14 Apr 1995 (fr), J. Pipoly et al. 19551 {PPI 19796} (BRIT, PNH), (fr), 19555 {PPI 19800} (BRIT, PNH), ar Recnecemer Mt. Kitanglad Range, 20 km SE of Dalwangan, Malaybalay ity, 87°5'N, 124°56 E, 2,385—2,825 m, 25-30 May 1993 (fr), B. Hernaez G M. ae oo {CAHUP 60561, 60562, 60563, 60564} (CAHUP). Myrsine amorosoana is most closely related to M. oblongibacca, but 1s sepa- rated by the symmetric, oblong to elliptic leaf blades with obtuse to rounded apices and revolute margins, and the globose fruits. 4. Myrsine oblongibacca (Merrill) Pipoly, comb. nov. (Fig. 1C). Rapanea ea oe Merrill, Philipp. : Sci. 20:429. 1922. Type: PHILIPPINES. Luzon Is- llo te Prov.: Mt. Palimlim, Aug 1918 (fr), M. Ramos BS 33256 (HOLO- TYPE: -: PNH. destroyed; Lectotype, here designated: A!; ISOLECTOTYPE: K!). Glabrous shrub to 3 m tall. Branchlets: somewhat angulate at very tip, soon terete, 2—3 mm diam. Leaves: buds glabrous with margins sparsely glandular ciliolate; blade chartaceous to subcoriaceous, asymmetric, nar- rowly oblong to narrowly oblanceolate, 3—6(—7.4) cm long, 0.8-1.8 cm wide, apically subacuminate to obtuse, but blunt at tip, basally cuneate, decurrent on the petiole, smooth, prominently pellucid punctate and nitid above, prominently pellucid punctate and black punctate-lineate below, midrib prominently raised above and below, the secondary veins 7—12 pairs, not visible above, inconspicuous below, the submarginal collecting vein ca. 0.5 mm from margin, densely and prominently black punctate below, en- tire, and flat along margin; petioles canaliculate and marginate, 3-8 mm long, glabrous. Staminate inflorescence: unknown. Pistillate inflorescence: subsessile, or on an apparently annual, peduncle 1.5—2 mm long, up to | mm diam.; floral bracts chartaceous, linear, 0.5—0.7 mm long, 0.1—0.2 mm wide, apically acute, densely black punctate, entire and glabrous along the margin; pedicels cylindric, 2-3 mm long, sparsely black punctate-lineate. 130 Sipa 17(1) Pistillate flowers: unknown, fruiting calyx 4—5-merous, cotyliform, 1—1.2 mm long, the tube ca. 0.2 mm long, the lobes irregular, ovate, 0.8—1 mm long, 0.5—0.7 mm wide, apically acute to acuminate, densely and promi- nently black punctate medially, densely glandular-ciliate along the mar- gin. Fruit oblongoid, 5-8 mm long, 3—5 mm diam., the persistent stigma scar stylopodic, densely and prominently pellucid | aerate lineata. Ditrshadioae desire oblongibacca is endemic to the Philippines, occur- ring on the Island of Luzon, in Cagayan, Ilocos Norte, Nueva Ecija, Quezon, and Rizal provinces, and on the Island of Dinagat, just north of Mindanao Island. While no precise elevation has been listed on collections, one might surmise that it is a submontane species, and as such, would be expected at perhaps 800—1,500 m elevation. Ecology and conservation status. —Virtually nothing is known of the ecol- ogy of M. oblongibacca. It is known only from historical collections, and recent expeditions made to the areas from which it was previously known have not resulted in recollections. However, Mt. Palimlim (Ilocos Norte), and Dinagat Island are areas containing significant pockets of vegetation on ultramafic soils. Therefore, it is possible that this species is another ultramafic specialist. Given the scanty remnants of forests left in the coun- try, it 1s doubtful that we will ever know the natural distribution of M. oblongibacca. \t is assumed that the species is highly endangered if not al- ss, ready extinct. Etymology.—The specific epithet refers to the ob among species of the genus. Common names.—“Supak” ([gorot language, Madulid 1992). — ongoid fruit, a rarity Specimens examined. PHILIPPINES. Luzon Istanp. Cagayan Prov.: Mt. Cagua Vol- cano, Oct-Nov 1929 (fr), G. Edaiio BS 78357 (K, NY, PNB), (fr), G. Edaito - la ) (NY), (ster.), i Edano BS 78424 UK, NY, PNH), Mc. Dos Cuernos, Apr 1929 (ft "), A Ramos BS 77034 (NY); Bauan District, Me. Tabuan, Mz ay 1929 (fr), M. Ramos ce ee (NY): Me. sere Nov 1929 (fr), G. Edafio BS 78533 (NY). Hocos Norte Prov.: Mt. Palimlim, Aug 1918 (fr), M. Ramos BS 33311 (A, K, PNH, US). Nueva ie Prov.: Mc. Umingan, hos Sep 1916 (fr), M. Ramos G G. Edatio 26418 (A, K, PN US). Quezon {Tabayas} Prov.: Mt. Dingalan, Aug—Sep 1916 (fr), . Ranos & G. aes BS 26548 (A, US). ore Prov.: Mt. Susong-Dalaga, Aug 1917 (fr), M. Ramos & G. Edaiio BS 29386 (A, K, PNH). DinaGar IsLanb. Surigao del Norte ee an 1919 (A bud), M. Ramos GJ. Pasgasio BS 35194 (L). Myrsine oblongibacca is most closely related to M. amorosoana, from which it is easily distinguished by its oblongoid fruits, and asymmetric, narrowly oblong or oblanceolate leaves with flat margins. 5. Myrsine cruciata (Philipson) Pipoly, comb. nov. a 1H, 1). Rapanea cruciata Philipson, J. Bot. (London) 77:105. 1939. Tyr: LAYSIA. Saban {Brit- is Nort Borneo}: Pava Cave, Mc. Kinabalu, Upper ae 2,400—3,050 m, 29 Mar 1932 (stam. fl), J. GM. . Clemens 28941 (HOLOTYPE: BM, n.v.; isorypes*: Al, Pipoty, Synopsis of Myrsine in the Philippines 1gt K!, L!, NY!). *The isotype at Kew contains two distinct elements, one-half of the sheet bearing a twig of M. cruciata and the other an entity close to, if not conspecific with, M. dasyphylla Stapf, and both sides were correctly annotated by Philipson. However, the sheet at A bears two twigs of still another unknown Myrszne species entity with very prominent secondary and tertiary leaf venation. Thus no portion of ie & M. S. Clemens 28941 housed at A corresponds to R. cruciata. On the sheet at NY, ortion corresponds to M. cruciata, while the right hand portion to M. iar Stapf. The L sheet is entirely M. cruciata. Shrub or tree to 5 m tall. Branchlets: angulate, 2.5—3.5 mm diam., some- what lenticellate, glabrous. Leaves: buds with margins sparingly glandular- ciliolate, glabrescent; blade cartilaginous, symmetric, oblanceolate to ob- long, (3-)6-9.5 cm long, 1.8-2.5 cm wide, apically rounded to obtuse, retuse to emarginate at very tip, basally cuneate, smooth and laccate above, pallid and conspicuously black-lineate and prominently black punctate along margin below, the midrib impressed above, prominently raised below, the secondary veins 16—22 pairs, scarcely or not visible above, scarcely visible below, the submarginal collecting vein ca. 0.5 mm from margin, entire and revolute along the margin; petiole marginate, 3—5 mm long, the mid- rib elevated medially adaxially, glabrous. Staminate inflorescence: subsessile, 2—5-flowered; peduncle ca. 2-3 mm long, ca. 1 mm diam.; floral bracts coriaceous, deltate, 1.5—2 mm long, |—1.3 mm wide, apex acute or obtuse, the margin entire, glandular cioliolate; pedicels angulate, 2-3 mm long. Staminate flowers: 4—S-merous, chartaceous 3—3.5 mm long; is xin, 2.3—2.8 mm long, the tube 0.3-0.4 mm long, the lobes ovate, 2—2.4 m long, 1.2-1.5 mm wide, apically obtuse to acute, medially thickened, con- spicuously (not prominently) black punctate, entire and densely glandu- lar-ciliate along the margin; corolla rotate, 3.3—3.5 mm long, the tube 0.8—1.1 mm long, the lobes elliptic 2.3-2.5 mm long, 1-1.2 mm wide, apically obtuse, densely and prominently black-lineate medially, black punctate marginally, dene granulose along the margin and near the margin within; stamens 3—3.2 mm long, the filaments 0.8—1.1 mm long, connate and eee adnate to the corolla tube, the anthers ob- long, 1.8—2 mm long, 1—-1.2 mm wide, apically apiculate, basally cordulate, dehiscent by wide longitudinal slits, the connective eglandular; pistillode conic, ca. 0.3 mm long and diam., hollow. Pzsti//late inflorescence: like stam1- nate but peduncle accrescent annually to 3 mm long, ca. 1 mm diam.,; pedicels 2—2.5 mm long; floral bracts 1.5—2 mm long, 1-1.3 mm wide. Pistillate flowers: like staminate but 2.8—-3 mm long, the tube 0.8—1.1 mm long, the lobes 1.9-2.2 mm long, 1—1.2 mm wide; staminodes 2.3—-2.8 mm long, the filaments 0.6-1.1 mm long, the antherodes ovate, 2—2.2 mm long, 0.4—0.6 mm wide, apically acute, basally sagittate; pistil ellip- soid to subglobose, 2—2.2 mm long, 1.3—1.5 mm wide, the stigma ligu- late, tortuous, ca. 1 mm long, stylopodic basally, the placenta subglobose, 132 Sipa 17(1) the ovules 6—8, biseriate. Fruit ovoid to oblongoid, (3—)5—7 mm long, 3—5 mm diam., densely and prominently black ees and punctate-lineate. Bi lane —TIn the Philippines, My ‘yuciata is known from the hills of Mt. Beaufort, above Puerto Pence on the island of Palawan, and from unknown localities in Abra and Zambales provinces. Outside of the Phil- ippines, it is known otherwise only from Mt. Kinabalu in Sabah, Malaysia. The altitudinal range of the species is 850 to 2,500 m elevation. Ecology and conservation status. —Myrsine cruciata appears to be restricted to sandy soils on quartzite rocks. Regarding its conservation status, no recent collections have been made from Luzon, and those populations may be extinct. However, the populations in Palwan appear to be surviving so far. Given these circumstances, Myrsine cruciata should be considered a threat- ened species. Etymology.—The specific epithet refers to the tetramerous perianth parts on the type specimen. Common name.—Tongog” (Bagobo language). — Specimens examined. PHILIPPINES. Luzon Istanp. Abra Prov.: sine loc. esp. Jan Feb 1909 (fr), M. Ramos BS 7263 (NY, US). Zambales Prov.: sine loc. esp., Dec 1907 (fr), M. Ramos BS 5050 (NY, US). PALAWAN IsLAND. Municipality Puerto Pincesa, Mt. Beau- fort, W spur, 815 m, 28 Mar 1984 (fr), C. Ridsdale SMHI 225 (A, K, L, PNH); Narra, Victoria Peaks, Trident Mining Co. Concession, 490-590 m, 18 May 1984 (fr), C. Ridsdale SMHI 1732A,C. err SMHI 1741 (A, K, L, PNH), 620 m, 20 May 1984 (fr), A. Podzorski SMHI 2125 (A, CAHUP, K. L, PNH), A. ee ie 2130 (A, K, L, PNH), 500 m, J. Dr mre 1292 (A, K,L, PNH). MALAY A. SABAH: Mc. Kinabalu, Pig Hill, 7,000 ft., 18 Feb 1964 (fr), W. Chew & E. Corner ate 4372 (K), 2,300 m, 24 Feb 1964 (fr), W. Chew & E. Corner RSNB 4493 (K), Upper Kinabalu, 1,830—4,115 m, 2 Jun 1932 (stam. fl), J. & M. Clemens 30258 (K); Mari Pari, 1,524 m, 27 Mar 1933 (fr), J. & M. Clemens 32378 (K),. While the population of Myrsine cruciata (sensu stricto) from the type lo- cality has mostly oblong leaves instead of oblanceolate ones, and the tex- ture is somewhat thicker, I have no doubt that the Philippine and Bornean populations are conspecific. This species has often been confused with Myrsine oblongibacca, but may easily be separated from it by the wider, symmetric leaf blades, angulate branchlets, and larger, mostly ovoid fruits. In most of its range through the Philippines, Myrsine cruciata is most readily com- pared to M. densiflora, but is immediately distinguished by the cartilagi- nous and laccate leaf blades with retuse to emarginate tips, and the larger fruit. Myrséne cruciata inhabits sandy soils on quartzite rocks, apparently adjacent to ultramafic substrates in submontane and montane habitats, while M. densiflora occurs on ultramafic soils behind coastal beaches. 6. — densiflora Scheffer, Comm. Myrs. Archip. Ind. 50. 1876. (Fig. 2 ). Rapanea densiflora (Scheffer) Mez in Engler, Pflanzenr. 9 (IV. 236):365. 1902. Type: INDONESIA. Irian Java [West New Guineal: S coast, along Daurga Pipoty, Synopsis of Myrsine in the Philippines os) River, Princess Marianne Strait, May 1828 (stam. fl), A. Zzppelzus 5.n. (HOLOTYPE: L, n.v., fragment, A!). Shrub or tree to 3—5(—10) m tall. Branchlets: angulate, (3—)3.5—5 mm diam., longitudinally ridged, lenticellate with age, glabrous. Leaves: buds with margins densely glandular-ciliolate, glabrescent; blades coriaceous, symmetric, elliptic to obovate, rarely oblanceolate, (5.5—)6—-14(-16) cm long, 3—5(—6) cm wide, apically broadly rounded to acute, basally cuneate, slightly decurrent on petiole, dull or somewhat nitid above, scrobiculate above and below, densely and conspicuously black punctate and punctate- lineate below, the midrib impressed above, prominently raised below, the secondary veins 13—17 pairs, prominulous above and below, connected by an inconspicuous submarginal collecting vein less than 0.5 mm from the margin, inrolled, densely and prominently black punctate below along the margin. Staminate inflorescence: fasciculate, 3—9-flowered; peduncle 2—3.5 mm long, 1-2 mm diam. (the longer peduncles apparently all on one year's growth); floral bracts coriaceous, deltate, 0.8—1.0 mm long and wide, apically rounded or obtuse, densely and prominently black punctate apically, the margin entire, glabrous; pedicels cylindric, 1.5—2 mm long, prominently reflexed at anthesis, conspicuously black lineate. Staminate flowers: 4-merous, chartaceous, 1.8—2.3 mm long; calyx cotyliform, 0.8—1.0 mm long, the tube ca. 0.2 mm long, the lobes widely ovate, 0.6-0.8 mm long, 0.7—0.9 mm wide, apically acute, medially prominently black punctate and punc- tate-lineate, the margin opaque, irregular, glabrous; corolla subrotate, 1.7— 2.2 mm long, the tube ca. 0.1 mm, the lobes oblong or elliptic, 1.7—2.2 mm long, 0.7—1.0 mm wide, apically obtuse to rounded, medially densely and prominently black punctate throughout, punctate-lineate apically and medially, entire, and densely glandular-granulose along the margin, gla- brous within; stamens 1.3—1.5 mm long, the filaments flat, adnate to adaxial corolla tube and lobe, inconspicuous, 0.2—0.3 mm long, the anther ovate to obcordate, 1.1—1.3 mm long, 0.5—0.7 mm wide, apically acute, basally broadly cordate, the connective densely and prominently brown punctate dorsally; pistillode conic, 0.2-0.3 mm long, 0.3—0.4 mm diam., hollow, without style or stigma. Pisti/late inflorescence: like staminate but peduncle to 3 mm long, 2 mm diam.; pedicels 1.5—2.5 mm long. Pistillate flowers: like staminate but 1.3—1.5 mm long; calyx 0.6—-1 mm long, the tube ca. 0.2 mm long, the lobes 0.5—0.7 mm long, 0.6—0.8 mm wide; corolla 1.3— 1.5 mm long, the tube ca. 0.1 mm long, the lobes 1.2—1.4 mm long, 0.4— 0.6 mm wide; staminodes like stamens but 0.7-0.9 mm long, the fila- ments ca. 0.1 mm long, the antherodes irregular, flat, deltate, 0.6-0.8 mm long and wide, apically acute, base not distinguishable; pistil 1.3-1.5 mm long; the ovary ellipsoid, 0.3-0.5 mm long, 0.2-0.3 mm diam.; stigma ligulate, 0.8—1 mm long, 0.2—0.3 mm wide, apically truncate; placenta 134 Stipa 17(1) N2WIO03SdS Fic. 2. A. Myrsine re Scheffer, showing leaf shape convergent with M. philippinensis, (H. Lam 3373). B-C. Myrsine avenis (Blume) A. DC. B. Pistillate branch, showing long fruiting medicals and inconspicuous venation of leaf blades (A. E/mer 142 ice C. elahaulate branch, showing leaf variation and sparse inflorescences (Loher 14097). D. M growth form from Palawan, Se resembling M. aralioides(C. Ridsdale 1084). E- E Myrsine philippinensis A. DC. E. Pistillate branch, showing globose fruit (M. Ramos GJ. Pagasio BS 3475 be F. Staminate branch, showing inflorescence and leaf apices (isotype).G Myrstne mindan is (Elmer) hie showing terete branchlets and globose fruits; lisctuee) H- L Myrsine avenis Blu ) A. DC. H. Pistillate branch (R. Williams 1346). I. Staminate branch. (C. R. Robinson me Black scale bar equals 1 cm, smallest unit on white scales equals | mm. Pipoty, Synopsis of Myrsine in the Philippines ie) subglobose, the ovules 2—3, uniseriate, immersed in the placenta. Fruzt depressed-globose, 5-6 mm long and in diam., densely and prominently black punctate and punctate-lineate apically, the same punctation types but pellucid basally. Distribution.—In the Philippines, this species is known from the Islands of Batan to Luzon(Pangasinan and Zambales provinces), Panay (Capiz Prov- ince), Negros (Negros Oriental Province), Samar Island (Western Samar Province), Dinagat (Surigao del Norte Province), Mindanao (Surigao City), and Palawan Island. Outside the Philippines, it is known from the Talaud Islands, Moluccas, Key Islands, Irian Jaya and Papua New Guinea, from sea level to 300 m. Ecology and conservation status.—Myrsine densiflora occars on ultramafic soils, just behind beaches in grasslands along streams at or near coastlines throughout its range. With ample habitat still remaining, the species does not appear to be threatened. The occurrence of Myrsine densiflora on Mt. Iraya on Batan Island might first be considered very unusual, but recent evidence shown by Richard et al. (1986) indicates that “Batan Island is noteworthy for the occurrence, within Mt. Iraya lavas, of ultramafic xeno- liths (harzburgites, dunites, Iherzolites, wehrlites, websterites) which have mantellic origin and had undergone strong interactions with metasomatic fluids.” Thus, similar geologic conditions, such as those found on Mindanao Island in the Provinces of Surigao del Norte and Surigao del Sur, and areas of southwestern Luzon (Zambales), allow us to predict that Myrsine densiflora may occur in these areas on ultramafic soils behind mangrove formations. Etymology.—The specific epithet refers to the dense clusters of slender pedicels and flowers, often appearing congested in early flowering. Common names.—“Tongog” (Bagobo language); “Maga” and “Magaspang (Bisayan, Panay dialects). ” Representative specimens examined: PHILIPPINES. Baran IsLANpD. Batanes Prov.: Mt. Iraya,122°00'E, 20°28'N, 750 m, 4 Jul 1989 (stam. fl), E. iis. 1003 (LBC), 800 , 9-16 Nov 1964 (fr), 8. oe & M. Sato 29059 {PNH 111132} (PNH). Luzon AND. Pangasinan Prov.: c 1907 (fr), M. Ramos s.n. ONY). Z a. Prov.: Acoje Mine Concession Area, ss oe uz, ca. 15°46'N, 120°00'E, low ee 23 May 1986 (pist. fl bud), C. Ridsdale & E. Reynoso 1440 (A, K, L, LBC, PNH). Panay IsLanp. Capiz Prov.: Below Mt. Salibongbong, Jul 1919 (fl bud), A. Martelino & G. Edaito BS 35612 (A). SAMAR IsLAND. Western Samar Prov.: Barangay — Marabut, 13 Feb 1992 (fr), sili sia PPI 6094 (A, BOG, BRIT, K, L, PNH). DinaGart IsLanb. Surigao del Norte rov.: Loreto Municipality, 26 Sep 1991 (fr), F Cae et al. PPI 3476 (A, BRIT, K, a oe , (fr), G. Ahern FB 444 (NY). NeGros Is-anp. Negros Oriental Prov.: Sibulan, Kabalinan, Lake Balinsasayao, 23 May 1991 (fr), E. Reynoso et al. PPI 1087 (A, BO, BRIT. K,L, PNH, US). PALAWAN IsLANbD. Palawan Prov.: N pe ca. 17 km N of Puerto Princesa, 27 Oct 1985 (stam. fl), C. Ridsdale 1084 (A, K, L, PNH). MINDANAO ISLAND: Surigao Prov. {del Norte ?}: Apr 1919 (fr), M. Ramos G J. Paseo BS 34625 (NY), Jun 1919 (fr), M. Ramos & J. Pasgasio BS 34755 (A). Surigao del Norte Prov.: Surigao City, — 136 Stipa 17(1) Apr—May 1911 (pist. fl), C. Prper BS 237 (US). INDONESIA. Surawesi [CELEBEs}. Talaud Islands: Miangas Paui Baronlo, me 1926 (stam. fl), H. J. Lam 3373 (BO, K); Karakelang, G. Piapi, 300 m 31 May 1926 (stam. fl), H. Lam 3258 (A, BO, K); Talaud, eas 100 , 13 Jun 1926 (stam. fl), H. Lam 34 20 (BOG, K); Salawati, P. Rumbobo, J. ee (BO, K). Motuccas: Obi. Paui Santari, 2 m, 28 Nov 1937 (pist. i. sole 5 (BO, K). IR1AN JAyA: West Irian Jaya: Seacoast below Sukarnapura (=Hollandia), 11 we 966 (stam. fl), A. Kostermans & Soegeng on on rh PAPUA NEW GUINEA: W Sepik Dis- trict: Selio Island Aitape subdistrict, °10'S, 142 °30'E, sea level, 31 silee! we (stam. fl), A. Millar & J. eee ene (sk, LAE). Apmirattry Is3 . Man Island. Manus Prov.: Lorengau Subprovince, Lorengau, 02°2'30" S, ae 30" E, ee m, 22 Mar 1981 (stam. fl), K. Kerenga et al. LAE 77490 (K, LAE); Milne Bay, Subprovince Misima, 11°22'S, 154°11'E, 22 Mar 1979 (fr), O. Gideon LAE 76035 (A, LAE); Archip. Key, 1889 (pist. fl), O. Warburg 21830 (A-2 sheets); Eastern Highlands District, 06°23'S, 145°S8'E, 2,300 m, 3 Jul 1963 (fr), T. Hartley 11955 (A, CSIRO, LAE). Myrsine densiflora is very closely related to M. philippinensis, but is readily separated from it by the coriaceous leaves scrobiculate above and below, the leaf apices not emarginate or retuse at tip, and the smaller pedicels. The species has populations which at times contain individuals with extremely coriaceous leaves and rather robust, albethey short, peduncles (C. Ridsdale 1084), resulting in confusion with Myrsine aralioides (Philipson) Pipoly. However, the depressed-globose fruit is unique among species of Myrsine in the region. Myrsine densiflora is restricted to ultramafic soils behind beach formations at low elevations, while M. ara/zoides is clearly a montane forest species. The edaphic specificity of the species and narrowly defined eco- logical tolerances account for its rather spotty distribution. While Myrsine densiflora may appear to be parapatric with M. philippinensis, the latter is a mangrove species, occuring in the mangrove proper with such other spe- cies as Aegiceras corniculatum (Myrsinaceae). a sa peregrina (Mez) Pipoly, comb. nov. (Fig. 3F, G). Rapanea peregrina Mez, Repert. Spec. Nov. Regni Veg. 16:424. 1920. Tyr: PHILIPPINES. MINDANAO IsLAND. Davao del Sur Prov.: Camp Keithley, Lake Lanao, Jun 1907 (fr), M. S. Clemens 917 (HOLOTYPE: PNH-destroyed; Lecrorype, here designated: US!; isoLecrotyPE: A!). — venosa Elmer, Leafl. Philipp. Bot. 2:672. 1910. non wees venosa (A, DC.) z, Pflanzenreich 9(1V. 236):386. 1902, nec Myrsine venosa A.DC., Ann. Sci. Nat., . > 16:86. 1841. Type: PHILIPPINES. Minpanao IsLanp. a Prov.; Mc. Apo Giedayay. District of Davao, 2,000 m, Aug 1909 (stam. fl), A. Elmer 11445 (Ho1o- TYPE: PNH-destroyed; Lecrotryre, here designated: GH!, isotecrorypes: A!, BISH!, L!, NY). Tree to 6 m tall. Branchlets: angulate, 2-3 mm diam., glabrous. Leaves: buds with margins sparsely translucent-ciliate, glabrescent; blades coria- ceous, asymmetric, oblanceolate to elliptic, 5-12 cm long, 1.5—3.5 cm wide, apically short-acuminate, the acumen ca. 3-7 mm long, basally obtuse or rarely acute, nitid, densely and prominently pellucid punctate above, pal- lid below, midrib immersed above, prominently raised below, the second- Pipoty, Synopsis of Myrsine in the Philippines 13} ary veins 21—32 pairs, prominulous above, less conspicuous below, the sub- marginal collecting vein ca. 0.5 mm from margin, entire, revolute, and bordered by numerous prominently pellucid punctations below along mar- gin; petiole marginate, 6-11 mm long, glabrous. Staminate inflorescence: umbellate, sessile to subsessile, 3—9-flowered, the peduncle obsolete to 2.5 mm long, ca. 1.0 mm diam.; floral bracts coriaceous, widely ovate, 0.8—1 mm long, 0.9-1.2 mm wide, apex rounded, the margin entire, sparsely glandular-ciliate, glabrescent; pedicels obconic, 3-5 mm long, reflexed in anthesis. Staminate flowers: chartaceous, 4-merous, 2.2—2.5 mm long; calyx cotyliform, 1-1.3 mm long, the tube ca. 0.1 mm long, the lobes ovate, 0.9-1.2 mm long, 0.5—0.8 mm wide, apically acutish to obtuse, densely and prominently red punctate, sparsely red punctate-lineate, irregular and glandular-ciliate along the margin; corolla rotate, 1.8—2.2 mm long, the tube ca. 0.2 mm long, the lobes 1.6—2 mm long, 0.8—1 mm wide, apically obtuse, cucullate, densely and prominently red punctate-lineate medially, red punctate near margins, on the margins densely glandular-granulose, stamens 1—1.5 mm long, the filaments ca. 0.3 mm long, developmentally fused to adaxial corolla tube and adjacent lobe, the anthers broadly ovate, 1-1.2 mm long, 0.8—1 mm wide, apically obtuse to rounded, basally sub- cordate, dehiscent by wide longitudinal slits, the connective brownish punc- tate apically and just above point of filament attachment, pistillode conic, ca. 1 mm tall, 0.8 mm wide, the placenta with two minute, infertile ovules. Pistillate inflorescence: \ike staminate but peduncle 1-5 mm long, ca. 1 mm diam. (breaking off at extreme length); pedicels 0.5—3.5, accrescent in fruit. Pistillate flowers: like staminate but 1.8—2.2 mm long, calyx 1.2-1.4 mm long, 0.5-0.4 mm wide, apex acute; corolla 1.8—2 mm long, the tube 0.5 mm long, the lobes 1.3—1.5 mm long, the apically acute; staminodes 1.3— 1.5 mm long, the tube adnate to the corolla tube, the antherodes deltate, malformed, 0.5—0.7 mm long, apically acute, basally sagittate, hollow; pistil obnapiform, the ovary globose, 1.3—1.5 mm long and in diam., the stigma stylopodic, lobed, 0.3—0.5 mm long, the placenta ovoid, the ovules 4, uniseriate. Fruit subglobose, longer than wide, 2.5—3.5 mm long, 2.2—3 mm diam., prominently red punctate-lineate. Distribution. —Myrsine peregrina is endemic to the Philippines, occurring from northern Luzon Island (Batan) southward through Luzon, Mindoro and Panay, to Mindanao Island from 2,090—2,800 m elevation. Ecology and conservation status.—Myrsine peregrina occurs in local popula- tions, at the interface of the submontane and montane “mossy forests,” in areas with high incident light. The type of Rapanea peregrina was collected in a swamp near Lake Lanao. I have observed it to be extremely localized, and where it occurs, there are approximately 12 individuals per hectare, rarely reaching 9 cm DBH. This species seems to thrive in areas of distur- Sipa 17(1) — oS om | i SPECIMEN Ms pertaria a DATE. see st ! or Pipoty, Synopsis of Myrsine in the Philippines iReky, bance, such as gaps produced by natural or man-made treefalls, so it ap- pears resilient enough to consider it not threatened at this time. Etymology.—The specific epithet means “travelling and migratory,” per- haps alluding to this species broad distribution atop mountains through- out the country. Common names.—‘Marimbakawan” (Palawefio and Tagbanwa languages), “Korios” (Bisayan-Cebuano dialect)- generic term related to long terminal bud; “Tongog” (Bagobo language). Specimens examined: PHILIPPINES. Baran IsLanp. Batanes Prov.: Mt. [raya, Jun— Jul 1930 (stam. fl), M. Ramos BS 80268 (A). LUZON ISLAND. Mountain Prov. {Bontoc}: Mt. Caua, Mar 1920 (stam. fl), M. Ramos & G. Edafto BS 37982 (K, PNH); Bontoc Sub- Prov., Bauko, On slopes of Mt. Data 2,100—2,500 m, 10-13 Feb 1964 (fr), W. Stern G J. Rojo 2299 [PNH 92363](CLP, PNH). Nueva Ecija Prov.: Mt. Umingan: Aug—Sep 1916 (fr), M. Ramos & G. Edatio BS 26397 (A, US). Minporo Istanp. Mindoro Oriental Prov.: Me. Halcon, Mar 1922 (pist. fl, fr), M. Ramos G G. Edafio BS 40712 (A, K, L, PNH, US), NE summit, 2,000—2,450 m, Apr-May 1995 (ster), E. Mandia 555 (CAHUP, DLSU, Panay IsLANb. Capiz Prov.: Mr. Bulilao, Jun 1919 (stam. fl), A. Martelino & G. Edaito BS 35682 (A). Iloilo Prov.: Iwa Iaya {[Iwaya } to Coyaoyao {Koya}, 10 Sep 1936 (stam. fl), T. Hosokawa 8560 (BISH). MINDANAO IsLANb. Bukidnon Prov.: Mt. Katanglad, Mar 1949 (fr), M. Sulit PNH 10054 (A, L, PNH, US), 14 Apr 1949 (fr), M. Sulit PNH 10125, (A, K, L, PNH), Mar-Apr 1949 (fr), G. Edatto PNH 11595 (A, PNH); Municipal- icy Lantapan, a Sungco, Mt. Kinasalapi, part of Kitanglad Mt. Range, W of Analib River, ca. 08°00'05" N, 124°30!26" E, 2,090-2360 m, 16 Nov 1994 (fr), J. Pipoly et al. PPI 16584 an PNH); Bukidnon, Kaatoan, above Cinchona plantation, 1,000 m, 10 Oct 1953 (ster.), B.B. Britton 421 {PNH 19646} (L, PNH). Davao Prov.: Mt. McKinley, E slopes, 2,250 m, 25 Sep 1946 (fr), G. Edaito 1051 {PNH 1564} (L, PNH); Mt. Apo, near Sulphur Spring, 2,100 m, 15 Nov 1946 (fr), G. Edaito 1536 {PNH 1540} (L, PNH). Davao del Sur Prov.: Mc. Apo, [Todaya], N slope, Lake Lanao, 2,100 m, 30 Oct 1946 (fr), G. Edafto 1232 {(PNH 1412} (L, PNH); Mt. Kampalili, Mar-Apr 1949 (fr), G. Edaito PNH 11565 (A, K, L-2 sheets, PNH, US); PNH 11595 (A, L, PNH, US); Mt. Apo, Baklayan, 1,800 m, 11 Nov 1946 (fr), G. Edaiio 1445 (PNH 1557) (A, L), 15 Nov 1946 (pist.fl), G. Edafio 1535 (PNH 1553) (A), Mt. Apo, 1,970 m, 13 Mar 1956 (fr), J. Pancho 2587 {CAHUP 4124} {PNH 34613} (CAHUP, PNH). Misamis Oriental Prov.: NE slopes of Mt. Malindang, Lake Duminagat, 1,763 m, 5 Jan 1993 (fr), F Gaerlan et al. PPI 10949 (A, BO, BRIT, K, L, PNH); Mt. Malindang, M. Mearns G W. Hutchinson FB 4573 (FS). North Cotabato Prov.: {Todaya} Mt. Apo, 2,134 m, Aug 1909 (pist. fl, fr), A. Elmer Fic. 3. A-B. Myrsine aralioides (Philipson) Pipoly. A. een branch, showing ee ce J. & M. Clemens 27801). B. Staminate branch. (J. & {. Clemens SO807).C. Myrsine glandulosa (Elmer) Pipoly, pistillate branch, showing e pees peduncles (isotype). .Myrsine fastigiata (Elmer) Pipoly, aon ite branch. (M. Ramos & G. Edatto 38494 (isotype)F-G. Myrsine peregrina (Mez) P es F. Pistillace ies showing leaf texture hd .Myrsine fastigiata (Elmer) (isotype, Ree venosa Elmer). G. Pistillate | Pipoly, staminate ranch, showing crowded ae (M. me 7564). 1. Myrsine penibukana (Philipson) Pipoly, staminate branch showing perennating peduncles forming “short shoots” J. & M. Clemens 50295). Black scale bar equals | cm, smallest unit on tie scales equals 1 mm 140 Sipa 17(1) 11456 (A, BISH, GH, K, L, NY, PNH, US); Mt. Apo, 7,000 ft, 15 Nov 1946 (pist. fl, fr), G. Edafio PNH 1540 (A, oe ee Nov rae a fl), G. Edafio PNH 1553 (A, PNH); Kidapawan Municipality, M th roject, Lake Venado, 11°32'N, 125°21'E, Oct 1992 (fr), D. cee 870 (PUHD). Site a 06°59.8'N, 125°14'E, 1,725 m, Oct 1992 (fr),D. Lagunzad 961 (PUH). South Cabs Prov.: Mt. Matutum, 7,000 ft. Apr 1932 (fr), M. Ramos & G. Edatio BS 85048 (A), 15 Apr 1992 (fr), E Gaerlan et al. PPIS361 (A, BRIT, K, L, PNH, US). PALAWAN IsLaNnp. Palawan Prov.: Lipun Island, Lipun Peak, 215 m, 28 Apr 1964 (fr), D. Mendoza PNH 91353 (L, PNH) Myrsine peregrina can be confused with M. densiflora, but is easily distin- guished by the longer, obconic pedicels, short-acuminate and asymmetric leaf blades 8. Myrsine mindanaensis nae Pipoly, comb. nov. (Fig. 2G). seg mindanaensts Elmer, Leafl. Philipp. Bot. 2:671. 1910. Type: PHILIPP MINDANAO ISLAND. North es Prov.: [Todaya], Mt. Apo, Aug 1909 i 7 cee 11332 (HoLoTYPE: PNH-destroyed; LEcrorypre: K !; 1sorecrorypeEs: A!, ISH!;GH!; L!, NY!). Shrub or cree to 8 m tall. Branchlets: terete, longitudinally ridged, 2.5— 3.5 mm diam., sparsely and minutely rufous-papillose, early glabrescent. Leaves: buds essentially glabrous, the margins minutely rufous glandular- ciliolate, early glabrescent; blades chartaceous, slightly asymmetric, nar- rowly oblanceolate to narrowly oblong, 4.5—-11.5 cm long, (1.8—)2—3.5 cm wide, apically acuminate, the acumen 0.5—1 cm long, basally acute, decur- rent on the petiole to base, scrobiculate and inconspicuously punctate above, densely and prominently black punctate and punctate-lineate below, the midrib deeply impressed above, prominently raised below, the secondary veins all inconspicuous, ca. 8-10 pairs, the submarginal collecting vein inconspicuous, ca. | mm from margin, the margin slightly revolute through- out; petiole marginate to base, 6-7 mm long, flat oe Staminate inflorescence: unknown. Pastillate inflorescence: a sessile umbel, 3—9-flowered; peduncle obsolete to 2 mm long, ca. | mm diam.; floral ae chartaceous, oblate, 1.3—1.5 mm long, 2—2.2 mm wide, apex broadly rounded, densely glandu- lar-granulose at first, glabrescent, densely and prominently red glandular- punctate and punctate-lineate, the margin opaque, densely glandular-cili- ate; pedicels cylindric, 3-5 mm long. Pistillate flowers: unknown. Fruiting calyx 4-merous, 0.8—1 mm long, the calyx tube ca. 0.1 mm long, the lobes ovate, 0.7—-0.9 mm long, 0.5—0.7 mm wide, apex subacuminate, densely red punctate and punctate-lineate medially, the margin flat, opaque, en- tire, minutely glandular-ciliolate, glabrescent. Fruit globose, 2—3 mm long and in diam., densely and prominently red punctate and punctate-lineate, the persistent stigma ligulate, 4-winged. Distribution.—Endemic to the Philippines, from southern Luzon Island — Pipoty, Synopsis of Myrsine in the Philippines 141 (Camarines Sur Province) southward through Leyte Island (Biliran Subprovince) to Mindanao Island (Davao del Sur and North Cotabato prov- inces), at 895—2,200 m elevation. Ecology and conservation status. —Myrsine mindanaensis is a narrow endemic found from southern Luzon Island southward through Leyte to Mindanao. The species appears to be a subcanopy species on Mt. Apo, in the mid- montane forest dominated by Lithocarpus spp. and Agathis philippinensis. It is locally common in secondary growth areas on Leyte. However, no studies have shown whether M. mindanensis can withstand prolonged disturbance. Therefore, pending further study, I prefer to describe its conservation sta- tus as threatened. Etymology.—The specific epithet refers to the site of original description, from the island of Mindanao. Common names.—“Dipalla” (Bagobo language, Ramos BS 41559); “Tongog” (Bagobo language); “Baliyuk” (Bagobo language, Madulid 1992). Specimens examined. PHILIPPINES. Luzon IsLaAnb. Camarines Sur Prov.: Tinambac, Mt. Isarog, Haribon Foundation, Tinambac Caw lees access trail N to Ulag Ganabo, Balagbag Kulod facing Isarog Tilhang, Ulag and vicinity, 895-1020 m, 20 Feb 1993 (fr), B. Hernaez & M. Cajano {CAHUP 6092, 60893, ave (CAHUP). Leyte IsLanbs. Biliran Subprov.: Libtong, Naval, 12 Aug 1992 (fr), Barbon et al. PPI 8601 (A, BOG, BRIT, K, L, PNH). Leyte Prov.: Cabalian, Dec 1922 (fr), M. Ramos BS 41559 (A, NY, PNH, US). MINDANAO IsLAND. Davao del Sur Prov.: Davao, Aug 1933 (fr), R. Kanehira s.n. (NY). North Cotabato Prov.: Napces Mt. Apo, Sep 1909 (fr), A. E/mer 11820 (A, BISH, GH, K, L, NY, PNH, US); Mt. Apo, NW slopes, along Marber River, 07°05'N, 125°14'E, 1,600 m, 23 Oct 1990 (ster.), L. Co 3125 (A, PUH, L); Kidapawan Municipality, Mt. Apo Geothermal Project, near Lake venado, 07°10'N, 125°16'E, 1,800—2,200 m, Gee 1992 (ster.), D. Lagunzad 916 (PUH). Myrsine mindanensis is closely related to M. philippinensis but can be dis- tinguished by its scrobiculate upper leaf surface, minutely rufous papillose branchlet apices, and minute calyx. 9, Myrsine philippinensis A. DC. in DC., Prodr. 8:94. 1844. (Fig. 2E, . Rapanea ela (A. DC.) Mez in Pacle, Pflanzenr. 9 (IV. 236):364. 1902. Type: PHILIPPINES. Luzon Istanp. Albay Prov.: 1841, H. Cuming 868 (HOLOTYPE: G, n.v., ae A!; isoTyPEs: K-2 sheets!, L!). i> Shrub or small tree to 5 m tall. Branchlets: terete, 2.5—3.5 mm diam., essentially glabrous. Leaves: buds minutely red glandular-papillate and trans- lucent glandular-granulose, early glabrescent, the margin minutely trans- lucent ciliate, early glabrescent; blade coriaceous, symmetric, oblanceolate or oblong, rarely elliptic, (S—)6-10(-12) cm long, 2—3(—3.8) cm wide, apically obtuse to rounded, the uppermost tip slightly emarginate, basally cuneate, decurrent on the petiole, smooth and inconspicuously pellucid punctate above, black or brown punctate and punctate-lineate below, the 142 Sipa 17(1) midrib slightly raised above, prominently raised below, the secondary veins 13-19 pairs, not visible above, inconspicuous below, the marginal collect- ing vein ca. | mm from margin, flat, entire, and bordered by numerous prominent black punctations along the margin; petiole canaliculate and marginate, 6-8 mm long, glabrous. Staminate inflorescence: umbellate, 9— 16-flowered; peduncle 1.5-2 mm long, 1-1.5 mm diam.; floral bracts chartaceous, ovate, 0.8—1 mm long, 0.3—0.5 mm wide, apically obtuse, prominently and densely pellucid punctate-lineate, the margin entire, glan- dular-ciliate; pedicels cylindric, 3.2-5.0 mm long. Staminate flowers: 4- merous, membranaceous, 2.6—3.6 mm long; calyx cotyliform, 1.2—1.5 mm long, the tube ca. 0.1 mm long, the lobes ovate, 1.1—1.4 mm long, 0.4—0.6 mm wide, apically obtuse, prominently black punctate near apex, erose and densely glandular ciliate along the margin; corolla 2.5—-3.5 mm long, subrotate, the tube ca. 0.3 mm long, the lobes oblong, 2.2—3.2 mm long, 0.6-0.8 mm wide, apically rounded, prominently and densely black punc- tate and punctate-lineate, densely glandular-granulose along the margin; stamens 1.7—2 mm long, the filament adnate to corolla tube, 0.3—0.5 mm long, the anthers oblong, 1.3-1.5 mm long, apically long-apiculate, the apiculum densely glandular-barbate, basally subcordate, dehiscent at first by large subapical pores, opening into wide longitudinal slits, the connec- tive and apiculum prominently black punctate ventrally and dorsally; pistillode conic, subglobose, ca. 0.1-0.2 mm long, 0.4—0.5 mm diam., hollow, the stigma apiculate. Pisti/late inflorescence: like staminate but pe- duncle 2—4 mm long, to | mm diam.; pedicels 2—3 mm long, accrescent in fruit co 5 mm. Pisti/late flowers: like staminate but 1.7—2 mm long; calyx 1.0-1.1 mm long, the tube ca. 0.2 mm long, the lobes widely ovate, ca. 0.8-0.9 mm long, 0.9—1.0 mm wide, apically rounded, the margin irregu- lar, densely glandular-ciliate; corolla 1.7—1.9 mm long, the tube 0.1—0.2 mm long, the lobes 1.3—1.5 mm long, 0.3—0.5 mm wide; staminodes 1.3— 1.5 mm long, the filaments 0.1—0.2 mm long, the antherodes lanceolate, 0.8—1 mm long, 0.2—0.4 mm wide, apically minutely apiculate, base sub- cordate; pistil obnapiform, 2.2—2.5 mm long; ovary globose, 1—1.2 mm long, 0.4—0.6 mm diam., the stigma ligulace, 1.4—1.6 mm long, the base stylopodic at the ovary apex, translucent, eglandular, the placenta obpyramidal to obovoid, ovules 3-5, uniseriate, exposed laterally. Fruit globose, 2.5—4 mm long and in diam., densely and prominently pellucid — punctate and punctate-lineate. Distribution.—Endemic to the Philippines, from southern Luzon Island (Batangas, Camarines Norte and Quezon provinces), southward through Mindoro, Sibuyan and Panay Island, then wesward to the Cuyo Island Group and eastward to Mindanao, including Sulu Island, at sea level to 305 m elevation. Pipoty, Synopsis of Myrsine in the Philippines 143 Ecology and conservation status. —M yrsine philippinensis is very common along coastal arelike secondary vegetation, and on sands in mangrove swamps, and as such does not appear to be threatened at the present time. Etymology.—The specific epithet refers to the distribution of this en- demic species. Common names.—Myrsine philippinensis has more common names than any other species in the genus, owing to its high frequency and visibility in the mangroves along coastlines. The following are common names from label data with no languages given: “Mamumboga’ (Panay Islands: Guimaras Subprovince, Guimaras Island); “Hampuy puyos: ’ (Panay Islands, Hoilo Prov.); “Maga” (Luzon Island: Quezon Prov.); “Naling Saging” (Samar Island: Northern Samar Prov.: Waray). Other names (Madulid 1992): “Anumbaga’ (Bisayan-Panay dialect); “Go-od go-od” (Tagalog language); “Gutimomo” (Ibanag language); “Hanigad” (Bisayan-Cebuano dialect); “Kalumanay” (Tagalog language); “Lipote” (Bukidnon-Binukid Manobo dialect- prob- ably from vicinity of Cagayan de Oro); “Luklukso” (Ibanag language); “Maga, Magaspang” (Bisayan-Panay dialect- probably mistaken and refers to Myrs7ne densiflora), “Malatukba” (Batangan, Mangyan dialect); “Marupig” (Negrito language); “Supak” (gorot language); “Supek” (Kankanay language). Specimens examined: PHILIPPINES. Luzon Is_anp. Batangas Prov.: 1841 (pist. fl H. Cuming 1601 (A, K). Camarines Norte Prov.: Paracale, Nov 1884 (fr), S. Vidal 1516 (K, L). Quezon Prov.: Polillo Island, Bo. Lube, sea level, Jun a (bisex. and pist. fl), F yes 1057 (PNH 3715) (A, L); Real, Saale me an, NBG, 4 Sep 1991 (fr), Baron al. PPI 2267 (A, BRIT, PNH); Infanta, 3 m, g 1959 (stam. fl), L. Orantia LBC 348 ee Tagkawayan, sea level, 20 Mar 1951 eee fl), S. Jacoban 20 {LBC 966} (LBC), : Jan 1951(stam. fl), E. Agaceta 6 {LBC 917} (LBC); Calauag Bay, Caluag Municipality, near sea level, Mar 1990 (stam. Al), D. Lagunzad 758 (PUH). Minporo IsLaANbD. Mindoro Ori- ental Prov.: Baco, Jan 1903 (pist. fl), E. Merri// 1206 (K, NY). SipuyANn IsLaAnb. Prov. Romblon: Magallanes (Mc. ie aes along P er River, 305 m, Apr 1910 ae fl), A. Elmer 12295 (A, BISH, GH, _, NY, PNH, US). Cuyo IsLAND Group: Palaw Prov.: Pamalican Island, Hill 38, summit area, 1 1° et 'N, 120°43.6'E, <50 m, 5 Apr 1990 (ster.) L. Co. 3073 (PUH). PANAy IsLAND. Guimaras Subprov.: Guimaras Island, Mar. ne (ster.), 8S. Vidal 3183 (K), Jan 1904 (ster.), ae Sine FB 272 (K, NY, PNH, US), ie i 4 (stam. fl), J. Gammill FB 276 UK, NY, US), Feb 1904 (fr), J. Ganmill PB ; K, NY, PNH), Feb—Mar 1950 (fr), M. Sw/it sn (PNH 11769} (A, K, L, PNH, US), in. M. Sulit 11800 oh PNH, US), Gimaras, s.¢. (Bud), S. Vidal 3156, 3183 (A). LEYTE oe ee > 19 Jun 1915 (pist. fl), C. A. Wenzel 1368 (A). SAMAR ISLAND. Sam Prov.: . apul on and, 27 Mar 1957(ster.), ¥. Kondo & G. Edatio 113 {PNH 590. ae a (BISH, PNH); Tarangan, sea level, s.d. (stam. fl), E. Rodulfa EVR-7 {LBC 3792} (LBC). Ae IsLAND. Surigao Prov.: Jun 1919 (fr), M. Ramos GJ. Pasgasio BS 34755 (K, PNH). Joro [Suu] IsLanp. Samales Group: ARMM, 15 Oct 1915 (stam. fl), M. S. Clemens 9332 (A, L). Myrsine philippinensis belongs to a complex of species including M. umbellulata (Wallich) A. DC., M. sumatrana Miquel, M. cochinchinensis A. DC., M. korthalsiz Miquel, and M. rawacensis A. DC. Field studies of popu- 144 Sipa 17(1) lation biology and intrapopulational variation will be necessary to deter- mine if the quantitative differences in pedicel length, floral number, and leaf reticulation provide useful characters upon which taxa may be consis- tently separated. These highly variable quantitative characters, combined with differences attributable to sex expression (staminate, pistillate, bi- sexual) may account for the proliferation of names in this group. In par- ticular, there appears to be only minor petal ornamentation differences which tentatively separate Myrsine philippinensis from M. cochinchinensis. It is hoped that current studies underway across the entire Malesia Floristic Province will remedy the current taxonomic chaos. In the Philippines, Myrszne philippinensis may be superficially confused with M. mindanensis, but is easily recognized by its glabrous branchlets, longer calyx and mangrove habitat. It is probably most often confused with M. densiflora, from which it is distinguished by the extremely small, glo- bose fruit, more robust, longer pedicels, more numerously-flowered inflorescences, and much longer calyx. LO. ae avenis (Blume) A. DC., Trans. Linn. Soc., Bot. 17:107. 1834. (Fig. 2H I). Ardisia avenis Blume, Be dr. 691. 1825. Repenes avenis (Blume) Mez in Engler, Piven OTV. 236):357. 1902. Type: a aad Java: sine loc. esp. (fr), H. Zollinger 883 (HOLOTYPE: L, n.v.; ISOTYPE 7 Tree to 8 m tall. Branchlets: angulate, 3-5 mm ‘diam., densely rufous tomentose at first, glabrescent. Leaves: buds densely rufous tomentose and with minute rufous sessile scales, the margins densely rufous-ciliate, gla- brescent; blade coriaceous, symmetric, elliptic to oblong, rarely narrowly oblanceolate, (3.8—)4—-7(—9.5) cm long, (1.0-)2—3(-4) mm wide, apically sharply acute, with a small emargination at tip, basally obtuse to rounded, not decurrent on the petiole, dull but prominently pellucid punctate above, densely and prominently pellucid punctate below, bearing minute rufous sessile scales when young, glabrescent, midrib immersed above, to base of petiole, prominently raised below, the secondary veins not visible above or below, entire and subrevolute, especially basally, along the margin; petiole marginate, 10-15 mm long, glabrous. Staminate inflorescence: subsessile, the peduncle 1—2 mm long, ca. 1 mm diam.; floral bracts chartaceous, ovate to suborbicular, 0.8—1 mm long, 0.6—0.8 mm wide, apically obtuse, densely and prominently black punctate, the margin glandular-ciliate; pedicels cylindric, (2—) 3-11 mm long. Staminate flowers: 4-merous, chartaceous, 1.5—1.8 mm long; calyx cupuliform, 1—1.2 mm long, the tube ca. 0.2 mm, the lobes ovate, 0.8—1 mm long, 0.7—0.9 mm wide, apically obtuse, promi- nently red punctate-lineate medially, entire, densely and prominently red punctate and sparingly glandular-ciliolate along the margin; corolla cam- panulate, 2.3—2.5 mm long, the tube ca. 0.2 mm long, the lobes asym- Pipoty, Synopsis of Myrsine in the Philippines 145 metrical, oblong, 2—2.3 mm long, 1-1.3 mm wide, apically obtuse to rounded, densely and prominently red punctate and punctate-lineate me- dially, densely red punctate and glandular-granulose along the margin; stamens 2—2.2 mm long, the filaments adnate to the corolla tube 0.2—0.3 mm long, the anthers ovate, 1.8—1.9 mm long, 1.2—1.5 mm wide, apically acute, almost apiculate, basally cordulate, the apiculum and connective darkened; pistillode conic, ca. 1 mm long and wide, hollow. Pist/late inflorescence: as in staminate but 10—15-flowered; peduncle 2.5—3 mm long, mm diam.; floral bracts suborbicular, 1-1.2 mm long and wide, apically widely rounded, the margin glandular ciliolate; pedicels 1.5—3 mm long. Pistif/late flowers: like staminate but calyx 1-1.3 mm long, the tube ca. 0.3 mm long, the lobes widely ovate, 0.8—1 mm long, 0.8—l1 mm wide, apically obtuse; corolla 2.3-2.5 mm long, the tube ca. 0.3 mm long, the lobes oblong, 2—2.2 mm long, 0.8—1 mm wide, apically rounded, densely and prominently red punctate and punctate-lineate medially, punc- tate and glandular-granulose along the margin; staminodes 2 mm long, the filament fused to the corolla tube 0.3 mm long, the antherodes ovate, 0.5—0.7 mm long and wide, the apically acute, slightly apiculate, non functional, basally deeply cordate, the connective darkened apically and on filament back; pistil tetragonal in cross-section, 1.2-1.6 mm long; ca. 0.8—1 mm diam., densely black punctate-lineate, the stigma 4-lobed, the lobes perpendicular, ca. 0.2 mm long, the placenta ovoid, the ovules 4, in one series. Frit globose, 1.5—2.5 mm long and in diam, red when fresh, densely and prominently orange and pellucid punctate, the persistent stigma base appearing stylopodic. Distribution.—Myrsine avenis is found from northern Luzon Island south through Mindoro to Negro and finally, Mindanao Island.Outside of the Philippines, it is known from Brunei, Sabah, Sarawak, Java, Sumatra, Buru, East and West Kalimantan, and Sulawesi, at 862 —3,000 m elevation. Ecology and conservation status. —Myrsine avenis is most frequently found on windswept ridgetops in forests dominated by Ericaceae and Podocarpaceae, in forests termed “mossy forest,” “montane forest” or “up- per submontane forest.” The climate is extremely hostile and there is pre- cious little commercially valuable timber on these sites. However, with increasing demands for telecommunication installations on the tops of many mountains throughout Malesia, this species may be considered threatened. Etymology.—The specific epithet, ‘avenis’ refers to the secondary veins, not readily visible from either the abaxial or adaxial side of the leaf. Common names.—‘Berig” (Bisayan, Panay dialect, Madulid 1992); “Paspas” (Bukidnon, Binukid Manobo dialect, probably from the north- ern portion of the Kitanglad Mountain Range, Madulid 1992). 146 Stipa 17(1) Specimens examined: PHILIPPINES. Luzon IsLanp. Baatan Prov.: Lamao River, Mt Mariveles, May 1905 (stam. fl), R. Meyer 3114 (K, PNH). Benguet Prov.: Mc. Pulogloco, Sep 1921 (pist. fl, fr), M. Ramos & G. ae BS 40394 (US); Mount = Tomas, 26 oe 1935 (fr), M. 8S. Clemens 51876 (A CH), Mar 1904 (stam. fl), A. Elmer S811 (A PNH, US), Jun 1904 (fr), A. Elmer pe (K, PNH, US), 3 Dec 1953 i E. Walker ar (US), 28 Oct 1904 (pist. fl, fr), R. Williams 1346 (GH, K, NY, US); Mar 1907 (fr), A. E/mer 8745 (L, NY, US); sine loc. esp., Mar 1916 (fr), C. Garcia 25507 (US); Baguio and a May 1911 (bisex. & pist. fl), C. B. Ee ta 14088 (K, L, PNH, US), Mar {1907]1913 (fr), A. Elmer 14286 (BISH, GH, NY, US); Mc. Tonglon, Bagio, Dec 1908 (fr), i Merritt FB 14153 (US). te ae noe Prov.: Vicinity of Paracale [probably near Parang}, sd. (fr), S. Vidal 1526 (L); Camarines Sur Prov.: Tinambak, Haribon Founda- tion for the Conservation of Natural Resources, Luzon, Mt. Isarog National Park, Tinambac Cawayan access trail N to Ulag Ganabo, Balagbag Kulod facing Isarog Tilhang. Ulag and vicinity, 895—2,000 m, 20 Feb 1993 (pist. fl bud), B. Hernaez & M. Cajano CAHUP 60019 (CAHUP), Balagbag Kulod facing Isarog Tilhang Trail to Kabagtuan, Gapugapuan and vicinity, 895-1065 m, 22 Feb 1993 (pist. Al bud), Hernaez & es ey eek CAHUP 60020 (CAHUP). Ilocos Norte Prov.: Mt. Palimlim, M. Ramos BS 33285 (NY). Laguna Prov.: Mt. Banahao aie. ao, ca. 14°04'N, 121°29'E], Jan 1913 (fr ape BS 19576 (US), Apr 1925 (fr), M. Salt BS 30073 (NY); Los Bafios, Mt. ane ling, Mar 1906 (ster), A. Loher 6140, Pie 917 (ster), A. ae ae (A, K, L, NY, PNH, US), (fr), A. Ed/mer 18270 (A, BISH, Ik, L, NY, PNH, 1 Jul 1964 (stam. fl), J. Hernaez 867 (CAHUP 18393} (CAHUP); Peak 2, 1060 m, Ee ane 1966 (stam. A), B. Hernaez 867 {CAHUP 58864, 38865, 58866} (CAHUP), 10 Jun 1966 (stam. fl), N. Orlido CAHUP 17927 (CAHUP), summit, Mt. Makiling, 1,100 m, 18 Jan 1913 (fr), F Gates 5196 (CAHUP, UC); Mc. Maquiling, wee 1913 (fr), Forestry School, UP Los Battos FB 20145 (US). Laguna/ Quezon Prov. border: Mt. Banaho, s.¢. (fr), 8. Vidal 15 1661s (A). Rizal Prov.: Mt. Susong- Dalaga, Aug 1917 (fr), M. Ramos & G. Edafio BS 29282 (A, K, US); Montalban, Sep 1909 (stam. fl), A. Loher 12009 (A), (fr), A. Loher 12038 (A); Sumag, Apr 1914 (stam. fl), A. Loher 14097 (A), sine loc. esp. May 1907 (fr), M. Ramos BS 2650 (BISH, US). Quezon Prov. [Tabayas} Dec 1993 (fr), A. Lober 3850; (UR, PNH, US); Banahao, Feb 1906 (ster.), A. Loher, 6125 (K, PNH), s.d@. (fr), Videl 1516 (A, PND: Lucuban, 1,829 m May 1907 (fr), A, Elmer 9351, 9352 (A, K, L, NY, PNH, US). Mountain Provy.: Bayninan, Banaue, Ifugao, 1,219 m, 26 Mar 1963 (pist. fl, fr), H. C. Conklin & Buwaya I-1023 {PNH 79612} (K, L, PNH). Mindoro ae Oriental Mindoro Prov.: Mt. Hi alcon, 862 m, 16 Feb 1948 ( (fr), G. Edato 192 (PNH 3574) (A, L); Mt. Yagaw, S slopes, 400 m, 30 Sep 1953 (ster.), H. Conklin 529 {PNH nee (L), Mt. Yagaw, E slope, SE s ue xe, 330 m, 28 Dec 1953 (ster), H. Conklin 8604{PNH 19321} (L, PNH). NeGros Istanb. aoe Occidental Prov.: SW face of Mt. Canlaon near summit crater, above Guintubdan, 31 Mar 1992 (fr), B. Stone et al. PP10471 (A, BRIT, PNH), Apr 1910 (fr), E. Merr#// BS 6998 (US). MINDANAO ISLAND. North Cotabato Prov.: Todaya, Mt. Apo, 1905 (fr), R. Williams 2603 (A) — — Myrsine avenis belongs to a complex of perhaps 15 taxa, within which Myrsine affinis A. DC. is the name most often misapplied to it. It is prob- ably most closely related to Myrsine capitellata Wallich (non alior), differing in much longer pedicels, greater numbers of flowers per inflorescence, and the pubescent branchlet apices. In the Philippines, Myrs/ne avenis was frequently labelled “Myrsine philippinensis” by such workers as Elmer, Loher, and even Merrill on occa- sion. This led to much confusion as later collections were apparently matched Pipoty, Synopsis of Myrsine in the Philippines 147 to those determined by these prodigious students of the Philippine flora. However, Myrsine avenis resembles M. philippinensis only in the highly vis- ible pedicels. In addition to the pubescent branchlets of M. avenis, it is restricted to upper montane habitats, while M. philippinesis is strictly a mangrove species. 11. Myrsine penibukana (Philipson) Pipoly, comb. nov. (Fig. 31). Rapanea pentbukana Philipson, London J. Bot. 78:105. 1939. Type: MALAYSIA. Sabau [Brit- isH NortH Borneo}: Mount Kinabaulu: Penibukan, flat place on ridge at 1,219 m., 18 Oct 1933 (stam. fl), J. & M. Clemens 20764 (HOLOTYPE: BM, n.v.; IsoTyPEs: A*!; NY*!). *The sheets ac A and NY bear the number J. G M. Clemens 40764, written in pencil at the bottom of the label, but the A sheet was annotated by a a before ence of the species. Given that the number was not included y him as a paratype along with - other collections, and that there are several other ne en out and partially corrected on the label, it is assumed that the A and NY sheets’ number is a typographical error and that they are, isotypes. Tree to 20 m tall. Branchlets: terete, 2.5—3 mm diam., sparsely ferrugin- eous glandular-granulose, glabrescent. Leaves: buds minutely and sparsely ferrugineous glandular-granulose, glabrescent, the margins rufous glandu- lar-ciliate; blades thinly coriaceous, Sas ae oblanceolate, elliptic or obovate, (3.5—)7—10 cm long, (2.2—)2.5—4 cm wide, apically obtuse, ba- sally acute, decurrent on the petiole to base, subnitid and conspicously but not prominently pellucid punctate above, pallid and conspicuously black punctate-lineate below except prominently black punctate between sub- marginal vein and margin, the midrib impressed above, prominently raised below, the secondary veins inconspicuous on both sides, 9-12 pairs, the submarginal collecting vein ca. 0.5 mm from margin, marginally flat and entire; petiole marginate, flat above, 6-8 mm long, glabrous. Staminate inflorescence: fasciculate; peduncles forming “short shoots” (3.5—)4—12.5 mm long, 1-2.5 mm diam.; floral bracts coriaceous, oblate, 0.5—0.7 mm long, 1—1.2 mm wide, apically broadly rounded, medially prominently pellucid punctate, with hyaline, scarious, erose-fimbriate, and long white glandu- lar-ciliolate margin; pedicels obsolete to cylindric and 0.5(—0.7) mm long. Staminate flowers: 5-merous, chartaceous, 2.8—3 mm long; calyx cupuliform, 5mm long, the tube ca. 0.1 mm, the lobes ovate or oblong, 1.2—1.5 mm long, 0.5—0.7 mm wide, apically obtuse to acutish, densely pellucid to orange punctate and punctate-lineate medially, irregular, entire, and glan- dular-ciliate along the margin; corolla campanulate, 2.8—3 mm long, the tube 0.5—0.7 mm long, the lobes elliptic to oblong, 2.3—2.5 mm long, 0.7-1.0 mm wide, apically obtuse, cucullate, densely and prominently pellucid punctate-lineate medially, prominently red punctate along entire margin, glandular-granulose within and on margin, the margin entire; sta- mens 1.8—2 mm long, the filaments completely fused to corolla tube, ca. 148 Stipa 17(1) 0.5 mm long, the anthers ovoid, 1.3—1.5 mm long,] mm wide, apex api- culate, base cordate, the connective prominently red punctate-lineate; pistillode vestigial, conical and hollow, ca. 0.5 mm long, or absent. Pzstz/- late inflorescence: unknown. Fruit subglobose, 4.5—5 mm long, 3.8-4 mm diam., densely and prominently pellucid punctate-lineate. Distribution.—Known only from the type locality, Mt. Kinabalu, Sabah, and Luzon Island, in Abra, Quezon and Rizal provinces, Philippines, at 1 ,000—1,200 m. Ecology and conservation status.—Myrsine penitbukana is a highly restricted species new to the flora of the Philippines. At this time, nothing is known of its ecology or its conservation status. Etymology.—The specific epithet refers to the type locality, Mt. Penibukan, a part of the Mt. Kinabalu Range, in northen Sabah, on the island of Borneo, Malaysia. Common names.—W hile no common names have been reported for this taxon per se, it is expected that “Tongog” (Bisayan-Panay dialect, and Bagobo languages) would be used for this taxon, as for all members of the genus Myrsine. Specimens examined: PHILIPPINES. Luzon Istanp. Abra Prov.: Mt. Posuey, Feb 1927 (fr), M. Ramos BS 20710 (A). Rizal Prov.: Jan 1913 (stam. fl), A. Lober 13871 (A), Feb 1913 (stam. fl), A. Loher 14221 (A), (stam. fl), A. Lober 14230 (A); sine loc. esp., May L907 (fr), M. Ramos BS 2040 (L). Quezon Prov.{Tabayas]: Mt. Dingalan, Aug—Sep 1916 (fr), M. Ramos & G. Edafio BS 26612 (US). MALAYSIA. SaBau {British Nortu Borneo}: Me. Kinabalu, Penibukan, 16 Jan 1933 (stam fl), J G@ M. 8. Clemens 31119 (A, BM). 8 Nov 1933 (stam. fl bud), J G M. 8. Clemens 50295 (A, BM, NY). Myrsine penthukana is a new record for the Philippines. Within the coun- try, it can be confused with M. fastigiata. However, the erose-fimbriate floral bract margins with long white glandular cilia, the much longer short shoots, corolla lobes with red punctations along the margins and pellucid punctate-lineations medially, and red punctate-lineate anther connective of M. penibukana clearly distinguish it from M. fastigiata. Myrsine pentbukana is most closely related to another Sabah species, de- scribed by Merrill as Rapanea multibracteata. However, because I recognize all species of Rapanea as members of Myrsine, the following combination is necessitated: lla. Myrsine multibracteata Oye Pipoly, cee nov. Rapanea multi- A bracteata Merrill, J. Asia Soc. Malay 1 192 a pe: MALAYSIA. Sapan [Britisu Nortu Borneo}. Sandakan, Jan 192), a a ), a ee 1345 (HOLOTYPE: BM, v.; soTyPES: A* !, K!). *The tsorype at A bears i number “Ramos 1354" instead of "Ramos 1345” as cited in the a oie However, because it is an obviously retyped label, it is most likely that this represents a true isotype. Pipoty, Synopsis of Myrsine in the Philippines 149 Myrsine multibracteata is most closely related to M. penibukana, because of the long, multibracteace short flowering shoots, the coriaceous, glandular- ciliate, and oblate floral bracts, 5-merous flowers with ovate to oblong ca- lyx lobes, elliptic to oblong corolla lobes, and the coriaceous leaves. How- ever, they may easily be distinguished by the following key: 1. Leaf buds and branchlets ferrugineous glandular-granulose; ees thinly coriaceous, nitid above, conspicuously black punctate-lineate below; calyx 1.3-1.6 mm long, the lobes 1.2—1.5 mm long, 0.5—0.7 mm ae densely selhierd to orange punctate and punctate-lineate micdiany, the margin ir- regular, entire, glandular-ciliate; corolla chartaceous, 2.8-3 mm long, the lobes cucullate, densely and prominently pellucid punctate-lineate medi- ally, prominently red punctate; plants of montane forests ............ 11. M. penibukana 1. Leaf buds and branchlets rubiginous glandular-papillate; leaf blades thickly coriaceous to cartilaginous, sordid above, densely and prominently pellucid and red punctate and punctate-lineate below; calyx 0.8—1.0 mm long, the lobes 0.6-0.8 mm long, 0.3—0.4 mm wide, sparsely pellucid aaa me- dially, the margin entire, glabrous: corolla membranaceous, 1.5—2 mm long, the lobes flat, sparsely pellucid punctate-lineate; plants of mangrove swamps M. multibracteata Ac this writing, M. mu/tibracteata has not been found in the Philippines. It is a characteristic element of the mangrove vegetation of Sabah. 12. Myrsine fastigiata Cane Pipoly, comb. nov. (Fig. 3E, H). Rapanea fastigiata Elmer, Leafl. Philipp. Bot. 2:670. 1910. Type: PHILIPPINES. MiInpANAO IsLaND. North Cotabato aa {Todaya}], Mt. Apo, 2000 m, Aug 1909 (stam. fl), A. Elmer 11447 (noLotype: PNH, destroyed; tecrorype, here designated, GH!, IsoLEcTOTYPES: A!, K, L, NY!). Tree to 8 m tall. Branchlets: angulate, 2.5—-4 mm diam., densely rufous papillose-tomentose, glabrescent. Leaves: buds densely rufous papillose to- mentose, especially on adaxial petiole surface, the bud margin densely ru- fous ciliolate; blades coriaceous, symmetric, elliptic, oblong or rarely oblan- ceolate, (3-)6—-7(—9) cm long, (1.5—)2—2.5 cm wide, apically acute to acuminate, the acumen with an obtuse tip, basally cuneate, decurrent on the petiole, nitid and smooth above, scrobiculate with obscure black punc- tations and punctate-lineations below, midrib immersed above, prominently raised below, the secondary veins 15—20, prominulous above and below (with magnification), the submarginal collecting vein ca. | mm from mar- gin, with the margin entire, revolute; petiole marginate, 5-8 mm long, almost flat above, glabrous at maturity. Staminate inflorescence: fasciculate, 4—9-flowered; peduncle 3.5—4.0 mm long, 1—2.5 mm diam.,; floral bracts chartaceous, widely ovate to suborbicular, 2.0—2.5 mm long and wide, apically broadly rounded, cucullate, densely and prominently black punc- tate and punctate-lineate medially, the margin scarious, hyaline, densely 150 Sipa 17(1) rufous glandular-ciliolate, entire; pedicels obsolete or cylindric to 2 mm long. Staminate flowers: pale green; 5—G-merous, chartaceous, 2.8—3.3 mm long; calyx cotyliform, 1-1.5 mm long, the tube 0.3—0.5 mm long, the lobes ovate to suborbicular, 0.7-1.0 mm long, 0.7—1.0 mm wide, apically rounded to obtuse, densely and prominently black punctate medially, the hyaline, erose apically, and densely glandular-ciliolate; corolla subrotate, S—3 mm long, the tube 0.8—1.0 mm long, the lobes narrowly ovate, 2— 2.2 mm long, 0.8—1.3 mm wide, apically subacuminate, glabrous, densely and prominently black punctate and punctate-lineate medially, with the margin entire, densely glandular-ciliate throughout its length; stamens 2.2— 2.5 mm long, the filaments 1.3—1.5 mm long, basally fused to the corolla tube ca. | mm, the apically free portion ca. 0.5 mm, the anther ovate, 0.8— 1 mm long, 0.3—0.5 mm wide, the apex apiculate, the base cordate, dehis- cent by introrse, wide longitudinal slits, the connective epunctate; pistillode conic, ca. 1 mm long, 0.5 mm wide, hollow, the stigma and style not dif- ferentiated. Bisexual flowers: like staminate but 2.5—3 mm long, calyx 1.2— 1.4 mm long, the tube ca. 0.2 mm long, the lobes subdeltate 1—1.2 mm long and wide; corolla 3—-3.5 mm long, the tube ca. 0.5 mm long, the lobes narrowly ovate to lanceolate, 2.5—3 mm long, 1.3—1.5 mm wide; stamens 2.2—2.5 mm long, the filaments 0.5 mm long, entirely fused to corolla tube, the anthers oblong to ovate, 1.8—-2 mm long, 0.4-0.6 mm wide; pistil subglobose, slightly costate, 1.3—1.6 mm long, 0.8—1.3 mm diam., the placenta globse, apically apiculate, ovules 3, deeply immersed in pla- centa, stigma conical, with a stylopodic base, slightly lobed. Pistil/ate flowers: like staminate but 3—3.5 mm long; calyx 1.3—1.5 mm long, the tube 0.2— 0.3 mm long, the lobes widely ovate, 1.1-1.3 mm long and wide; corolla 3—3.5 mm long, the tube 0.3-0.5 mm long, the lobes ovate, 2.5—3 mm long, 0.8—1 mm wide, apically somewhat acuminate; staminodes 2.3—2.5 mm long, the filaments 0.5 mm long, entirely fused to corolla tube, the antherodes lanceolate, 1-1.5 mm long, the apically acute, basally sagit- tate, without pollen; pistil 2.3-2.5 mm long, the ovary subglobose, 2.2— 2.3 mm long and diam., the stigma conic, with a stylopodic base, slightly 4-lobed, the placenta subglobose, the ovules 6, inserted and exposed in one series near placenta. Fruit subglobose, 2.5—3 mm long and in diam., promi- nently orange punctate and punctate-lineate. Distribution.—Endemic to the Philippines, from the Mountain Province of Luzon southward through Camarines and Leyte to Bukidnon Province of Mindanao, at 2,000—2,500 m elevation Ecology and conservation status.—Myrsine fastigiata is a conspicuous ele- ment of short, dense stands of upper montane moist forests. These few remaining forests are found on a very limited number of mountains in the country, and therefore, the species is considered endangered. I N \ Pipoty, Synopsis of Myrsine in the Philippines 151 Etymology.—The epithet ‘fastigiata’ refers to the clustered, erect branches of the tree. This is more accurately described by its architectural model, “Rauh’s Model “ (sensw Halle et al. 1978). Trees exhibiting Rauh’s Model are characterized by their rhythmically growing, readily distinguishable trunks, which develop tiers of branches morphogenetically identical to those trunks. All branches are orthotropic and monopodial, with lateral (“axil- lary”) inflorescences that do not affect shoot development. In Myrsine, like other Myrsinaceae, branching is sylleptic, easily recognized by the absence of basal bud-scales and by the elongate hypopodium. Therefore, given room to grow freely, and without experiencing reiteration phenomena, all spe- cies of Myrsine could indeed appear “fastigiate” in the sense the term is used here. Common names.—Supak” (Igorot). Because this species has heretofore been confused with Myrsine avenis, it is probable that other names, such as “Berig” (Bisayan-Panay dialect) and “Paspas” (Bukidnon-Binukid Manobo dialect) will also apply to it. Specimens examined. Luzon IsLanp. Bataan Prov.: Mt. Mariveles, upper Lamao River, 1,060 m, 23 Mar 1904 (fr), R. Williams 750 (NY, US), Mar 1905 (bud), E. Merrill BS 3954 (NY, US). Benguet Prov.: Atok Municipality, Mt. Pauai, Apr-Jun 1918 (bisex. fl), J. Santos BS 31782 (A, K, PNH, US), (pist. fl, fr), J. Santos BS 31995 (A, K, PNH, US), Mc. Paua, 2,380 m, 15 Apr 1994 (fr), L. Co 4135 (BRIT, CAHUP, PUH), “Heights in the Oaks,” 2,134 m, Jul 1907 (fr), E. A. Mearns 4270 (L); sine loc. esp. Dec 1926 (fr), M. Clemens Dee Baguio, Mar 1907 (fr), A. Elmer 8597 (A, K, a (fr), A. mee 8807 (A, K US); Mt. Santo Tomas, beyond jeep track, | Jul 1958 (pist. fl, fr), J. Sinclair G G. Bdaiio ie (L, PNH, SING); Mt. Singakalsa, Mar 1931 (fr), E. Quinmiing & M. en BS 82441 (A). Camarines del Sur Prov.: Paracale, sd. (fr), S. Vidal 1525"A” (A). Mountain Prov {Bontoc}: s.d., (fr), §. Vidal 1525 (K); Mt. Catua, Bontoc, Mar 1920 (bisex. fl), M. RintoeE G. Edattio BS 37982 (A, L, US); Mt. Pulog, 10 Mar 1961 (stam. fl), M. Stezmer 1921 (L), Mt. Pulog, 16°36'N, 120°54'E, 2,350—2,450 m, 22 Jan 1968 (pist. fl, fr), M. Jacobs 7039 (A, K, L, LBC, PNH); W slopes of Mt. Pulog, Mar 1948 (bisexual fl), M. Celestzno et al. 0-268 (PNH 4330) (A); Mt. Pulog, Jan 1909 (bud), H. Curran et al. FB 18038 (US), (pist. fl), H. Curran et al. FB 18139 (BISH); Mt. Nafigaoto, Km 62 below rod, 2,500 m, 3 Feb 1948 (stam. fl), M. Sulit 2343 (PNH 7687) (A); 2,400 m, 24 Jan 1948 (stam. fl), M. Sulit 2197 (PNH 7485), (A, L); Mt. Tabayoc, 16°42'N, 120°53'E, 2,240 m, 18 Feb 1968 (pist. fl, fr), M. Jacobs 7532 (A, K, L, LBC PNH), 2,000—2,100 m, 19 Feb 1968 (stam. fl), M. Jacobs 7564 (A, K, L, LBC, PNH). Ifugao Prov.: Mt. Polis, Feb 1913 (stam. fl), R. M° Gregor BS 19646, (K, L, PNH, US), BS 19833 (K, L, PNH, US), Mt. Polis, 16°59.2'N, 121°02.4'E, 1,880 m, 18 Apr 1993 (ster.), L. Co 3812 (A, CAHUP, CANB, K, L, PUH, US). Zambales Prov.: Santa Cruz, 8. Vidal 1517 (A, L). Leyte IstANp. Leyte Prov.: Mt. Suiro, N slope, 900 m, 5 May 1954 (fr), M. Sait 5450 [PNH 21617 (L, PNH). MINDANAO IsLAND. Bukidnon Prov.: Mt. Lipa, Jun—Jul 1920 (fr), M. Ramos G G. Edano BS 38494 (K, PNH, US). Myrsine fastigiata was placed into synonymy under Myrsine avenis by Merrill (1923), but can be separated from it by the tomentum of the branchlets (rufous papillose-tomentose vs. rufous tomentose with rufous sessile scales), longer petioles, subsessile, fasciculate inflorescences, and much 152 Sipa 17(1) shorter pedicels. Its habitat is similar to that of Myrsine avenis but label data imply that M. fastigiata occurs in more sheltered thickets of those forests. 13. Myrsine Serious weve eee cae nov. (Fig. 3A, B). o aralioides Philipson, ee J. Bot. 78:104. 1939. Type: MALAYSIA. BAH {BRITISH NORTH BORNEO}. Mount oo lu, aie: at head : - di- vide of Dahombong Creek, at 1,52 — ; : see 1933 (pise. fl, fr), J. & M. S. Clemens 40664 (HOLOTYPE: BM, n.v.; IsOTYPES: , LI, NY!). Tree to 7 m tall. Branchlets: terete, 7-10 mm diam., drying sulcate, sparsely covered with minute translucent or rubiginous glandular lepidote scales, glabrescent. Leaves: buds densely covered by minute translucent or rubiginous glandular lepidote scales, the margin sparsely rubiginous glan- dular ciliate basally, glabrescent; blades cartilaginous to thickly coriaceous, symmetric, elliptic to oblanceolate or obovate, 8.5—15(—21) cm long, 3.5— >(—8.5) cm wide, apically obtuse, rounded or acutish to an obtuse tip, ba- sally rounded and often abruptly tapering, decurrent on the petiole to base, prominently punctate and scrobiculate and subnitid above, densely and prominently red punctate and black lineate-punctate below, the midrib prominently raised above and below, the secondary veins 20—25 pairs, the submarginal collecting vein inconspicuous, ca. 1.5 mm from margin, the margin entire, revolute; petiole amply marginate and canaliculate, 5—10 mm long, glabrous at maturity. Staminate eae unknown. Bisexiwal inflorescence: a 6—9-flowered umbel: ; peduncle 3.5-5 mm long, (3—)3.5—4 mm diam.; floral bracts coriaceous, oblate, 1.2— ‘ > mm long, 1.5—-2 mm wide, apically acute, cucullate, inconspicuously pellucid punctate, the margin entire, densely villous-glandular-ciliate; pedicels cylindrical, 2.6— 4 mm long. Bisexual flowers: 5-merous, 3.3—3.5 mm long; calyx cotyliform, 1.5—2 mm long, the tube to 0.5 mm, the lobes deltate, 1—1.5 mm long, 1- 1.4 mm wide, apically acute, medially transversely rugose, densely and prominently black punctate marginally and black punctate-lineate medi- ally, the effect rendering the lobe verruculose, the margin entire, glandu- -3—3.5 mm long, the tube ca. 0.2 mm long, the lobes > lar-ciliolate; corolla 3 elliptic, 3.1-3.3 mm long, 1—1.2 mm wide, apically acute, densely black punctate-lineate apically and toward the margin, densely glandular- granulose apically within and on the margin eitself without; stamens 2.5— 2.7 mm long, the filament ca. 0.2 mm long, the anther lanceolate, 2.3—2.5 mm long, ca. | mm wide, apically acute, basally deeply cordate, the con- nective sparsely and minutely but prominently black punctate; pistil obconic, 1—1.4 mm long, the ovary ca. 1 mm long, the style and sigma morchelliform, ca. 0.3 mm long, the ovules 2—3, uniseriate. Fruit globose, 3-5 mm long and in diam., densely and prominently red punctate. Pisti/- late inflorescence: a 12—15-flowered umbel; peduncle a “short shoot” 8—10 Pipoty, Synopsis of Myrsine in the Philippines 153 mm long, 2-3 mm diam.; pedicels 2.6-4.0 mm long; floral bracts chartaceous, ovate, 1.5—1.7 mm long, 0.6-0.8 mm wide, apically acute, densely and prominently punctate medially, the margin entire, densely glan- dular-ciliate. Prsti/late flowers: S—6-merous, like bisexual but calyx 1.3-1.5 mm long, the tube ca. 0.2 mm long, the lobes ovate, 1.1—1.3 mm long, 0.6—-0.8 mm wide; corolla 2.2—-2.5 mm long, the tube 0.3 mm long, the lobes 2.1—2.2 mm long; staminodes resembling stamens but reduced in size, 1.2—1.4 mm long, the filaments entirely adnate to the corolla tube, ca. 0.3 mm long, the apical free portion ca. 0.1 mm long, the antherodes 0.6— 0.8 mm long, 0.3—0.4 mm wide, apically acute, devoid of pollen; pistil conic, 2—2.2 mm long, the ovary ee 1—1.2 mm long, 0.8—1 mm diam., the stigma translucent, carnose, 4-lobed, the placenta depressed-globose, ovules 4—G6, immersed in the arn Pruit like bisexual fruit. Distribution.—In the Philippines, Myrsine aralioides is known only from the Island of Mindanao (Surigao del Norte Province) and the Island of Pala- wan, Palawan Province, at 150-200 m elevation. Outside the Philippines, the species is well known from Marai Parai and Penibukan, of the Kinabalu Mountain Chain in northern Sabah, Malaysia, at 1,524 m elevation. Ecology and conservation status.—Myrsine aralivoides is an exceedingly rare taxon, especially in the Philippines, and as such should be considered en- dangered. Given its distribution, it may be an ultramafic specialist. Unfor- tunately, because it is known from mostly historical collections, with few modern data, its ecology is in further need of study. Etymology.—The specific epithet refers to the ramigerous and ramuligerous umbels, especially conspicuous when in fruit, which are reminiscent of Schefflera (Araliaceae) species when viewed from afar. Common names.—“Lanito” (Bisayan-Davao del Sur). PINES. MINDANAO IsLAND. Davao Oriental Prov.: South eee Mati [ca. ee he oe 13'E}, along Biga Creek, 8 May 1929 (fr), L. Aguilar 53 ume 31044} (NY). Surigao [del Sur ?} Prov. Wee 1919 (fr), M. Ramos & J. Pasgasio BS 34674 (A). PALAWAN ISLAND. Palawan Prov.: Mt. Bloomfield Sabang, Aug—Sep 1986 (fr) rE. Reynoso & J. Proctor et al. 489 (BISH, Ls PNH): Mt. Bloomfield, WNW to W face, near chromite mine, {ca. 10°12'N, 118°52'E], 150-200 m, 6 May 1984 (fr), C. Ridsdale et al. SMHI 1623 (L, PNH). MALAYSIA. SaBau [British Norty Borneo}: Ranau District: along E Mesilau River between Mt. aN golf course site and eee Cave 06°03'N, 116°36'W. 1,700—1,900 m, 20 Dec 1983 (f ), J. Beaman 7992 (GH, K, L, MSC, UKMS); Pig Hill, 2,134m, 18 Feb 1964 i (pis. fl, fr), W. Chew & E. Corner RSBN 4370 (K, L); npper Kinabalu, 1,829—4,115 ae 1931 (fr), J. G M. Clemens 27801 (K, L, NY), 3,350 m, Dec 1933 (stam. Ae G M. Clemens 5O807 (NY); Penibukan, 1 219 1,524 m, i 1933 (fr), J & M. Clemens ey (A, L, NY); Ranau District, Marai Parai, 1,524 m, 30 Mar 1933 (fr), J. & M. Clemens 32346 (A, L, NY), 10 Apr 1933 (fr), J. & M. Clemens 32612 (A, L, NY), 1,524—-1,829 m, 17 Sep 1965 (fr), A. Kanis 53996 (L, SAN); Small valley N of Carson's Camp, 2,700— 2 ,900 m, 15 Jan 1989 (ster.), M. Hotta 3899 (L). In the Philippines, Myrszne aralioides can be most often confused with 154 Stipa 17(1) M. glandulosa, but is easily distinguished by its thick, sulcate branchlets, the branchlet and leaf bud vestiture of minute translucent or rubiginous lepidote scales, the marginate and canaliculate petioles, larger thickly co- riaceous to cartilaginous leaf blades, and the deltate calyx lobes. There may be an as yet undescribed, more closely related taxon from upper montane forest on the Selangor/Pahang Genting Highlands access road in Malaya, which differs primarliy in having longer petioles, much longer, angulate pedicels, and larger fruits. 14. Myrsine glandulosa (Elmer) Pipoly, comb. nov. (Fig. 3C, D). Rapanea glandulosa Elmer, Leafl. Philipp. Bot. 8:2784. 1915. ee PHILIPPINES. MINDANAO IsLAND. Agusan Prov.: Cabadbaran (Mt. Urdaneta), | 4m., Oct 1912 (fr), A. E/mer 14148 (HOLOTYPE: PNH, destroyed; LECTOTYPE, nga geen A!; ISOLECTOTYPES: BISH!, GH!, L!, K! NY! —_— Epiphytic shrub to tree to 3.5 m tall. Branchlets longitudinally ridged to angulate, 3—3.5 mm Ae densely rufous glandular stellate tomentose at first, early glabrescent. Leaves: buds densely appressed rufous glandular stellate and dendroid-tomentose, the margins long translucent ciliate, gla- brescent; blades coriaceous, obovate, oblong or oblanceolate, (4—)S—7(-9.5) cm long, (1.6—-)2—2.8 cm wide, the apex obtuse or rarely, subacuminate and ending in an obtuse tip, the base cuneate, decurrent on the petiole for 3/4 its length, nitid, densely and prominently translucent punctate above and be- low, midrib raised for its entire length above to petiole base, prominently raised below, the secondary veins 6—9(—15) pairs, somewhat prominulous above and below, the submarginal collecting vein ca. | mm from margin, the margin entire, revolute, with numerous prominent pellucid puncta- tions along its length below; petiole marginate, the midrib prominently raised above, 3—5 mm long, glabrous. Stamznate seelas: unknown. Pis- tillate inflorescence: umbellate, 3—12-flowered; peduncle 2—6 mm long, 1-2 mm diam.,; pedicels angulate, 3-6 mm long, densely and i ecomninesely black punctate-lineate; floral bracts chartaceous, widely oblate, 0.3—0.5 mm long, ().8—1.1 mm wide, apex broadly rounded, densely and prominently brown punctate, the margin minutely glandular ciliolate. Prsti//ate flowers: un- known. Fruiting calyx 4-merous, chartaceous, unequally divided, 0.6— mm long, the tube ca. 0.1 mm long, the lobes very widely ovate, 0.5—0.7 mm long, 0.6—8 mm wide, apically obtuse, sparingly but prominently black punctate, the margin irregular, entire except minutely erose apically, gla- brous. Fruit globose, 3.5—4.5 mm long and in diam., densely and promi- nently black punctate, the remnant stigma base stylopodic. Distribution.—Myrsine glandulosa is apparently endemic to the Philip- pines, thus far known only from southern Mindanao and nearby Negros Oriental, at 1,350—2,242 m elevation. — PrpoLy, Synopsis of Myrsine in the Philippines 155 Ecology and conservation status.—This species is poorly known, and has been found only in submontane forest. Etymology.—The specific epithet refers to the extremely prominent pel- lucid punctattions of the adaxial and abaxial surfaces of the leaf blades. Common name.— Nagas” (Magobo language, Madulid 1992). Sp xamined. PHILIPPINES. NeGros ee Negros Occidental Prov.: SW of Mec. Canlaon, sue Guintubdan Forest Hut, 1,350 m, 29 Mar 1992 (fr), B. Sas et a Pi (A, BO, BRIT, K, L, PNH, US). Ne ae ee Prov.: Dumaguete, Cuern , 1,680 m, May cn (fp), A. Elmer se (A, K, NY, PNH, US), (fr), A. E/mer 10186 a % L, PNH, US). Leyte Istanp. Leyte Prov.: sine esp., 5 Jun 1914 (fr), C. A. Wenzel hig (A), 15 Aug 1914 (fr), C. A. Wenzel ae (A, GH). MINDANAO ISLAND. Bukid ov.: Kaatoan, Malaybalay, 11 Oct 1953 (stam. Al), B.B. Britton 441 {PNH 19666} (L, ae sine loco esp, Jul-Aug 1914 (fr), L. Escritor BS 21404 (L, NY). Davao del Sur Prov.: Davao, Aug 1933, R. Kanehira 2672 (NY); Mt. McKinley, 2,000 m, 24 Aug 1946 (ster.), G. Edato 757 {PNH 1013} (A, L, PNH) 7,400 ft, 13 Sep 1946 (ster), G. Edaiio 987 {PNH 976} (A, L, PNH), 1002 {PNH 1052} (A, K, L), (pist. fl, fr), 999 (PNH 979} (A); 995 {PNH 994} (pist. fl, fr) 895 (PNH 987}, 2466 m, 25 Sep 1946 (fr), G. Edafio 1059 {PNH 1568} (A). South Cotabato Prov.: Mt. Matutum, ee 1932 (ster.), M. Ramos & G. Edatio o 55015 (A), Mt. Matutum 2,242 m, 15 Apr 1992 (fr), EF Gaerlan et al, PPI 5361 (A, BO, BRIT, K, L, PNH); Klaja Hill, Barrio Konel, 7 Sep 1950 (pist. bud), P. Avtonuevo 297 ee 13598} (A, L, PNH). Myrsine glandulosa may easily be confused with M. aralioides in the Phil- ippines, but differs by the thinner branchlets, less coriaceous leaves, promi- nent leaf punctations, red-papillose tomentum of the leaf buds, raised leaf midrib decurrent to the petiole base, longer petioles, thin peduncle and smaller sepals. ACKNOWLEDGMENTS I owe my sincerest appreciation to Sy Sohmer and Domingo Madulid, co-Pl’s of the Philippine Plant Inventory Project at BRIT and PNH te- spectively, for exposing me to Philippine botany. The company, hospitality and teamwork of the PPI field team members, Ernesto “Mang Estong” Reynoso, Toy Sagcal, Raymond Fuentes, Humberto Garcia, Efren Romero, “Ding” Fernando, Ferdie Gaerlan, Ernesto Barbon, and Raymond Garcia were much appreciated. The special collaboration and collegiality of Victor and Cecilia Amoroso (CMU) was invaluable, as was the hospitality of the Talandig Tribe from Sungco, Bukidnon and Mr. John Tonsciongco of Magdiwang, Sibuyan Island. Important insights into the distribution of ultramafic formations was obtained through the courtesy of the staff of the Geology Division (Luis T. Omafia, Roberto SP de Ocampo, and Priscilla Ong), Philippine National Museum. Technical, administrative and other assistance from June Cunningham, Melanie Medecilo, Maricar Sampson, Irma Castro, Maribel Agoo, “Magen” Genevieve Lusica, James Rivers, and 156 Stipa 17(1) Myrna Resurecci6n, was very much appreciated. My collections work at PNH and attendance at the Flora Malesiana Symposium and visit to K were supported by the John D. and Catherine T. MacArthur Foundation, while fieldwork and work at BRIT is supported by the Philippine Plant Inventory, NSF Grant DEB 9300910. While at K, I enjoyed the cordial hospitality of its director, Prof. Sir Ghillean Prance, as well as the privilege of working and collaborating with Ynette, “Tivvy” Harvey. Also at K, Nicholas Hind, John H. Beaman, Robert Johns, David Frodin, Dick Brummit, “Lulu” Rico, Diane Bridson, Mark Coode, Martin Cheek and Barry Conn, gave freely of advice in geographic specialties, thus making my trip even more productive. During my visit to US, the cordiality and hospitality of Warren Wagner, Vicki Funk, Carol Kelloff, and Rusty Russell was immensely helpful. At A/GH, Michael Donoghue, Emily Wood, Dave Boufford, Anthony Brach, Peter Stevens, Paul Goff, Walter Kittridge, Benito Tan, Peter Ashton, James La Frankie, Santiago Madrinan and Stuart Davies greatly facilitated my work with their collaboration and hospitality. At NY, Scott Mori, Carol Gracie, Pat Holmgren, Brian Boom, Sara Hunkins, Pam White, Lisa Campbell, Amy Litt, Steve Churchill, Favio Gonzalez and Jackie Kallunki all helped make my visit a success. In the Philippines, the curators and staffs of CAHUP (Norma Aguilar, E. Mandia), CLP Justo Rojo), LBC (Edwino Fernando), and PUH (Daniel Zagunzal, Leonard Co), received me with warm hospitality and collaboration. I also thank the cu- rators of cited herbaria for providing loans of specimens. Finally, I am ae to Gordon McPherson (MO), Kancheepuram Gandhi (GH) and Barney Lipscomb (BRIT), whose thought-provoking re- views of the manuscript and helpful suggestions greatly improved the quality and clarity of the paper. REFERENCES BACKER, . and R.C. BAKHUIZEN VAN DEN BRINK, JR. 1965. Myrsinaceae. Fl. Java 2: 194-2 BATHIE, . ae RRE DE LA. 1953. 161° Famillie Mysinacées. In. H. Humbert, ed. Flore de Madagascar. Typographie Firmin-Didot. Paris. Pp. 1-148. CANDOLLE, ALPHONSE DE. 1834a. A review of the natura Penden. 12:95-138. order Myrsineae. Trans. Linn. Soc. «18345. Revue de la famille des Myrsinées. Ann. Sci. Nat., Bot., Ser. 2, 9:285—301 184 1a. Second mémoire sur la famille des Myrsineacées. Ann, Sci. Nat., Bot., Ser 2 Sone 9Q7, | ). 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Flore ilustrée des See ahi de Guadeloupe et de Martinique. Institut National de la Reserche Agronomique. Par GREEN, P. 1986. Notes relating to the floras of ‘Norfolk Island and Lord Howe Islands, II. J. Arnold mee 67:109-122. eee oie relating to the floras of Norfolk Island and Lord Howe Islands, III. Kew Bull. yer 235-255. GuzMAN-TEARE, M. 1992. Wns In: L. Brako and J. Zarucchi, ed. Catalogue of the oo Plants and Gymnosperms of Peru. Missouri Botanical Garden. St. Louis. Pp. 30- Hire r, R OLDEMAN and P. ToMLINSON. 1978. Tropical forests: An architectural analy- sis. Springer-Verlag. Berlin. Hatuipay, P. 1984. eee 7 - Polhill, ed. Flora of Tropical East Africa. A. Balkema. Rotterdam. Netherlands. Pp. 1— Harvey, Y. and J. PIpoty. tc a ee In: B. Stannard, ed. Flora of the Pico das Almas, Chapada Diamantina, Bahia, Brazil. Royal Botanic Gardens, Kew, Richomond, U.K. Pp. 487-491. Hosaka, E. 1940. A revision of the Hawaiian species of Myrsine (Suttonia, Rapanea) (Myrsinaceae). Occ. Pap. Bishop Mus. 16:25—76. Howarp, R. 1989. Myrsinaceae. In: R.A. Howard, flora of the Lesser Antilles. Eaconyle dons, Vol. 6. Arnold Arboretum of Harvard University, Jamaica Plains. Pp. 34-53. Koorpers, S. and T. VALETON. 1900. Myrsinaceae. In: S$. Koorders and T. Valenton, ed. oomsoorten op Java; Bijdrage 5. G. Kolff & Company. Netherlands. Pp. 215-278. Kupicua, F. 1983. Myrsinaceae. In: E. oe ed. Flora Zambesiaca. Flora Zambesiaca Managing Committee, London. Pp. 198— Li, H.-L. 1963. wee flora of Taiwan. Mors Arboretum, Philadelphia. «978. Myrsinaceae. In: H. T-S Liu, T-C Hu oe T. Koyama and C. DeVol, ed. Fides of Twas, Epoch ee Taipei. Pp. 47- LioGier, ALAIN [HERMANO ALAIN}. 1959. Myrsinaceae. In: H. 0s FlomdetCube Vel: e La Salle, Habana, Cuba. Pp. 106-113. . Novitates antillinae. Myrséve in Hispaniola. Brittonia 21:140-144. _ «1: 989. Myrsinaceae. In: In: H. Le(n. La flora de La Espafiola. V. Universidad Central del ‘Este Santo Domingo, Dominican Republic. Pp. 34—53 LITTLE, ., R. Woopsury and F. Wapswortu. 1988. Arboles de Puerto Rico y las Islas Virgene s, U.S.D.A. Puerto Rico. LUNDELL, C.L. 1966. Flora of Guatemala: Myrsinaceae. Fieldiana, Bot. 24(8):1—35. «971. Flora of Panama: Myrsinaceae. Ann. Missouri Bot. Gard. 58 285-353. 158 Sipa 17(1) 1981. Neotropical Myrsinaceae- IV. Phytologia 48:137-142. —_____. 1984. Neotropical Myrsinaceae- XV. Phytologia 56:413—418. MADULID, = 1992, A dictionary of Philippine plant names. National Museum, Manila. Merrill, E. 1923. Myrsinaceae. In: E. Merrill. Enumeration of Philippine plants. Bureau of Prining, Manila. Pp.253-274. Meyer, F. and E. Waker, eds. 1965. Flora os by J. Ohwi, translated and updated. eae a Institution Press, Washington, DC Mez, C. 190 eee : J. Urban, ed. oe Antillinae 2:397—500. 1902. Moiese In: A. Engler, ed. Das Pflanzenreich 9(1V. 236): 1— Nicuotson, A. 1991. Myrsinaceae. In: D. Nicholson, ed. Flora Dominica. Part 2: Dicotyledonae. Smithsonian Institution Press. Washington, DC. Pp. 159-161 Otiver, W. 1951. The flora of the Three Kings Islands: Additional notes: with notes on Suttonia. Auckland Inset. Mus. 4:111—112. Pipoty, J. 1981. Contributions toward a monograph of Cybranthus ey ee I. Subge- nus /teordes and the identity of Conomorpha loretensis. Brittonia 91-493 —______. 1983a. Contribucions toward a monogr sas of Cybianthus (Myrsinaceae): II. A revision id subgenus oe Brittonia 35:61—80 —______.. 1983b. Contributions toward a monograph of Cybianthus (Myrsinaceae): IV. Notes on ae amir and Stapha. Wrightia 7:2 4, —___.. 1987. A systematic revision of the genus Cy ee ‘ (Myrsinaceae). Mem. New York Bot. Gard. 43:1—76. —_______. 1988. Contribuciones para una monografia del género Gea (Myrsinaceae): V. Dos especies nuevas de la Guayana venezolana. Ernstia 50:32—40 —_______.. 199 la. Systematic studies in the genus Myrsine L. (Myrsinaceae) in cite Novon 1:204—210. L991b. New species of Cybtanthus subgenus Conomorpha (Myrsinaceae) from Amazonian Peru. —— ea 46% il— Loot ~ a c.Nt | s del gé se ee a Martius subgénero Conomorpha (Myrsinaceae) de Colombia. Clee 16(78):2 64 1992a. A further note on Myrsin 7 iuciflora Pipoly (Myrsinaceae). Novon 2:176. . 1992b. Estudios en el género Myrsive (Myrsinaceae) de Colombia. Caldasia 1992c. Notes on the genus Myrsime (Myrsinaceae) in Peru. Novon 2: —_______.. 1992d. The genus Cybranthus subgenus Conomorpha (Myrsinaceae) en Guayana. Ann. oo Bot. Gard. 79:908-957. ee . Notes on a subgenus Cybtanthus (Myrsinaceae) in south- eastern Brazil. ae 3:459—-4 2, 1994, Noteson te genus ¢ aia subgenus Cyhianthus (My iimatene) in Colombian ‘Agaczoma Sida 16:333-3 and C. Chen. Nome nc: itural notes on the Myrsinaceae of China. Novon 5:357-361. RicHarp, M., R. Maury, H. Betton, J-F. SrepHan, J-M. Borrat and A. CALDERON. 1986. Geology 3 Mt. Iraya Volcano and Batan Island, northern Philippines. Philipp. J. Volcan. 3:1-27. Rock, J. 1974. The indigenous trees of the Hawaiian Islands. Pacific Tropical Garden and OES E. Tuttle Company. Hawaii. SACHET, ens Flora of the Marquesas, |: Ericaceae-Convolvulaceae. Smithsonian Contr. Bot 3: PIpoLy, Synopsis of Myrsine in the Philippines 159 SLEUMER, H. 1986. A revision of the genus Rapanea Aublet (Myrsinaceae) in New Guinea. Blumea 31:245—2 SmitH, A.C. 1973. Studies Pacific Island plants X XV. Myrsinaceae of the Fijian Region. J. Arnold Arbor. 54:1— —_C*d'*O*'B LL. oe In: A.C. es ae Vitensis Nova, Vol. 2. Pacific Tropical Garden: Lawai, Kauai, Hawaii. Pp. 78 ene P. 1938. Myrsinaceae. = Flora of C eae a Field Mus. Nat. Hist., Bot. Ser 18(2):898. 1938. Pp. 898— STEARN, W.T. 1969. A synopsis ee Myrsinaceae. Bull. Brit. Mus. Nat Hise. Bot. Ser. 18:884—-900. Taron, A. 1980. Myrsinaceae. In: P. aes ed. Flore D’ Afrique Centrale. Jardin Botanique National de Belgique. Meise. Pp. 1— WacGner, W.H., D. Hersst eae 6. ScuuER 1990. Manual of the flowering plants of Hawai'i. University i: Hawaii Press and Bishop Museum Press. Honolulu Watker, E.H. 1940. A revision of the eastern Asiatic Myrsinaceae. Philieg, J. Sci. 73: 1- 258 _—«*d'S. AA revision of the Myrsinaceae of Taiwan. Quart. J. Taiwan Mus. 12:161-194. Wirpur, R.L. 1965. Nomenclatural notes on Hawaiian Myrsinaceae. Pacific Sci. 19:522. NUMERICAL LIST OF TAXA 1. Myrsine medeciloae Pipoly 9, Myrsine philippinensis A. DC. 2. Myrsine apoensis ee Pipoly 10. Myrsine avenis (Blume) A. DC. 3. Myrsine amorosoana Pipoly 11. Myrsine penibukana (Philipson) 4. Myrsine o sblonigibaces (Merrill) Pipoly Pipoly 5. Myrsine cruciata ere Pipoly lla. Myrsine multibracteata (Merrill) 6. Myrsine densiflora Scheffe Pipo 7. Myrsine peregrina (Mez) Pipoly 12. Myrsine fastigiata (Elmer) Pipoly 8. Myrsine mindanaensis (Elmer) 13. Myrsine aralioides (Philipson) Pipoly ipoly 4, Myrsine glandulosa (Elmer) Pipoly LIST OF EXSICCATAE The figures in parentheses refer to the numbers from the numerical list of taxa. Collection numbers in boldface type indicate type specimens. Collection numbers before [ } brackets indicate field collection numbers; numbers in brackets indicate herbarium sheet numbers. Both are given because it appears that some duplicates have one number or the other, or oth on the mounted specimen. BS indicates herbarium numbers in the Philippine Bureau of Science series, FB indicates the herbarium numbers in the Forestry Bureau Series, and PNH indicates herbarium numbers in the Philippine National Herbarium Series. Other letters indicate herbarium acronyms, except for SMHI, which is a series from the Hilleshog Expedi- tion to Palawan, a joint project of K, L, and PNH, and PPI, a series from the Philippine Plant Inventory, a joint project co-sponsored by BRIT and H 160 Agaceta, E. 6 [LBC 917} (9) Aguilar, L. 33 [PNH 31044} (13) Ahern, G. FB 444 (6) Afnonuevo, P. 297 [PNH 13598} (14) Argent, G. & E. ane Bes 89112 (3); 89113 Barbon, E. et al. PPI 22 Beaman, J. 7992 (13) Britton, B.B. PNH 19666 (14); 421 {[PNH 19646} (7); 441 [PNH Celestino, M. et al. 0-268 [PNH 4330] (12) Chew, W. & E. Corner RSNB 4370; RSNB 4 Clemens, J. & M.; 20764 (11); 27801 (13); bao: a ); RSNB 4493 (5) 11); 32346 (13); 3237 Clemens, M. 917 (7); 9332 (9); 17218 (12);51876 (1( Co, L. 3073 (9); 3125 (8); 3649 (2); 3812 (12); ne Canciin. H. 529 [PNH peace (10); 864 [PNH 19321] ( Conklin, H. & Buwaya I-1023 [PNH 79612} (10) Coode, M. et al. 5635 (3); 5765 (3) Cuming, H. 868 (8);1601 (9) Curran, H. & M. Merritt BS 8072 (1) Curran, H. et al. FB 18139 (12) Dransfield, J. SMHI 1292 (5) Edanfio, G. 192 [PN [PNH 1032} (2); 895 [PNH 987} (14); 984 [PNH 1048] (2); 995 [PNH 994] (14); 999 [PNH 979] (14); 1 1564} (7); 1059 [PNH 1568] ¢ 1232 [PNH 141 1557} (7); 1535 [PNH 1553} (7); 1536 [PNH 1540 — ar BS 29386 (4); BS 76198 (3); BS 76268 (3); BS 77034 (4); BS 77079 (3); BS 7); PNH 1553 (7); PNH (4); BS 78412 (4); BS 78424 (4); BS 78533 (3); PNH 1540 ( 11565 (7); PNH 21960 (3) Elmer, A. S811 (10); 6539 (10); 8597 ( 11820 (8); 12295 (9); Escritor, L. BS 21404 (14) Fernando, E. 1003 (6) Forestry School, UP Los Bafios FB 20145 (10 saerlan, F et al. PPI 3476 (6); PPI 5361 Gammill, J. FB 272 (9); FB 276 (9); FB 301 Garcia, C. 25507 (10) Gates, F. 5196 (10) Gideon, O. LAE 76035 (G6) Haenke, T. 440 (3); 447 (3); 472 G) Hartley, T. 11955 (6) Hatusima, S. & M. Sato 29059 {[PNH 111132] (6) Hernaez, B. 867 Stee 58864, 58865, 58866} (10); 20. 60563, 60564] (3): 3 (3) Hernaez, B. & N 32378 (5); 32612 (13); 40664 t (1 3% 5( )295 (11); 50807 ¢ (12);8745 (1 a 8807 (12); 9351 (10); 4); 10186 (14); 10629 (2); 11332 (8); 11445 (7); 11447 (12); 1 14148 (1 4); 14286 (10); 17815 (10); 18270 (10) Sipa 17(1) a) 57 (9); PPI 6094 (6); ee 8223 (3); PPI 8601 (8); PPI 8845 (3) 19666} (14) 1 (5); 30258 (5); 30989 (13);31119 13) H 3574] (10); 630 [BS 76225] (3); 757 [PNH 1013} (14); 821 ); 987 [PNH 976] (14); 002 [PNH 1052] (14); rat eae 14); 1090 [PNH 1595} (7); 1095 [PNH 1 2}; 1266 [PNH 1382) (2) 1267 [PNH 1383] (2); (7); 8195 [PNH 22003] 5 78357 9352 (10): 1456 (7); (7); PPI 5361 (14); PPI 10949 (7) 38 [CAHUP 60561, 60562 {, Ci ino a AHUP 57282 (3); CAHUP 57283 (3); CAHUP 57284 (3); CAHUP 60019 (10); CAHUP 60020 (10); CAHUP 60892 (8); CAHUP 60893 (8); CAHUP 60894 (8) Hosakawa, T. 8560 (7) Pipoty, Synopsis of Myrsine in the Philippines 161 Hotta, M. 3899 (13) Iwatsuki, K. 719 (1 Jacoban, S. 20 [LBC 966] (9) Jacobs, M. 7039 [PNH 103962} (12); 7227 [PNH 103861] (3); 7274 [PNH 103905] (3); 7333 [PNH 103962] (3); 7488 [PNH 104120} (3); 7532 (12); 7564 (12) Kaneira, R. s.n. (8) Kanis, A. 53996 (13) Kerenga, K. et al. LAE 77490 (6) Kondo, Y. & G. Edafio 113 [PNH 36799, PNH 36817] (9) Kostermans, A. & Soegeng 311 (6) cae D. 758 (8); 870 (7) 916 (8); 961 (7) Lam, H. 3258 (6); 3373 (6); 3420 (6) Lit, I. - 84-001 [C AHUP 40296} (3) Loher, A. 3830 (10); 6125 (10); 6128 (3), 6140 (10); 12009 (10); 12038 (10); 13871 (11);14097 (10); 14221 (11); 14230 . 1); 18270 (10) Mandia, E. 421 (3); Mandia, E. 555 Martelino, A. & G. Edano BS 35612 i BS 35682 (7) M‘Gregor, R BS 49833 (7); BS 19646 (12): BS 19833 (12) Mearns, M. & W. eee FB 4573 (7) Mendoza, D. PNH 913 Merrill, E. 1206 (9); ve 5734 (3); 5735 (3); BS 6998 (10) Merritt, M. BS 4449 (3); FB 14153 (10) Meyer, R. BS 3114 (1); 3114 (10) Millar, A. & J. Vandenberg NGF 40880 (6) Nedi, X 665 (6) Orantia, L. LBC 3481 (9) Orlido, N. CAHUP 17927 (10) Pancho, J. 2587 [PNH 34613] (7) Piper, C. BS 237 (6) Pipoly, J. et al. 19551 (3); 19555 (3); PPI 16584 (7); PPI 16610 (3); PPI 16647 (3); PPI 16684 (3); PPI 16871 (3) Podzorski, A. SMHI 2125 {[PNH 159319] (5); SMHI 2130 [PNH 158938} 6) Quisumbing, E. & M. Sulit BS 82441 (12) Ramos, M. s.n. (6); 1345 (11a); BS 1524 (3); BS 1525 (3); BS 1961 (3); BS 2640 (11); BS 2650 (10); BS 5050 (5); BS 7623 (5); BS 19576 (10); BS 20710 (11); BS 22009 (3); BS 23512 (3); BS 23616 (3); BS 32757 (1); BS 33243 (1); BS 33256 (4); BS 33311 (4); BS 41559 (8); BS 77079 (4); BS 80268 (7) Ramos, M. & G. Edafio BS 26397 (7); BS 26418 (4); BS 26548 (4); BS 26577 (3); BS 26612 (11); BS 28758 (3); BS 29282 (10); BS 29386 (3); BS 34625 (6); BS 37982 (12); BS 38494 (12); BS 38528 (3); BS 38918 (3); BS 40394 (10): ); BS 40712 (7); BS 44936 (3); BS 45602 (3); BS 85015 (14); BS 85048 (7) Ramos, M. & J. Pasgasio BS 34674 (13); BS 34755 (8); BS 34755 (6); BS 35194 (4) Re », E. et al. 489 (13); PPI 1087 (6) ee e, C. SMHI 225 [PNH 157145} (5); SMHI 1623 (13); SMHI 1732 AG); SMHI 1741 [PNH 159486] (5); 1084 [PNH 162204] (6) Ridsdale, C. et al. ay (6); 1703 (3) Robbins, R. s.1 Robinson, C. “ et 14088 (10) Rodulfa, E. EVR—7 [LBC 3792} (9) Salvoza, F. 1057 [PNH 3715} (9) 162 Sipa 17(1) Santos, J. BS 31782 (12); BS 31995 (12) Sinclair, J. & G. Edafio 9800 (12) Steiner, M. 1921 (12) Stern, W. & P. Rojo 2299 [PNH 92363] (7) Stone, B. et al. PP] 6471 (10); PPI 6643 (14); PPI 6906 (3) Sulit, M. 2197 [PNH 7485} (12); 2343 [PNH 7687] (12); 3383 [PNH 10054] (7); 3467 [PNH 10125] (7); 3630 [PNH 11769] (9); 5450 [PNH 21617] (12); BS 30073 (10); PNH 11800 (9) Teysmann, J. 7471 (6) Univ. San Carlos 586 (2) VWidal,.$.:1516 ©); 1516. bis (10) 15.17 (12921525 (12) 1525 -A 12): 1526 (10); 3156 (9); 3183 (9) Walker, E. 7546 (10) Warburg, O. 21830 (6) Wenzel, C. A. 945 (14); 1063 (14); 1368 (9) Williams, R. 750 (12); 1346 (10); 2603 (10) Zippelius, A. s.n. (6) Zollinger, H. 883 (10) — MORPHOLOGY OF PENNISETUM ORIENTALE (POACEAE: PANICEAE) J. RAMU Department of Soil and Crop Sciences Texas AGM University, College Station, TX 77843, U.S.A. S.L. HATCH! Department of Rangeland Ecology and Management Texas AGM University, College Station, TX 77843, U.S.A. M.A. HUSSEY Department of Soil and Crop Sciences Texas AGM University, College Station, TX 77843, U.S.A. E.C. BASHAW USDA-ARS Southern Crops Research Laboratory Texas AGM University, College Station, TX 77843, U.S.A. ABSTRACT Pennisetum orientale Rich., a polymorphic species, is potentially important as an orna- mental and as a valuable source for germplasm in forage crop improvement programs. Unfortunately little information is available on the diversity within U.S. collections. This investigation was initiated to provide an extensive morphological analysis of each of eleven P. orientale accessions and to characterize the germplasm. Cytological studies revealed that all accessions were tetraploids (27=36) and that the method of reproduction being sexual or aposporus apomicts. Based on plant heights, leaf blade lengths and widths, inflorescence density, and growth habit in College Station (with a latitude of 31.5°), Texas, the acces- sions 269961, 330717, 271595, 433, and Cowboy appeared to have the best forage poten- tial. However, this has yet to be supported by studies on nutritional value. There were also some distinct taxonomic features such as the longest ligule length in PI 315867; central spikelet length in PI’s 215600 and 315867 which was significantly longer than all the other accessions. Furthermore, long anthers and filaments were observed in PI’s 330717 and 315867. Additional germplasm acquisitions from non-represented areas could expand genetic diversity in the germplasm collection. RESUMEN Pennisetum orientale Rich. es una especie polimérfica que tiene importancia potencial como planta ornamental y como fuente valiosa de germoplasma en los programas de mejora de forrajes. Desgraciadamente hay poca informacién disponible sobre la diversidad en las colecciones estadounidenses. Esta investigaci6n se inicié para realizar un analisis morfolégico amplio de cada una de once adquisiciones de P. orientale y para caracterizar el germoplasma. 'Corresponding author. Sipa 1701); 163-171. 1996 164 Stipa 17(1) Los estudios citol6gicos revelaron que todas las muestras eran tetraploides (27= 36) y que el método de reproduccién es sexual 0 apomixis aposporica. Basandose en la altura de las plantas, longitud y anchura de la lamina de la hoja, densidad a a inflorescencia y habito de crecimiento en re © — Station (con una latitud 31.5°), Texas, las muestras 269961, 330717, 271595, , y Cowboy parecen tener el mejor es forrajero. Sin embargo ésto aun no ie an ees eae ado con estudios del valor nutritivo. También se encontraron a ae caracteristicas taxonomicas diferentes, tales como la mayor longitud de la ligula en PI 315867, longitud de la espiguilla nial en PI’s 215600 y 315867 que era Par eens mas larga que en todas las otras muestras. Ademas se observaron filamentos y anteras largos en PI’s 330717 y 315867. Muestras adicionales de germoplasma de dreas no representadas podrian ampliar la diversidad genética de la coleccién de germoplasma. The genus Pennisetum is classified in the tribe Paniceae. Within this tribe Clayton and Renvoize (1982) placed Pennisetum in the Cenchrinae Dumort, a subtribe characterized by the presence of bristles and with the disarticu- lation below those bristles. Thirteen genera, including Pennisetum, are pres- ently included in this subtribe. Pennisetum, with about 90 species, is the largest genus of the Cenchrinae. Several Pennisetum species, in the tertiary gene pool of pearl millet [P. glaucum (L.) R.Br. are partially or completely apomictic (Jauhar 1981). Pennisetum ortentale L.C. Rich. is a polymorphic species in this gene pool consisting of different cytological races. This species, considered by Stewart (1945) as a Mediterranean grass, is distributed from 600—2,700 m eleva- tion in the western Himalayas of India westward through Asia Minor and N. Africa to Morocco. The cultivar, “Cowboy” (Pennisetum orientale) released in Texas, has been reported to be more winterhardy (Hussey et al. 1989) than buffelgrass [Pennisetum ciliare (L.) Link} suggesting that P. orzentale could be a valuable source for germplasm in forage improvement programs. Several interspecific hybrids have been recovered between P. orientale and P. ciliare. Any discussion of the morphology of P. orientale must also address apo- mixis because this taxon is believed to consist primarily of obligate apomictic accessions (Chatterji & Timothy 1969; Choda and Sharma 1981). There- fore any interpretation of the results must be viewed with the understand- ing of the effects of apomixis on grass morphology. The effects are that there 1s less morphological variability within populations. Apomixis, pro ably best defined as “ —_— nature's method of cloning plants through seed,” is a vegetative method of reproduction in which the embryo is formed without fertilization. Agamic complexes are usually arbitrary taxonomic groups of sexual and apomictic species that closely resemble each other morphologi- cally. Since P. ortentale is a polymorphic species, use of this species in forage improvement programs requires an extensive morphological and cytologi- cal analysis to identify and characterize the germplasm. Ramu et AL., Morphology of Pennisetum orientale 165 Richard (1805) originally described P. orientale from plants from the Orient. Later a general taxonomic description of P. orientale was given by Hrishi (1952). Sisodia (1972) compared the morphology of two P. orientale races and found the tetraploid race with longer leaves, longer tiller intern- ode length, and longer panicle length compared to the diploid race. A triploid race of this species was described by Vishnuvardhan and Lakshmi (1983). These descriptions covered only a few characteristics of some races from different geographical regions. Morphological characteristics vary from one region to another depending on the environmental conditions. Hence, there was a need for better understanding of the nature of morphological differences among accessions. This study, wherein different P. orientale ac- cessions were grown in a common location, will help to clarify relation- ships and to characterize the germplasm. Avdulov (1931) first reported the somatic chromosome number of P. orientale as 2n=36 although numbers of 27=18, 27, 35, 36, 45 and 54 have been reported as well (Chatterji & Timothy 1969; Dujardin & Hanna 1987). This suggests that x=9 is the base chromosome number and that aneup- loidy frequently occurs. However, Sahni and Bir (1985) report a gametic chromosome number of 7=7 for P. orientale. Confirmation of the chromo- some numbers and the method of reproduction for the accessions of this species are reported by Ramu (1994). These are essential for moving the breeding programs forward. MATERIALS AND METHODS Eleven plant introductions (PI's) of P. orientale were selected for a study of morphological characteristics. These accessions, their country of origin, and collection number of the voucher specimens that are deposited in the S.M. Tracy Herbarium (TAES) are listed in Table 1. Initial plantings of ten plants per accession were grown in the green- house during the fall of 1991. The plants were transplanted to the field west of College Station, Texas in early 1992. The soil at the field site is a Lufkin sandy loam with a pH of 8.3. The character list for the descriptive study is in Table 2. In addition, general growth habit of the plant was also recorded. For exact information on how each character was recorded refer to Ramu (1994). A pair-wise comparison among 11 accession means were tested for selected characters using the General Linear Model (SAS 1989) at alpha 0.05 level and the results tabulated in Table 3. Their ranges, means, and standard deviations for all the characters considered from the eleven accessions are listed in the Appendix of Ramu (1994). Chromosome counts were made from root tips using a modified Fuelgen technique (Hignight et al. d O01): 166 Sipa 17(1) Table 1. Pennisetum orientale accessions included in the study. Accession Collection site TAES* PI 215600 India, between Kalka and Simla along road, Pepsu. 30,000’ (1954). 201 PI 219610 India, Agriculture Research Station, Peshawar (1953), 202 PI 269961 Pakistan, pine forest, rocky hills, diacian 1200 m (1960). 203 PI 271593 unknown 204 PI 271596 India, steep wooded hills 20 km from Kalka, Simla Road to Nahan. 205 PI 314994 India. 206 PI 330717 Iran, Southern Esfahan. 207 433 India, seed obtained from ICRISAT. 208 PI 271595 India, steep wooded hills, high rainfall area, 28 miles from Kalka, Simla Kalka Road. 209 COW BOY Released by G.E. Pogue Seed Co., Kenedy, TX. 210 PI 315867 Czechoslovakia. 201 * Collection numbers of the first author are deposited in the 8.M.Tracy Herbarium (TAES). RESULTS AND DISCUSSION Species description of Pennisetum orientale L.C. Rich. Morphological measurements were taken on the eleven accessions grown ina common garden at College Station, Texas. Pennisetum orientale is a rhizomatous, perennial grass. RAzzome scales coriacious. Mid-rhizome scale length 2.2-8.8 cm, mid-internode length 0.4— 5.2 cm long. Flowering culms 38-149 cm tall, erect, much branched basally. Ligules 0.2-1.4 mm long, membranous. Leaf blades 12-35 cm long and 1.3—7.7 cm wide, linear to lanceolate, acuminate, glabrous to scabrous with ciliate margins. Inflorescences a panicle, 13.5-25.5 cm long and 2.6—5 cm wide, loose. First to second inflorescence internode lengths 0.5—10.5 mm long, second to third 0.5—7.7 mm long, and third to fourth 0.6-7.5 mm long. Involucre density 12—43 per basal 5 cm. Bristles in two series, length unequal, number inconsistent; inner bristles 6.5—18.5 mm long, ciliate with long hairs, green or purplish. Outer series of bristles 2.2-14 mm long, glabrous or scabrous, the two series are not always distinct. Spikelets per involucre ranged from one to five. Stipe scabrous to pubescent. Central spikelets 4.4—8.7 mm long, lanceolate; two florets per spikelet, occasion- ally three, lower floret triandrous or sterile, mostly staminate, upper floret perfect; first glumes 0.7—2.5 mm long, rudimentary, ovate to oblong, ob- tuse, veinless or faintly 1-veined; second glumes 0.8-4.8 mm long, ovate, lanceolate, veinless or 1- to 3-veined, usually 1-veined, acuminate; /ower floret lenmas 3.4—6.2 mm long, ovate, lanceolate, 2-(7)-veined, usually 4- (5S)-veined; paleas 2.3—5.7 mm long, oblong, obtuse, bifid, hyaline, 0- to 2- veined, usually 2-veined; pper floret lenmas 3.6-6.5 mm long, membranous, ovate, lanceolate, 5- to 7-veined, usually 5; paleas 3.5—5.9 Ramu ET AL., Morphology of Pennisetum orientale 167 Taste 2. Character list used in the morphological analysis, including the units of measurement. Vegetative: 1. Tallest tiller height (cm) 2. Shortest tiller height (cm) 3. Mid-rhizome internode length (cm) . Mid-rhizome scale length (cm) a Length of second uppermost leaf blade below the inflorescence (cm) 6. Width of sn uppermost leaf blade below the inflorescence (mm) 7. Ligule length of second uppermost leaf blade below the inflorescence (mm) ies ar . Length of the inflorescence (cm) i Width of the inflorescence (mm) 10. Inflorescence internode length (from the base)-Ist to 2nd (mm) 11. Inflorescence internode length (from the base)-2nd to 3rd (mm) 12. Inflorescence internode length (from the base)-3rd to 4th (mm 13. Number of involucres per lowest 5 cm section (# Spikelet: 14. Number of spikelets per involucre (mean of two involucres of the mid inflorescence) (#) 15. Length of inner bristle of the above spikelets (mm) 16. Length of outer bristle of the above spikelets (mm) 17. Length of the central spikelet (mm) 18. First glume length of the central spikelet (mm) 19. Number of the veins of the first glume (#) 20. Second glume length of the central spikelet (mm) 21. Number of the veins of the central spikelet (#) 22. Sexuality of the lowermos et 23. Lemma length of the lowermost floret (mm) 24. Number of the veins of the lowermost floret (#) 27. Lemma length of the uppermost floret (mm) 28. Number of the veins of the uppermost floret (#) 29. Palea length of the uppermost floret (mm 30. Number of the veins of the uppermost floret (#) 31. Filament length of the uppermost floret (mm) 32. Anther length of the uppermost floret (mm) 33. Stigma branch length of the uppermost floret (mm) 34. Caryopsis length (mm) 35. Caryopsis width (mm) mm long, lanceolate, acuminate with involute margins, 2- to 4-veined, usually 2. Stamens 3; filament 4.3-10 mm long, anthers 1.1—-3.2 mm long, yellowish or euaplich dehiscence by longitudinal pore; stigmas 2—6 mm long, two- branched, feathery; ovaries one-ovuled. Caryopses 1.2—2.6 mm long, 0.8-1.5 mm wide, elliptic. Flowering from spring until fall under favorable conditions. No disease or insect damage was observed at College Station, Texas. The culm height and leaf measurements of this study were not consis- tent with those reported by Sisodia (1972) as 243.8 cm and 12.5 mm, Tale 3. Mean and Duncan groupings* of selected morphological characters of Pennisetum orientale accessions. Accession Tallest Shortest Rhizome Rhizome Leaf Leaf Inflorescence Involucre/ Spiklet/ Caryopsis Caryopsis Ligule Central Upper Upper tiller tiller scale internode length width length — basal cm involucre length width length spikelet filament anther length length length length (cm) (mm) (cm) # # (mm) (mm) (mm) length length length : (cm) (cm) (cm) (mm) (mm) (mm) PI 215600 104.90 c* 50.88 f 4.30 abcd 2.99 abc 19.47cd 3.85 b 20.50 ab 23.50 d L.85 f 1.93 be 1.08 b 0.56cd 6.40a 6.73¢ 2.00d PI 219610 115.39b 57.96 3.73d 2.50 bed 24.02 ab 4.51b 22.40a 24.80 d 2.50 cde 2.07 abe 104 be 0.57 ¢ 5.75 bed 7.76b 2.16 cd def PI 269961 136.32a 81.66a 4.99 abc 1.60d 26.76a S5.52a 22.00a 26.10 cd 3.75 a 2.12b 1.096 O.73b 5.30ef 7.09 be 2.17 cd PI 271593 111.33 be 60.71 de 4.05 bed 2.51 bed 21.77 be 3.76b 20.40 ab 30.80 ab 2.15 ef 1.68d 0.94 d O.Sl cd 5.24 £ 6.49¢ 2.00d PI 271596 119.05 b 64.77 de 4.82 abcd 2.49 bed 24.63 ab 4.05 b 17.60 de 30.20 abe = 2.95 be 2.13 ab 1.08 b 0.58c 5.62cde 7.81b 2.39¢ PI 314994 112.42 be 65.41 de 4.53 abcd 1.82 d 22.88 abc 4.10 b 17.70 de 30.00 abe 2.70 bed 2.21 a 1.01 bed 0.55 cd 5.56def 7.87b 2.36c¢ PI 330717 109.80 be 55.80 ef 3.83 cd 1.64 d 17.00d 5.51 a 18.50 cd 23.50 d 3.10b 1.94 be 0.98 cd 0.50cd 5.94be 88la 2.71b 433 119.33 b 70.69 be 4.22 abcd 2.15 bed 22.77 abe 4.33 b 21.60 a 32.30 ab 2.25 def 1.84 cd LOL bed 0.45 d 5.55 def 7.95 ab 2.14 cd PI 271595 132.21a 73.53b 5.15 ab 3.63 a 21.51 be 4.13 b 21.10 ab 34.334 2.56 cde 2.20a 1.02 bed 0.49 cd 5.29ef 7.73 b 2.16 ced Cowboy 128.98 a 75.74ab5.39a 3.09 ab 23.00 abc 4.15 b 19.60 be 34.00 a 2.85 be 2.07 abe O0.98cd 0.52 cd 5.21 f 7.28 be 2.23 cd PI 315867 87.12d 49.78 f 4.62 abcd 2.05 cd 23.30 abe 3.91 b 16.50€ 27.90 bed 1.05 g 2.28 4 1.26a 1.03a 6.04b 8.00 ab 3.00a 3) * Duncan means followed by the same letter within a column are not significantly different at < a eo) pha = 0.05. 89 (T)LT Vals RaMu eT AL., Morphology of Pennisetum orientale 169 respectively. This could probably be due to ecological conditions at our common garden. However, involucre density is consistent with reports of Sisodia (1972) and Vishnuvardhan and Lakshmi (1983) and certain spike- let character observations of Hrishi (1952), Chromosome counts from root tip cells showed that all the accessions examined were tetraploid (27=4x=36). However, chromosome numbers ranging from 2” = 18, 27, 35, 45 and 54 have also been reported in addi- tion to 2” = 36 (Chatterji & Timothy 1969; Dujardin & Hanna 1987). Morphological analysis A pair-wise comparison of tiller height of eleven accessions (Table 3) revealed that the tallest tiller heights were recorded in accessions 269961, 271595, and Cowboy. The tiller heights in accessions 219610, 271593, 271596, 314994, 330717, and 433 were intermediate, while 315867 had the shortest tillers. Leaf blade length measurements show that PI 269961 blades were significantly longer than those of PI’s 215600, 271593, 271596, and 330717. Leaf blade widths of PI’s 269961 and 330717 were significantly wider than the other accessions. Taking leaf blade length and width to- gether, PI 269961 had the largest leaves. PI 330717 had the shortest leaves and was significantly wider than all the other accessions except for PI 269961. Intermediate leaf blade sizes were recorded for all the other acces- sions. All accessions were rhizomatous, however, differences were also ob- served between accessions with respect to rhizome scale length and intern- ode length. Cowboy had a rhizome scale length significantly longer than PI’s 219610, 271593, and 330717. The smallest rhizome scale length was recorded in PI 219610. There appeared to be an association between rhi- zome scale length and internode length. The long rhizome scale length observed in Cowboy and other accessions may protect the axillary buds of the rhizome from winter damage. In con- trast, the shorter rhizome internode length and long scale length observed in 269961 indicates more meristamatic buds per unit length. This may further enhance the plant’s capacity for regrowth. The suspected association of rhizome characters with winterhardiness might be addressed by multi-location trials where winter temperatures are more severe than College Station. No attempt was made in this research to quantify non-structural carbohydrate content, number of meristems per unit rhizome length or their association with winter survival. The data for inflorescence length and width indicate that PI 219610 had the longest and widest inflorescences. Intermediate length and width were observed in Cowboy while the inflorescences of PI’s 269961 and 433 were long with intermediate width. The smallest size inflorescence was observed itil 31580. 170 Sipa 17(1) Cowboy and PI 271595 had more involucres per basal 5 cm of inflorescence than PI’s 215600, 219610, 269961, 330717, and 315867 With respect to spikelets per involucre, PI 269961 had the highest num- ber of spikelets followed by PI’s 330717, 271596, and Cowboy. However, seed set fertility data is needed to judge the agronomic usefulness of this morphological character. Caryopsis (seed) size distribution across eleven accessions shows that PI 315867 had the largest seeds with the smallest seeds being observed in PI Zid: With regard to growth habit, all the accessions were erect. PI’s 269961 433, 271595, and Cowboy had robust vegetative growth, whereas the re- maining accessions were moderate, except PI 315867 which exhibited poor vegetative growth. The distinguishable character of PI 269961 is the droop- ing inflorescence giving the appearance of a bottlebrush. Among the eleven accessions studied there were also some distinct taxo- nomic features such as the longest ligule length in PI 315867; and central spikelet length in PI’s 215600 and 315867 which was significantly longer than all the other accessions. Furthermore, long anthers and filaments were observed in PI’s 330717 and 315867 The results of this study provide a better understanding of the nature of morphological differences among eleven P. orientale accessions. Based on plant heights, leaf blade lengths and widths, inflorescence density, and growth habit ina single area, the accessions 269961, 330717, 433, 271595, and Cowboy appeared to have the best forage potential. However, this needs to be supported by studies on nutritional quality. This comprehensive basic descriptive study provides the range of diver- sity among the accessions introduced from five geographical regions of the world. However, there were limitations because the study was conducted in one location. Therefore applicability of the results obtained in locations other than College Station need to be tested. Efforts should be made to collect additional accessions of different ploidy levels and from other eco- logical regions to contribute additional diversity to the germplasm collec- tion. Research reported here was conducted by the Texas Agricultural Experi- ment Station. REFERENCES AvpbuLov, N.P. 1931. Karyo-systematische undersuchung der amilie gramineen. Bull. Appl. , Genet. and Plant Breeding. Suppl. 43, Leningrad Cuarreryl, A.K. and D.H. Timotuy. 1969. Apomixis and tetraploidy in Pennisetum orientale Rich. Crop Sci. 9:796-799. Crayton, W.D. and S.A. RENvoIze. 1982. Pennisetum. In: Flora. er East Africa. Graminae (Part 3). Royal Botanical Gardens, Kew, London. Pp. 672-6 Ramu e? AL., Morphology of Pennisetum orientale 171 ar eine M. and W. Hanna. 1987. Inducing male fertility in crosses between pearl mil- let and Pennisetum orientale Rich. Sci. 27:65-68. Higmieer, K.W., E.C. BasHaw, M.A. Hussey. igen eae and Cae ia se of native apomictic ee Link. Bot. Gaz. 152:21 18. ee N.J. 1952. Studies on the cytogenetics of six species of Pennisetum and their com- parative morphology and anatomy. Genetica 26:280—356. Hussey, M.A., E.C. BasHaw, J. ALDERSON, R.M. Jones, W.R. OcumpauGH, and H.J. WHEDEMANN. 1989. Evaluation of winterhardy buflelgrass (Cenchrus ciliaris L.) germplasm. Forage Research in Texas. TAES CPR-473 Jana P.P. 1981. Cytogenetics and breeding of pearl millet and related species. Alan R. , New York, NY. an * "1994. Morphology, cytology, and mode of reproduction of Pennisetum orientale L ich, accessions. . Dissertation, Texas A&M University, pena Station. Ricuarps, L.C. 1805. Giininene In: C.H. Persoon. Synopsis Plant and : S. Bir. 1985. SOCGI plant chromosome number reports - HI. J. Cytol. 206. ae os. 205— SAS INSTITUTE = 1989. SAS/STAT User's Guipe, Version 6, Fourth Edition, Vol. 2 Cary, NC. Sisopia, K.P.S. 1972. Cytomorphological studies in Pennisetum orientale Rich. Cytologia 37:309-326. Stewart, R.R. 1945. The grasses of North-West India. Brittonia 5:404—468. VISHNUVARDHAN, Z. and N. LaksuMi. 1983. A new triploid race of Pennisetum orientale Rich. exhibiting aoe numerical mosaicism and neo-centric activity. Proc. con Acad. Sci. (Plant Sci.). 92:259-264. 172 Stipa 17(1) ANNOUNCEMENT The Gray Herbarium Card Index of New World Plants and the Harvard University Herbaria Type Specimen Collection Database The Gray Herbarium Card Index and the Harvard University Herbaria Type Collection databases are now available on the World Wide Web through the Harvard University Herbaria Web page. 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Sipa 17(1): 172. 1996 NEW SPECIES OF UREDINALES ON BIGNONIACEAE FROM BRAZIL JOE FE. HENNEN Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4060, U.S.A. HELEN M.P. SOTAO Museu Paraense Emilio Goeldi DBO, Caixa Postal, 599 Belém, PA 66040.170, BRASIL ABSTRACT Five new species of rust fungi (Uredinales) from Brazil parasitizing plant host species in the family Bignoniaceae are described. The new species are: Porotenus bibasiporulus, Porotenus biporus, and Uredo amapaensis on Memora spp.; Prospodium amapaensis on undetermined Bignoniaceae, and Prospodinm laevigatum on Mansoa sp. RESUMO Cinco novas espécies de fungos (Uredinales) sao descritos, provenientes do Brasil, parasitando espécies de plantas hospedeiras da familia Bignoniaceae. As novas espécies sao: Porotenus bibastporulus, Porotenus biporus, e Uredo amapaensis em Memora spp., Prospodium amapaensis em Bignoniaceae indeterminada e Prospodium laevigatum em Mansoa sp. INTRODUCTION The Uropyxidaceae, a family of nearly 200 species of plant rust fungi (Uredinales) worldwide, is composed of eleven genera, eight of which are neotropical. About 70 species distributed in four genera infect hosts mainly belonging to the Bignoniaceae. As part of our continuing research to mono- graph the Uropyxidaceae for the Neotropics, we propose five new species. Porotenus is a small neotropical genus of five species, two on Memora spp. and one on Amphilophium sp., hosts belonging to Bignoniaceae. Two other species parasitize Lippia in the Verbenaceae (Cummins & Hiratsuka 1983). We add two new species on Memora. The four species of Porotenus on Memora are known only from Brazil. Uredo is an anamorphic rust genus. Anamorphic taxa are also referred to as “imperfect” and “form” taxa. Anamorphic taxa are used to classify fungal spore states that represent only a part of the life cycle of a species. Connec- tions to other spore states in the life cycle may or may not be known (Cummins & Hiratsuka 1983). Both Uredo huallagensis P. Hennings, known Sioa 17(1): 173-184. 1996 174 Stipa 17(1) only from the type from Tarapoto, Peru, and Uredo amapaensis, which we describe below, have morphology and hosts (Memora spp.) that suggest t will belong to Porotenus when other spore states are discovered — ey Prospodium, with nearly 65 species, is the largest genus of Uropyxidaceae. It is strictly American, exending from northern Argentina to southern Texas and Florida in the Northern Hemisphere. Like Pororenus, it also parasitizes some members of Verbenaceae (Lippia, Aloysia, and Lantana) but most spe- cies occur on at least 19 genera of Bignoniaceae (Cummins 1940; Cummins & Hiratsuka 1983). We add two new species of Prospodium on Bignoniaceae. The Bignontaceae, with about 100 genera and 800 species worldwide, is primarily tropical and especially abundant in northern South eee (Cronquist 1981; Gentry 1980, 1992). In Brazil about 41 species of fa are known to parasitize plants of the Bignoniaceae (Hennen et <1 982): eae species concepts of rust fungi are based primarily on morphol- ogy, the various host specificity patterns of rusts are useful traits for their taxonomy. Knowledge of host identity, at least to family or genus, is often necessary for efficient identification of rust species (Cummins 1971; Gallegos & Cummins 1981; Lindquist 1982; Hiratsuka, N. et al. 1992). To identify rust species on the family Bignoniaceae it is useful to know that even though the teliospores of Porotenus, Prospodinm, and some other genera of rusts that infect bignoniaceous plants may resemble closely the teliospores of the ge- nus Pwcinia, no species of Puccinia are known to produce teliospores on members of this family. This kind of host relationship is especially impor- tant for identifying species of Pwccinia, the largest genus of rusts with nearly 4,000 species worldwide. Hennen and Buritica (1980) discussed some ideas about the evolution- ary pathway mechanisms of rusts, where the fungus-plant relationship ap- pears to be a great influence in the evolutionary process. MATERIALS AND METHODS The data are based on specimens in herbaria of the Biological Institute of Sao Paulo, Brazil (IBI), the Emilio Goeldi Museum of Para at Belém, Brazil (MG) and the Botanical Research Institute of Texas at Fort Worth, ‘Texas (BRIT). Semipermanent microscopic slide preparations of spores and sori were made by scrape mounts and freehand thin sections from herbarium speci- mens. Spores or thin sections were mounted in standard lactophenol solu- tion, heated and sealed. For better germ pore visibility some preparations were mounted in a saturated solution of chloral hydrate. Photomicrographic illustrations were made with differential interference contrast optics (Nomarski) and Polaroid type 55 positive/negative film HENNEN AND SorAo, New species of Uredinales POROTENUS TAXONOMY Telial traits in the four species of Porotenus on Memora are similar. Sori are scattered on the abaxial side of the leaflets, subepidermal in origin, erum- pent, pulvinate, 0.1—-0.5mm across, and cinnamon-brown but become whitish as the spores germinate in situ. The spores are mostly two-celled oblong-ellipsoid to cylindrical but often narrowed above, rounded below, somewhat constricted at the septa, and pedicellate; the walls are uniformly 1.0-1.5 pm thick, pale cinnamon-brown to nearly colorless, and smooth. Each probasidial cell has one germ pore at its distal end through which a metabasidium emerges on germination. Germination occurs without any noticeable dormancy period. Except for some overlapping spore size differ- ences, telial traits are not useful for identifying species of Porotenus on Memora. Variations in aecial and uredinial traits are more useful for characteriz- ing these species. Sori are considered as aecial if they closely encircle small patches of spermogonia and develop from the same mycelium. In general, spermogonia and aecia are encountered much less frequently than uredinia. Uredinia are randomly scattered singly or in small irregular groups over the abaxial side of leaflets. They are not directly connected to spermogonia. Both aecia and uredinia are subepidermal in origin, become erumpent, with the broken epidermis usually remaining as a partial covering over the sorus. The sori are 0.1—0.5mm across and somewhat powdery. The many loose spores produced in the sorus are chestnut-brown in mass. Each spore breaks off of its pedicel, leaving a definite scar termed a hilum at the proximal end of the spore. The spores are bilaterally symmetrical or radially asymmetri- cal usually with a convex and a concave side. The four species of Pororenis and one Uredo, including our new taxa on Memora, can be identified by their aeciospores or urediniospores using the following key. 1. Pores near the hilum 2. Each spore with one a on its convex, echinulate side, the concave side with a smooth patc P. memorae eee 2. Each spore with two pores, both the convex and concave sides with smooth patch P. bibasiporulus Hennen & Sotado Pores on the sides of the spores, more or less equatorial 3 3. Spore walls more or less evenly strongly echinulate, without smooth patches ry i apaensis Hennen & Sotio Ye 4 3. Spore walls with one or two smooth patches 4. Wall with one smooth patch on the concave side, pores near edge Pas the smooth patch concavus Viégas 4. Wall with two smooth patches, one on the concave side, one on aie vex side, pores in the echinulate area of the spore ........... P. biporus Hennen & Sotiio Porotenus biporus Hennen & Sotao, sp. nov. (Figs. 1, 2) Spermogonia amphigena in gregis. Aecia 2 Narcan cingentia, sporis urediniosporis similibus. Uredinia hypophylla; sporis (33—)35—40(-42) x 24—30(—33) pm asymmetricis L7G Sipa 1L7(1) N v wf a A Fics. 1, 2. Porotenus biporus. Fig. 1. Urediniospores, median focus. Fig. 2. Teloispores. Bar = ca 13 pm. HENNEN AND SoTAo, New species of Uredinales 177 late ellipsoidea vel ovoidea; poris 2 agar Telia hypophylla; sporis (36-)40—49 x 17-22 pm oblongo-ellipsoidea vel plus minusque cylindrica; episporio 1—-1.5 pm crasso, brunneolo vel praecipue hyalino laev! i in cellula poro apicali; pedicello tenuitunicat hyalino typice in fragmento curto; sporis non aoueene rn, Spermogonia amphigenous in groups on discolored areas that finally fall from the leaf leaving a little hole. Aecia arranged around the spermogonia, brown, spores pedicellate, similar to the urediniospores. Uredinia hypophyllous, scattered, dark brown, spores (33—)35—40(-42) x 24—30(- 33) pm, asymmetrical, broadly ellipsoid or oval with pores in optical sec- tion, slightly reniform with pores in the optical axis, wall 1.5—2 pm thick, dark cinnamon-brown, with a longitudinal irregular band of echinulae in which there are two more or less equatorial pores. Telia hypophyllous, brown- ish, pulvinate, spores (36-)40—49 < 17-22 pm, oblong-ellipsoid or more or less cylindrical, wall 1-1.5 pm thick, pale brownish to essentially color- less, smooth, pore apical in each cell, pedicel thin-walled, colorless, usually broken short, spores germinate without dormancy. ee pe: BRAZIL. Para: near Belém, Mocambo Forest Preserve, on Memora flavida (A. C.) Bureau & K. Schumann, 9 Jul 1979, J.F & M.M. Hennen 79-153 (HoLtoryre: IBI, ISOTYPE: - Additio ecimens examined: BRAZIL. Amapa: ae 27 Dec 1989, . Hennen G H. Pais - 175 (BRIT, IBI); Serra do Navio, 7 Jan 1990, J. Hennen & H. Sotdo 90-22 ae IBI). Distrito Federal: DF Hwy 15, 12 Feb 1981, as Heringer et al 6195 (ee T). aranhao: Sao Luis, forest in Sacavem (CAEMA), 20 Jan 1990, J. Hennen & H. Sotdo 90- a (BRIT, IBI). Minas Gerais: 40 km E of Lavras on a Hwy 116, 12 Mar 1984, ].&6 M. Hennen 84-250 (BRIT, IBI); 40 km E of Lavras, J. G M. Hennen 83-351 (BRIT, IBI). Para: elém, on Memora ee (A. DC.) Bureau - Schumann, 5 Nov 1967, Albuquerque s.n. (PUR F17762); Belém, on the grounds of EMBRAPA, on Memora sp., 17 Jan 1990, J. Hennen & H. Sotdo 90-43 (BRIT, IB); ae Estagao Cientifica Ferreira Penna, May 1994, J.& M. Hennen & H. Sotdo 94-172 (BRIT, IBD). Etymology.—The specific epithet refers to the two pores in the urediniospores. Distribution.—Porotenus biporus seems to be widespread in northeastern Brazil on several species of Memora from the cerrado and Amazon forest regions. Further field work is necessary, especially in the Amazon area to learn the full extent of its geographic and host range. The concave and convex sides of the anamorph spores have large irregu- lar smooth patches. The germ pores are along the sides opposite each other in the irregular echinulate band that surrounds the spore and separates the two smooth areas. Careful focusing on the spore surfaces is required for observing these traits. Porotenus bibasiporulus Hennen & Sotdo, sp. nov. (Figs. 3, 4) Spermogonia et aecia amphigena in areis parvis nigrescentibus hypertrophicatis, ad 5mm diam.; aeciosporis et viteciniospotis similibus. Uredinia hypophylla sparsa fuligenea; sporis Sipa 17(1) Fics. 3, 4. Porotenus bibasiporulus. Fig. 3. Urediniospores and one teliospore. Fig. 4. Teliospores, note the unusual three-celled spore. Figs. 5, 6. Uredo amapaensis. Fig. 5. Spores, median focus. Fig. 6. Same as Fig. 5 burt surface focus, note distribution of echinulae. Bar = ca 13 pm. HENNEN AND Sorao, New species of Uredinales 1? (31-)34—43(-48) x (14-)18-24 (—25) pm, plerumque ellipsoideis 22—24 jim latis; telia hypophylla sparsa cinnamomea, coacti-pulveracea; sporis billocularibus (parce trilocularibus), (40—)45—55(—66) x (15—)18—23 pm, plerumque fere cylindricis, leniter constrictis ad septum; episporio constanter 1—1.5 jam crasso laevi avellino vel fere hyalino; poro in cellula quaque apicali; pedicello hyalino tenuitunicato ad 50 pm longo. Spermogonia and aecia amphigenous in small, blackened, hypertrophied areas to more or less 5mm diam; aeciospores similar to urediniospores. Uredinia hypophyllous, scattered, dark brown, spores (3 1—)34-43(—48) x (14—)18—24(—25) pm, mostly ellipsoid and 22—24 pm wide with pores in the optical axis, strongly uniform and (14—)17—20 pam wide with pores lateral, thus conspicuously asymmetrical radially, wall uniformly 1.5 pm thick, chestnut-brown, echinulate except with two irregular smooth areas near the pores, pores 2, near the hilum. Telia hypophyllous, scattered, cin- namon-brown, felty-pulvinate, spores 2-celled (rarely 3-celled), (40—)45— 55(—66) x (15—)18—23 pm, mostly nearly cylindrical except constricted at septa, wall uniformly 1-1.5 pm thick, smooth, pale golden to nearly color- less, pore apical in each cell, pedicel colorless, thin-walled, to 50 pm. en Tyre: BRAZIL. Gots: dirt road off of Hwy 153 between Rialma and Riandépolis, on Memora nodosa (Manso) Miers, 15 Jul 1979, M.M. & J. FE Hennen 79-182 (HoLoryPe: IBI; ISOTYPE: IT) Additional specimens examined: BRAZIL. Dist. Federal: Brasilia, on Memora nodosa, 20 May 1981, E.P. Heringer 2023 (BRIT). Goids: Luziania, on Memora sp., 20 May 1982, E.P. Heringer 2219 (BRIT); 18 km SSW of Jataf, km 364, 18 Jul 1988, /.F Hennen & M.M. ileum & R.M. Lopez 88-58 (BRIT, IBI). Minas Gerais: Trés Marias, 25 Jun 1972, Inneco MO 2630750, BRIT-leaf fragment); 13 km SE of Paracatu, 19 Nov 1983, /.E & M.M He 83-758 (BRIT, IBD). Etymology.—The specific epithet refers to the two pores in the lower part of the urediniospores. Uredo huallagensis P. Hennings (Hedwigia 43:161. 1904) on Memora sp., from Tarapoto, Peru, U/es.n., 1902 (in Ule’s exsiccata, Mycotheca brasiliensis, n.23), also has two basal pores but perhaps is not synonymous, the pig- mentation being much paler and the radial asymmetry much less pro- nounce Uredo amapaensis Hennen & Sotao, sp. nov. (Figs. 5, 6) Spermogoniis, aecidia et teliis ignotis. Soris anamorphis ny, pophyllis, sparsis, O0.Olmm diam., cinnamomeus-brunneus, erumpentibus; sporis 26—31(—33) x 17—23(—24) pm, asym- metric, late Ae, ovatis vel badius, echinulatis, 2 poris germinationis plus minusve equatorialibus. Spermogonia, aecia and telia unknown. Sori hypophyllous, scattered, 0.1mm across, cinnamon-brown, pulverulent, ruptured epidermis evident; spores 26—31(—33) X 17-23(—24) pm, asymmetrical, broadly ellipsoid or oval with pores in optical section, slightly reniform with pores in optical 180 SIDA 17(1) axis, wall 1—1.5 pm chick, pale chestnut-brown, more or less evenly strongly echinulate, pores 2, more or less equatorial. Type: BRAZIL. AmapdA: Camaipi, on Memora cf. magnifica (Martius ex A. DC.) Bureau, 15 Nov 1987, J.E Hennen & H. Sotdo 87-124 (HoLoryre: IBI; isorypes: BRIT, MG). A Additional specimens examined: B AL. Para: Caxiuana, re Cientifica Ferreira Penna, on Memora cf. ei lee Benen ex A. DC.) Bureau, 2 . 1994, J.F & M. Hennen & H. Sotdo 94-17 I, MG); Caxiuana, IBAMA, 3 May 1994, .F GM. ee & H. Sotdo 94- es a MG); 20 Nov 1995, H. one et al 95-308 folic IBI, MG). Etymology.—The specific epithet refers to the Brazilian state of Amapa. This species is closely allied to the three species of Porotenus on Memora. The more or less evenly and strongly echinulate urediniospores differ from other species, all of which have irregular smooth areas. The three speci- mens from Caxiuana (J. Hennen & H. Sotdo 94-173, 94-182, 95-308) have arger spores than those from the type, 33-48(—5 1) x (15—)18—35(—38) pm. — PROSPODIUM TAXONOMY Three informal groups of species were recognized by Cummins (1940) based on soral traits and kind of life cycle: 1) aecia, uredinia, and telia subepidermal in origin and erumpent, Exprospodium; 2) aecia subepider- mal—erumpent, but uredinia and telia suprastomatal, Cyathopsora, 3) microcyclic, only spermogonia and telia, Neph/yctis. Our two new species belong to Cyathopsora, a group of about 15 species characterized by a sorus that develops as a minute basket-like structure from hyphae that emerge through stomata (suprastomatal). Prospodium amapaensis Hennen & Sotao, sp. nov. (Figs. 7—9) Spermogonia et aecia ignota. recy ) ees sparsa, Suprastomatales, oF —0.5mm diam., cyathiformes, cum paraphysibus 31—40 x 6-9 jim; sporis late globoideis vel globosis, poris 2, (ae ialis, poris a latarela visis, peer bilaminato, bicapitato, strato exteriore aculeato. Telia instar uredinearum; sporis 33—47 « 24-29 am, ad septum leniter constrictes ad septum apicem, episporio constanter 2-3 pm crasso. Spermogonia and aecia unknown. Uredinia scattered on adaxial side of leaflets, suprastomatal, 0.2—0.5mm across, brown, pulverulent; sori com- posed of a circular basal peridium surmounted with paraphyses around the margin, paraphyses 31—40 x 6-9 pum; spores 21-30 x 19-29 pm; globoid to broadly ellipsoid, wall 1—1.5 um thick, yellowish-brown incompletely two layered, the outer layer forming a longitudinal ring of irregularly in- terconnected spines, the ring divides distally to form a low crown (Cummins & Hiratsuka 1983, use the term “bicapitate” for this trait). The area of the lateral walls between the ring has widely spaced spines and two equatorial and opposite germpores. Telia like the uredinia but light chestnut brown; spores 37-47 X 24-29 pm, broadly ellipsoid, broadly rounded above and HENNEN AND Sorao, New species of Uredinales Fics. 7-9. Prospodium amapaensis. Fig. 7. Two urediniospores, center, and tw teliospores, median focus, note “halo” of echinulations around Heres Fig. 8. Same as Fig. 7 but surface focus, note that some echinulae are joined to each other. Fig. 9. Suprastomatal basket-like soral peridium, lateral view. Bar = ca 13 pm 182 Sipa 17(1) below, with a small umbo above and slightly constricted at the septum; wall evenly 2—3 pm thick, echinulate with echinulae irregularly and widely spaced with echinulae sometimes basally interconnected; pore apical in upper cell and near the pedicel in the lower cell; pedicel usually without append- ages and broken near spore. Type: BRAZIL. AMAPA: ca 10 km from Mazagao (ca 40 km SW of Macapa), on undetn. Bignoniaceae, 14 Nov 1987, J. Hennen & H. Sotdo 87-120 (woLoryeer: IBI; isorypes: BRIT, MG) Etymology.—The specific epithet refers to the Brazilian state of Amapa. Prospodium amapaensis belongs to a group of six other Prospodium species which are characterized by suprastomatal sori that are “basket like” (sec- tion “Cyathopsora,’ Cummins 1949), and the urediniospores have an apical corona (“bicapitate,” Cummins 1949). Within this group, P. amapaensis is most similar to P. tabebuticola on Tabebuia trom Puerto Rico but P. tabebuticola has teliospores that are larger, not so obtuse above and below, and are obvi- ously constricted at the septum. Their walls are bilaminate, the outer layer thin, transluscent, and noticeable especially at the septum. Prospodium laevigatum Hennen & Sotao, sp. nov. (Figs. 10-12) Spermogonia plerumque epipays Aecia hypophyllis, confluens, pulverulentus, cinnamomeus-brunneus; sporis 21-24 x 20-23 pm. Uredinia ignotis. Telia See sparsa, suprastomatales; peridium flavidus, 40-55 jim crasso; sporis 28-35 —21 4 oblongus ellipsoideus et valde constrictus ad septum, episporio 1.5 jam crasso, aureus vel pellucidus cinnamomeus-brunneus, levi Spermogonia mostly ee few in a group. Aecia hypophyllous opposite the spermogonia, confluent in a more or less circular group, pul- verulent, cinnamon-brown; spores mostly 21-24 x 20-23 pm, globoid or essentially so, the inner wall clear chestnut-brown, the outer layer apically coronate, absent over the pores, beset with bacilliform papillae, forming a complete halo when pores are in face view, but showing only as two caps or “ears” laterally from the apex; pores 2, equatorial in the smooth sides. Ure- dinia not seen, probably not produced. Telia hypophyllous, scattered suprastomatal, cyathiform with a basal, yellowish peridial cup 40-55 pm diam., the rim of the cup with low knobs (scarcely paraphyses); spores mostly 28-35 * 17-21 pm, oblong-ellipsoid and strongly constricted at septum, the cells nearly globoid, wall uniformly 1.5 pm thick, golden or clear cinnamom-brown, smooth, with a low lenticular, hyaline umbo over each pore, pore apical in upper cell, at septum in lower cell; pedicel thin-walled, colorless, fragile, about 25 pm long, without appendages; germination oc- curs without dormancy. Type: BRAZIL. Para: Belém, Mocambo Forest Preserve. on Mansoa kerere (Aublet) A. Gentry, 9 Jul 1979, /.F. & M.M. Hennen 79-153A (HoLotTYPE: IBI; isoryPE: BRIT). 183 HENNEN AND SoTdAo, New species of Uredinales 10 > Fics. 10-12. Prospodinm laevigatum. Fig. 10. Accigpes, mein focus, note (arrows) the apical corona that has a bicapitate or “two e ” appearance. Same as Fig. 10 but surface focus. Fig. 12. eters telial ca Fig. 11. sorus. View of bottom of sorus with teliospores emerging from top. Bar = 13 pm. 184 Sipa 17(1) Etymology.—The specific epithet refers to the smooth walled telispores. Prospodium laevigatum is the only species of the Cyathopsora group with smooth teliospores and coronate aeciospores. Only four other species of Prospodium have teliospores with smooth walls. Uredinia are probably lack- ing from the life cycle. ACKNOWLEDGMENTS The first author acknowledges the National Science Foundation for financial assistance for some of the field work. He also thanks Dr. G.B. Cummins for cooperation in monographic studies of the Uropyxidaceae of which this research is part. The authors thank the late Alwin Gentry and Nelson Rosa for identification of some host plants, and acknowledge the Instituto de Estudos e Pesquisas do Estado do Amapa (IEPA), and Museu Paraense Emilio Goeldi (MG) for field work logistical support. — REFERENCES Cronguist, A. 1981. An a system of classification of flowering plants. Colum- bia Univ. Press, New CUMMINS, GB.1 1940. he genus Prospodinm pea Lloydia 3:1-78. 1971. The rust fungi of cereals, grasses and bamboos. Springer Verlag. —____ and Y. Hirartsuka. 1983. Illustrated genera of rust fungi. APS. Press, St. Paul, Minnesota. Ga.ecos, H. and G.B. Cummins. 1981. Uredinales (royas) de Mexico. INIA, Culiacan. \V- Gentry, A. 1980. Bignoniaceae - Part I (Crescentieae and Tourrettieae). Fl. Neotrop. 25:1-130 —______.. 1992. Bignoniaceae - Part II (Tribe Tecomeae). Fl. Neotrop. 25:1—370. HENNEN, J.F. an P. BuriricA. 1980. A brief summary of modern rust taxonomic and evolutionary ees se Tottori Mycol. Inst. Japan. 18:243-256. a VEN, and M.B. FiGuEiREDO. 1982. Indice das ferrugens (Uredinal * i Brasil. Arq. Inst. Biol., Sao Paulo 49 (suppl. 1):1—201. Hirarsuka, N., S. Saro, M. Kakisuima, 8. KANEKO, T. Sato, T. Hirarsuka, K. Karsuya, Y Hrratsuka, Y. ONO, Y. HARADA, and K. Nakayama. 1992. The rust flora of Japan. Tsukuba oo. Tsukuba. LINDQUIST, J.C. 2. Royas de la republica Argentina y zonas limitrofes. INTA, Buenos Aires. A NEW COMBINATION IN THEVETIA (APOCY NACEAE) JUSTIN KIRK WILLIAMS Department of Botany University of Texas Ain, TX 267150 3 ABSTRACT Recent studies of Mexican a ahora have led to a reevaluation of the taxonomic status of Thevetia — (Pers.) K. Schum. var. pinifolia Standl. & Steyerm. The ta cifically distinct and ere accordingly with oo. new een Sais ‘pinifalia Standl, & Steyerm.) J.K. Williams, comb. nov. RESUMEN Los estudios recientes sobre las Apocynaceae mexicanas han permitido una reevaluacién del estatus taxonémico de Thevetia peruviana (Pers.) K. Schum. var. pinifolia Standl. & Steyerm. Este taxon se ve aqui como una especie distinta y por consiguiente es elevado de rango con la nueva combinacion, Thevetia pinifolia (Standl. & Steyerm.) J.K. Williams, comb. nov. Key Words. Thevetia, Apocynaceae, Flora, Mexico In their original description of Thevetia pernviana vat. pinifolia, Standley & Steyermark remarked that they were inclined to recognize the new taxon as a distinct species (Leavenworth 1946). They refrained from this course, however, because they interpreted two herbarium specimens as having leaves that were intermediate in form between the putative taxa. A reexamina- tion of these plants, (MacDaniels 244 and Palmer 27, F), indicates that they o not share with the typical variety a set of newly discovered characters. Instead, these plants are merely narrow leaved forms (4 mm wide) of T. peruviana (6—20 mm wide). Furthermore, these “intermediates” are grow- ing without the range of the typical variety, and it is suspected that they are not native but rather cultivated individuals. This hypothesis is founded on two principles; (1) given the cultivated popularity of Thevetia peruviana, it is difficult to ascertain its natural habitat (Woodson 1938), and (2) the “intermediates” were collected from the vicinity of Acapulco, an area of artificial development. e only character that Standley & Steyermark used to distinguish var. pinifolia from vat. peruviana was leaf dimension: filiform vs. linear-lanceolate, respectively. Field observations, along with a significant accumulation of herbarium specimens, has revealed alternative characters that separate var. Sipa 17(1): 185-190. 1996 186 Sipa 17(1) 4mm Fic. 1. Bracts of Thevetia pinifolia (a) and T. peruviana (b). pinifolia from var. peruviana. The new traits are pubescence (vs. wholly gla- brous), spreading corolla lobes (vs. erect), ovate bracts (vs. lanceolate) (Fig. 1), and an isolated geographical range (Fig. 2) (vs. widespread throughout Mexico and Central and South America). The var. perwviana does share several features with var. pinifolia. There is essentially no difference in the shape and dimensions of their calyx, corolla, and fruit. Also, both taxa lack prominent secondary lateral veins on the surface of their leaves, an important diagnostic feature emphasized by Woodson (1938). The only other species of Theveria lacking lateral veins, T. gaumer? Hemsl., is wholly glabrous, has elliptic leaves, and ovate bracts. Although var. pinifolia does not have secondary lateral veins on its leaves, it possesses other characters similar to a species that does: T. ovata (Cav.) A. DC. Both var. pinifolia and T. ovata have pubescence along their stems, leaves, and inflorescence. They also have similar geographical ranges, spread- ing corolla lobes, ovate bracts, and both possess similar calyx and corolla dimensions. Another species of Tevetia with secondary lateral veins and pubescence, 7. thevetioides (H.B.K.) K. Schum., has flowers twice the size of var. pinifolia, T. peruviana, and T. ovata. Given that var. pinifolia has a combination of characters used elsewhere in the genus to discern species, it seems evident that the taxon is a distinc- tive element showing little if any character intergradation with Thevetia peruviana. Because of this var. pinifolia is elevated to the rank of species. The two species can be readily distinguished by the following key: WILLIAMS, Thevetia 187 103°48'W 99°59'W 20°28'N 20° 28'N + + ee _ + eee MICHOACAN ee Fic. 2. Documented distribution of Thevetia pinifolia. 1. Leaves filiform, 1-2 mm wide, (9—)10—17 cm long; ae with peduncles, branches of new shoots, petioles, underside of leaf, bracts, and calyx pubes- cent; corolla lobes spreading, tube glabrous inside; one ovate, 4-3 mm long, 2 mm wide Fig. 1); fruits with shallow median ridge; restricted to Michoacan and Guerrero (Fig. 2) T. pinifolia _ Leaves lanceolate, (4-)6—15(—20) mm wide, 8—12(—15) cm long; plants wholly glabrous; corolla lobes erect, tube with indument inside; bracts lan- — ceolate, 2—3.5 mm long, | mm wide (Fig. 1); fruits with large median ridge; eidespiead throughout Mexico and extending into Central and South merica, commonly cultivated in both hemispheres (Fig. 3) ........0006 . peruviana Thevetia pinifolia (Standl. & ee le ) J.K. Williams, comb. nov. Basionym: Thevetia peruviana (Pers.) K. Schum. var. pinifolia Standl. & Steyerm., Amer. Mid]. Naturalist 36:185. 1946. ie pk: MEXICO. Micuoacan: trail from Apatzingan to Tan- citaro, 2,500 fr, 7 Aug 1940, W.C. Leavenworth 505 (HOLOTYPE: F!; isoTyPEs: GH, NY). Shrub to 4.5 m; new branches sparsely pubescent, 2—5 mm wide. Leaves at the upper S—20 cm of new branch growth, sessile, filiform, entire, dark 188 Sipa 17(1) Fic. 3. Documented distribution of Thevetia peruviana in Mexico, documentation by Gensel (1969). green, (9—)10—17 cm long, 1-2 mm wide; blade 0.5 mm wide on either side of midrib, lateral veins absent, pubescent along the bottom 4 mm. Inflorescence terminal, of 2—5(—7) flowers; peduncles 1.5—3.0 cm long, pubescent; bracts ovate, 4-5 mm long, 2 mm wide, pubescent; pedicels 1.0—2.5 cm long, pubescent. Calyx of five sepals, connate at the extreme base; sepals ovate, 7-10 mm long, pubescent, 6-10 colleters per sepal. Corolla infundibuliform, yellow, 40-50 mm long; tube 10-15 mm long, pubescent within along the upper portion of tube where grading into the cup, glabrous along the bottom four-fifths of tube; cup 9-15 mm long, glabrous within; lobes 20-30 mm long, spreading, glabrous. Anthers five, free, along the top edge of the tube protruding into the cup, past the infrastaminal hairs. Fruits a four-seeded drupe, 3—4 cm tall, 3—5 cm wide, greenish when ripening, blackish when mature. Representative specimens. MEXICO. Guerrero: Mpio. Zirandaro, 29 km SW of Zirandaro, ay to poy Gee 13 Jun 1982, Salas 1336 (MO); Mexicaltepec, 8 km al NW of Inguala, 14 Oct 1981, Nunéz 3339 (MEXU); Mpio. de Coyuca de Catalan, in Placeres del Oro, 2( ene 1980, a 2038 (MEXU),; en Paso de ee 18 km al SW de Coyuca de Catalan, 17 May 1978, Nunéz 660 (MEXU); Placeres del Oro, 2 Aug 1937, Hae etal. 10532 (TEX; MO; US); Placeres del Oro, 28 Jul 1936, vate etal. 9178 (TEX S); Coyuca, 3 Apr 1935, Hinton et al. 7579 (MO; US). Michoacan: Road to aaa 2km N of Apatzingan, growing in ravine along creek, 23 Aug 1995, ].K. Williams 95-56 (TEX); carretera entre Gallina y Gabriel Zamora, 5 km antes es Gallina, Aug 1987, Luna 18263 WILLIAMS, Thevetia 189 (TEX); en Las Colonias, Huetamo, Martinez 3639 (MEXU); En Huetamo, en El Barrio Alto, poe Mar 1982, Nunéz 3890 (MEXU); 2 km al SE de Pinzandaro, 9 Aug 1978, Rzedowski 35724 (MEXU); 13 km W of Apatzingan, 4 May 1966, Rzedowski 22307 (TEX; US); Apatzingan, arid slope with scattered trees, 13 Aug 1941, Leavenworth 1477 (F), bank of Rio Apatzingan, 5 Aug 1940, Leavenworth 463 (F); Apatzingan, 15 Aug 1938, Hinton et al. 12018 (F; TEX; US); Huetamo, 1 Mar 1934, Hinton et al, 5716 (US). Distribution.—This plant is apparently endemic to the tropical scrub forests of Michoacan and Guerrero, growing chiefly along arid plains and canyons between 360—1,000 m. It is interesting to note that this species shares a similar habitat and range with another apocynaceous endemic in Apatzingan (the type locality for both taxa), Echites woodsoniana Monachino (Williams, pers. obs.), a taxon that was recently collected by the author (Williams 95-80; TEX) not more than 30 m from a stand of T. pinifolia. Another rare apocynaceous taxon, Fernaldia asperoglottis Woodson, has also been collected from Apatzingan (Hinton et al. 15329; TEX), and surround- ing Guerrero (the type locality). After submitting the manuscript for review, it was brought to my atten- tion (Nesom, pers. comm.) that Gensel (1969), in an unpublished masters thesis, had independently recognized T. pinzfolia. All but one of the charac- ters we used to differentiate 7. pinifolia from T. peruviana were different. I did not notice the presence of indument in the corolla tube of T. peruviana, nor did I recognize a difference in the median ridge of the fruit. Gensel di not recognize the difference in external pubescence nor the shape of the bracts. From field work, however, we independently noticed the spreading corolla lobes of T. pinifolia and the erect lobes of T. permviana. | have in- cluded the characters discussed by Gensel (1969) in the key. I should point out the Gensel annotated the two “intermediates” (see above) as T. pinifolia. The flowers of these specimens have been destroyed through the years and it is not possible to examine the inside of the corolla tube for indument. However, the specimens have lanceolate bracts and lack external pubes- cence, for these reasons I treat them as T. peruviana. In several villages near Apatzingan the plants appear cultivated, but upon questioning the natives I was informed that the plants were wild and that they had removed the surrounding vegetation for farming, leaving Thevetia for aesthetic purposes. Label data on Nwnéz 2038 notes that children eat the pulp of Thevetia fruits. Alchough Morton (1982) reports that all parts of the plant except the pulp contain toxic glycosides (ex. thevetin and peruvoside) consuption of the pulp by children is not recommended as the seeds are highly toxic (Woodson 1937). A child’s accidental swallowing of only one to two seeds will typically result in their death. The average toxic dosage for an adult is eight to ten seeds. 190 Sipa 17(1) ACKNOWLEDGMENTS Iam grateful to Guy Nesom for informing me of the Gensel thesis, and to the curators of the following herbaria for allowing me the opportunity to observe their specimens: F, BRIT, LL, MEXU, MO, TEX, — REFERENCES GrnseL, W.H. 1969. A revision of the genus Thevetia (Apocynaceae). Masters Thesis Uni- versity of Connecticut. LEaAvENWoRTH, W.C. 1946. A preliminary study of the vegetation of the region between >» Tancitaro ac the Rio Tepalcatepec, Michoacan, Mexico. Amer. Midl. Naturalist a : 37-2006. Morton, J.F. 1982. Plants poisonous to people in Florida and other areas. Southeastern Printing Co., Inc., Stuart, FL. - Morton, J.B, E. Atvarez, and C. QuIANONEZ. 1990. Loroco, pale ee (Apocynaceae): a arene ae e (swe of Central America. Eco ot 310. ares, R.E., JR. eae or foe noteworthy eee of nate America eee: oe en -12. 1938. ee - Amer. Fl. 29:103-192 UNA NUEVA ESPECIE DE AGAVE SUBGENERO AGAVE (AGAVACEAE) DE MEXICO JOSE A. VILLARREAL Q. Departamento de Botdnica Universidad Auténoma Agraria “Antonio Narro” Buenavista, Saltillo, Coahuila 25315, MEXICO RESUMEN Se describe Agave montana como especie nueva de la regién de la Sierra Madre Oriental. Esta relacionada con A. gentryiy A. ak bas pero difiere de ambas por sus rosetas grandes con hojas mas numerosas, panic - as »vadas con mayor nimero de ramas floriferas y bracteas dentadas. Se presenta una ilustracién con los detalles de la planta. ABSTRACT Agave montana is described as a new species from the Sierra Madre Oriental. It is relatec to A. gentry: and A. parrasana but differs from both in its larger rosettes with more numer- ous leaves, ovate panicles with higher number of floriferous branches and armed bracts. A detailed illustration is included. Colectas realizadas en la cima de la sierra de la Marta en los limites de los estados de Coahuila y Nuevo Le6én, adicionan una nueva especie de Agave subgénero A gave. Agave montana Villarreal, sp. nov. (Figs. | y 2) Agave gentryi Ullrich affinis, sed rosula compacta |1.4—1.6 m in diametro, foliis numerosioribus ellipticis viridi-flavis, inflorescentia ovata, ramis floriferis numerosioribus, bracteis denticulatis et floribus 6-7 cm longis discrepans. Plantas multianuales, no rizomatosas; rosetas hemisféricas, compactas, 1.40-1.65 m didmetro, 90—1.25 m alto; hojas arregladas en 12—16 hileras, 84-112 por planta, cortamente elipticas, ligeramente céncavas a casi planas, 30—40 cm largo, 15-17 cm ancho, de color verde-amarillento, base ensanchada, apice acuminado con borde de color café-purpura, espina de 3— 5 cm largo, margen recto, con 16—18 dientes grisdceos antrorsos y retrorsos por lado, separados 2.5—3.5 cm; inflorescencia 3.5—4.5 m alto, ovoide, con alrededor de 30 ents florales, de 25-35 cm largo, ubicadas en las 2/5 partes superiores del escapo; bracteas elfpticas, 18-25 cm largo, 8-12 cm ancho, con dientes marginales y espina terminal, cOncavas, suculentas, café- rojizas, cubriendo completamente el tallo floral; tallo floral 12-16 cm didmetro; flores 6-7 cm largo, amarillas, suculentas; Soca 1-1.5 cm largo; ovario 3 cm largo; tubo del perianto 2 cm largo, 12—14 mm ancho; Sipa 17(1): 191-195. 1996 Fic. 1. Agave montana A. Hoja. B. bractea. C. flor, corte longitudinal. D. flor y E. cépsula. tépalos lanceolados, 2 cm largo, 3—4 mm ancho; filamentos 5—6 cm largo, 3 mm ancho, scenes insertos en la base de los tépalos; anteras de 19— - mm largo, 2 mm ancho, amarillas; ca4psulas oblongas, S—6 cm largo, 1.5— cm ancho, de color cafe oscuro; semillas lacrimiformes, 5-6 mm largo, 3-4 mm ancho, de color negro brillante. ee MEXICO. Nuevo LEon: Municipio de Rayones, cima de la sierra de la Marta, 42 km al E de San Antonio de las Alazanas, 25 09°N,100 23 W, matorral bajo de Arctostaphy- los pet Ceanothus i ee Pinus culminicola, P. hartwegit, Quercus gregeit, Ceanothus eregeil, Cercoarpus, entre otras, 3,300 m, 5 May 1995, ].A. Villarreal 8120, M.A. Carranza y JA. ee MEXU; tsotripos: ANSM, ENCB). VILLARREAL Q., Una nueva especie de Agave (ee TasLa 1. Comparaci6n de algunas caracteristicas entre Agave montana, A. gentryi y A. parrasana. Caracter Agave montana Agave gentryi Agave parrasana Didmetro de la planta) 140-165 cm 60-100 cm 30-50 cm Hijuelos rizomaticos ausentes presentes ocasionales Numero de hojas 85-115 30-45 40-60 Limbo de la hoja casi plano céncavo casi plano Taman de hoja 30-40 X 15-18 cm 60-100 X 17-26 cm 20-30 X 10-15 cm Tipo de dientes curvos casi rectos curvos Color de hoja verde-amarillencto verde glauco-grisaceo Forma de la panicula ovada elfptica elfptica Numero de ramillas 20-30 10-18 10-15 orales Borde de bracteas dentado entero entero Longitud de flores 6-7 cm 7-9 cm 5-6 cm Material adicional examinado: Localidad tipo, 6 Sep 1994, J.A. Villarreal 7905 y M.A. Carranza (ANSM). La especie descrita se localiza en la sierra de la Marta, en laderas de rocas calizas con exposicié6n W-SW entre los 3,200—3,400 m. La poblacién es esparcida extendiendose por toda la ladera en una comunidad mas xerofila que la de la exposicidn contraria de la misma sierra. Agave montana esta relacionada con A. gentryi Ullrich (nombre asignado por Ullrich (1990) al tradicionalmente conocido A. macroculmis) y con A. parrasana Berger por su tallo floral gueso y completamente cubierto por brdcteas céncavas y suculentas, asi como por su habitat en bosques de pino- encino de la Sierra Madre Oriental. Una comparacion de las caracteristicas mas sobresalientes de estos taxa se presenta en la Tabla 1. La nueva especie se diferencia por sus rosetas de aproximadamente 1.5 m de didmetro, con hojas de color verde amarillento y en mayor cantidad, asi como en el borde dentado de las brdcteas. Se distingue de A. gentry7, la especie al parecer mds cercana, por las hojas mds cortas y numerosas, casi planas y de color verde amarillento, panfculas ovadas, mds anchas cerca de la base, con mayor numero de ramillas florales y flores mas pequefias (Tabla 1). La descripcién para A. macroculmis en Gentry (1982) y la de A. gentryi por Ullrich (1990) som muy amplias e incluyen caracteristicas de las dos especies (A. gentryi y A. montana). La nueva especie propuesta puede separarse siguiendo las caracteristicas de la Tabala 1, por lo que A. gentry: se restringe a plantas con rosetas laxas de 60-100 cm de didmetro, hojas largamente elfpticas 60-100 x 17-26 cm, céncavas, de color verde oscuro, 194 Sipa 17(1) Oe Nee ey Bans Fic. 2. Agave montana en su habitat y plantas en floracién y fructificacién. ofe : f: VILLARREAL Q., Una nueva especie de Agave 195 inflorescencias en paniculas elfpticas, mds anchas en la parte media, con 10—28 ramas florfferas, bracetas enteras y flores de 7—9 cm largo Las tres especies comparadas en el Tabla 1 comparten una serie de caracteres y se les puede agrupar bajo las siguentes caracteristicas: Plantas pequefias a tamafio medio; rosetas compactas con hojas numerosas o laxas y menor numero de hojas; hojas menos de 1 m largo; pedunculo largo y grueso con bracteas elipticas, c6ncavas, suculentas, erectas, con tintes rojo-purpura, cubriendo totalmente o casi totalmente el tallo floral, con amontonamiento de bracteas debajo de la panicula; paniculas ovadas de 1/3—2/5 del escapo floral; floraci6n en primavera; distribucién en la Sierra Madre Oriental. AGRADECIMIENTOS Se agradece la revisién del manuscrito y la diagnosis en Latin al Dr. J. Rzedowski y la elaboracién del dibujo al bidlogo Miguel A. Carranza P. REFERENCIAS Gentry, H.S. 1982. Agaves of Continental North America. University of Arizona Press, son. ULtricy, B. eee Agave macroculmis Todaro en Agave gentryi Ullrich spec. nov. Succulenta 214, 69(10):21 196 Stipa 17(1) BOOK REVIEW NELSON, CHARLES E. and ALLAN Prosert. 1994. A Man Who Can Speak of Plants. (ISBN 0 9524847 06, pbk). privately published by Dr. E. Charles Nelson, 14 Connaught Parade, Phibsborough, Dublin 7, Ireland, $30.00, 181 pp, 8 color plates, 52 b/w. This book is the life story of Dr. Thomas Coulter (1793-1843), an Irish botanist that lived and worked in Mexico and California in the 1820s and 1830s. He was a physician and also a naturalist with broad interests in animals and plants. He discovered and his name is forever enshrined as part of the binomial for the big-cone pine (Pinus coulteri) and the white-blossomed poppy, matilija (Rommneya coulteri). His plant collections, especially of xican cacti, were distributed to eae gardens in Europe. His talents extended far beyond the study of botany, including insects and reptiles, and activities such as angling, shooting, managing silver and lead mines, and as a surveyor. ife from youth in a beginning chapter pen The book is organized chronologically, tracing his Child and university student (1793-1820); his middle years Student of Botany (1820— 1824) - 15 pages; Miner and physician in Mexico (1824-1829) - 60 pages; Explorer and plant-hunter in California (1829-1834) - 21 pages; ro his final failing h health and death ‘ wreck of a man’his final decade (1834-1843). His failing health and early death limited his publication productivity once his collecting days were over The botanical content of this book is rather limited when compared to Coulter's other life activities. One of the highlights of Coulter’s botanical career was his study and mono- graph of the Dipsacaceae (teasel and scabious family) under the tutelage of the ape botanist Augustin-Pyramus de Candolle of Geneva, Switzerland. Coulter's trip co lower California has commentary about the excessive heat, his poor health and the lack of inter- esting plants. Here he meets David Douglas who is a Scottish botanist collecting plants for William Hooker of the Royal Horticulcural Society of London. The title of this book comes from a letter written by Douglas that praises Coulter for his botanical work and refers to him as “a man who can speak of plants.” Many of Coulter's collections come from the environs of Monterey where he found the big cone a Pinus conlteri, and four other new species of pines. The beautiful white Howered Matilija poppy was named to honor Coulter by his friend Dr. Romney Robinson, ee ie This beautiful species is surely Coulter's finest memorial and was introduced into cultivation in Europe about 1875. William Harvey described new genera from Coulter's California's collections: Lyrocarpa coulteri and Dithyrea . nica (Brassicaceae, the cabbage family); Whitlavia grandiflora ind W. minor in the Hydrophyllaceae; and Berg/nia virgata . Many of Coulter's collections are at Trinity College, Dublin where he established this famous herbarium with his own w World collections. He is also noted for his famous pine cone collec . He ne published any papers on his Mexican and Cal ifornia collections and eer no new species ‘rom America. Appendix I provides a glossary of Spanish words and mining terms. Appendix I provides the original texts of Coulter's letters about upper California to the Royal Geographical Society of London. Appendix II has Thomas Coulter's herbarium specimens, but this is not a complete listing of all the new taxa based on Coulter's collections. Such a list may not have been possible to compile but it certainly would provide valuable reference information. Anyone interested in the floras of Mexico and California will welcome this book as an important source of botanical s on plant exploration in the early 1800s.—Harold W. Keller, Research Associate. BRIT Sipa 17(1): 196. 1996 THE MEXICAN GENERA OF THE APOCYNACEAE (SENSU A. DC.), WITH KEY AND ADDITIONAL TAXONOMIC NOTES JUSTIN KIRK WILLIAMS Department of Botany University of Texas Austin, TX 78713, USA. ABSTRACT A key to the apocynaceous genera of Mexico is presented. The study is based on litera- ture, field observations, and herbarium studies. The family is represented in Mexico by thirty genera (twenty-five native, four Old World cultivars, and one South American cul- tivar) and approximately ninety species (eighty-five native, four Old World cultivars, and one South American cultivar). Bibliographical references for the native genera are included, as are illustrations, a distribution list of genera by state, and a Ae of morphological novelties. RESUMEN Se presenta una clave de los géneros de Apocynaceae de México. El estudio esta basado en bibliograffa, observaciones de campo y estudios de herbario. La familia esta representada en México por treinta géneros (veinticinco autéctonos, cuatro del Viejo Mundo y uno Sudamérica). Se incluyen referencias bibliograficas para los géneros autéctonos, asi como ilustraciones, una lista de distribucidn de géneros por estados y una tabla con las novedades morfol6égicas. Key Worps: Apocynaceae, Asclepiadaceae, Flora, Mexico In the course of revisionary studies on various apocynaceous genera it came to my attention that the identification of numerous specimens as to genus, in several major herbaria, was erroneous. The following key has been constructed to serve as a supplementary aid to the identification of the apocynaceous genera of Mexico. Considerable detail and repetition has been included in the key to insure the utmost accuracy. I have avoided the use of minute (ex. calycine colleters) and variable characters (ex. phyllotaxy) when- ever possible, however, as there is considerable convergence within the family, at times their use was necessary. The characters used within the key do not necessarily represent the entire genus, but merely represent it for those species growing within Mexico. It should also be emphasized that the key is in no way meant to reflect natural groupings; after various efforts it was deemed more effective to arrange it artificially. The native Apocynaceae genera in Mexico are in an alphabetical list following the generic key. Recognition of genera and their accepted names Sipa 17(1): 197-213. 1996 198 Sipa 17(1) within the list and key has been adopted from Leeuwenberg (1994). Ge- neric synonyms within the list have been included only for those names that I have seen used in the current literature and various herbaria. The number of species in Mexico (approximate or exact) for each genus is given in the list. The species epithet is given within the key for all genera repre- sented in Mexico by one species. The most recent and pertinent revisionary studies (dating from 1930), as well as publications of newly described spe- cies (the state of the type locality is provided), have also been provided for all of the New World genera. I have not included references for any of the Old World, non-native, cultivated genera: Carissa, Catharanthus, Nerinm, and Vinca. All cultivated genera are represented in Mexico by one species (given in key). A list of general works on the Apocynaceae of Mexico and related areas (References), a tentative list of the distribution of all native genera in Mexico by state (Table 1), and a list of morphological novelties (Table 2) have also been provided. Illustrations have been included for only those characters that most readily represent the particular taxon in ques- tion and that distinguish it from morphologically similar genera. TABLE 1. A list of the distibutions, by state, of the native genera of the Mexican Apocynaceae. SPECIES BJN BJS SON CHI COA NUE TAM SIN DUR ZAC AGU SAN NAY JAL Alstonia x Amsonia xX Xx Xx XxX Apocynum X x xX xX Aspidosperma Camerari Echites Xx Fernaldia X x Forsteronia xX Haplophyton x x Xx aubertia XxX Xx Mandevilla x Xx x x Xx D:¢ x xX Xx Pentalir xX Plumeria x x XxX 4 xX xX Xx Prestonia Xx xX uvolfia x X x xX Rhabdadenia Stemmadenia xX xX x x x Xx Tabernaemontana Xx X xX Xx x Telosiphonia xX XxX X XxX X x X xX XxX xX NX xX Thevetia x x Xx x Xx x Thenardia Xx Tintinnabularia Vallesia Xx xX xX Xx x WILLIAMS, Mexican genera of Apocynaceae 199 Lastly, it should be noted that with the recent advent of cladistic analy- sis it has become evident that the Apocynaceae as traditionally circum- scribed is paraphyletic. Judd et al. (1994) have recently suggested the in- clusion of the Asclepiadaceae within the Apocynaceae. Although this view is becoming more widely accepted (Struwe et al. 1994) I treat the Apocynaceae (sensu A. DC., de Candolle 1844) here in its traditional sense (i.e., distinct from the Asclepiadaceae). The following key is based on literature, field observations, and her- barium studies from the following institutions: BRIT, F LL, MEXU, MO, TEX, and US. KEY TO THE GENERA OF MEXICAN APOCYNACEAE 1. Anthers free from pistil head, bases obtuse (prolonged into a fork in Allamanda, Tabernaemontana, and Stemmadenia), aestivation of corolla bud to the left (right in Haplophyton cimicidium A, DC.); plant an herb, shrub, or LEGe, cheate) scandent (A/llamanda); leaves alternate, opposite, or whorled; a dry or fleshy follicle, berry or drupe; seeds entire, winged, or ciliat he margin (A/stonia), not comose or if so then both apically and basally comose (Haplophyton) 2 COL GUA QUE HID MIC MEX MOR TLA PUE VER GUE OAX TAB CPS CAM YUC QUI Xx Xx xX x x Xx x Xx Xx Xx x x Xx x x Xx x x Xx Xx Xx x x x x Xx X x Xx x Xx x Xx Xx x Xx Xx x x x Xx x Xx x Xx Xx x x Xx x x x Xx Xx Xx Xx x x xX Xx x Xx x Xx x x Xx x Xx xX Xx Xx x Xx x x x xX Xx xX Xx X x x Xx Xx Xx x Xx x 4 x Xx x x Xx Xx x x x x x x Xx Xx x X Xx xX Xx xX x Xx x Xx Xx x Xx x x x xX XK x x x x x Xx Xx Xx Xx Xx x x xX Xx x x Xx x x xX XX xX x Xx x Xx Xx x x Xx Xx x xX Xx x 200 Sipa 17(1) Paste 2. External morphological novelties of Apocynaceae genera and species found in Mexico. Plants spiny: Carissa a (Ecklon) Plants densely yellow tomentose: Pieroni anda L.O. Wms., P. mexicana Donn. Smith Leaves alternate: Agents Aspidosperma, Haplophyton, Plumeria, Theveria, Vallesia Leaves whorled: A//amanda, Alstonia, Nerium, Rauvolfa Leaf hairs with multi-cellular bases: Hap/ophyton Leaves subeoratery various isepectts of Mandevilla Leaves L f petiole: Forsteronia, Mandevilla, Mesechites, Telosiphonia, Tintinnabularia Leaves with glands along the adaxial midrib: Mandevilla hirsuta (Rich.) K. Schum., M. subsagittata (R. & P.) Woodson, M. villosa (Meirs) Woodson Leaves with domatia in the axils of the abaxial midrib: Forsteronia, Tintinnabularia Corolla tube twisted: Eebites umbellata Jacq., Lanbert Corolla with thickened annulus around mouth: en Prestonta Calyx of ane sepals, the outer two fused, inner two free: Aspidosperma megalocarpon Mill. Arg. Calyx red: ue tonta portobellensts (Beurl.) Woodson (purple), Tintinnabularia (burgundy eae coiled: narvdia flortbunda Kunth YS nt wit pee amentous apical appendages: Cameraria, Nerium, Pentalinon, Tintinnabularia nther exs serted: na Steronia, Laubertia, Prestonia, Tabernaemontana amygdatlifolia Jacq. (only slightly so in T. alba Mill.), Thena Fruits of two ae folli x various species of Mandevilla, Thenardia Fruits spiny: A//amay Fruits red: Carissa, ae obfia, Thevetia abouai (L.) A. DC. Seeds eae Allamanda, Aspidosperma, Plumeria Seeds with both basal and apical coma: Hap/ophyton Seeds with ciliate margins: A/stonia — 2 (1). Plants with spines in che axils of the leaves; fruit an indehiscent browinsh-red berry, seeds embedded in a pulp; occasionally naturalized species Carissa macrocarpa (Ecklon) A. DC. 2. Plants without spines; fruic a follicle, berry or drupe; native or cultivar 3 (2). Anthers with distinct apical filamentous appendages; inflorescence of 1-5(—8) flowers, flowers white; fruits resembling single seeded samaras Cameraria latifolia L. 3. Anthers without distinct apical filamentous appendages; inflorescence of 1-80 flowers, flowers blue, pink, red, yellow, or white; fruits not samaras 4 4 (3). Flowers with two distinct nectaries adjacent to ovary; plant her- baceous; occasionally naturalized species 5 5 (4). Plant prostrate; flowers solitary in leaf axils, funnelform, blue; corolla tube not apically constricted; filaments bent and longer than anthers; anthers incompletely fertile; nectaries shorter than ovary Vinca minor L. . Plant erect; flowers 2—4 in axillary cymes, salverform, red, white, or pink; corolla tube apically constricted; filaments straight and shorter than anthers; anthers completely fertile; nectaries as long as or longer than ovary atharanthus roseus (L.) G. Don 4. Flowers with a solitary annular nectary or nectary absent; plant a herb, tree or shrub; native or cultivatec 6 A WitiiaMs, Mexican genera of Apocynaceae 201 6 (A). so pee herb; flowers yellow; inflorescence reduced to a solitary flower in leaf axil; petals of the bud overlapping to the right (H. ae A. DC.) or overlapping to the left (H. crooksii (L. D Benson) L.D. Benson); leaves alternate to subverticillate; leaf ie with multi-cellular bases; fruit a linear follicle; seeds with both es and apical coma Haplophyton . Herb, tree, or shrub; flowers bluish, white, or yellow; inflorescence flowers; petals of the bud overlapping to the left; leaves alternate, opposite, or whorled; leaves glabrous or if pubescent then with simple hairs; fruit a linear or circular follicle, berry, or drupe; seeds without coma, or if pubescent then with a ciliate margin (Alstonia) 7 7 (G). Leaves alternat 8 8 (7). Herbaceous Seal to 0.7 m high; flowers bluish; fruit a linear follicle with many simple seeds; restricted to northern Mexico Amsonia 8. Tree or shrub from 1—35 m tall; flowers white, yellow, or red; fruit a linear or circular follicle with many winged seeds, or drupe with one-four nutlets; distributed Spells MCHICO pase ccrsdsesuvas 9 9 (8). Tree from 7—35 m; calyx of 4 sepals, outer 2 fused and inner two free (A. Saws Mill. Arg.), or five regular oe (A. spruceanum Benth.); fruit a circular follicle, with winged seeds Aspidosperma 9. Sheil from 1—G6 m tall; calyx of 5 free sepals; fruit a broad follicle, berry or drupe 10 10 (9). Flowers to 1.5 cm long, typically less, salverform, white; fruic a one-seeded white fleshy drupe Vallesia I 10. Flowers much longer than 1.5 cm, funnelform or salver- form, white, yellow, or red; fruit a many-seeded follicle or 1-4 seeded fleshy drupe 11 11 (10). Flowers o. white, red, or yellowish; corolla orifice small; calyx without colleters; fruit a thick, stout, dry follicle; eed winge Plumeria 11. SS anast ee in T. abouat (L.) A. C.), bright yellow or creamy; corolla orifice large; ao with colleters; fruit a triangulate fleshy drupe; seeds ov Thevetia fe —. opposite or an d 2 (7). Leaves opposite; calyx with colleters; anther bases pro longed into a fork; seeds with arils i 13 (12). Inflorescence of 1—4(—10) flowers; corolla large and showy, tube (8—)15—30 mm long, funnelform, cream-colored or yellow; anthers included; sepals peer leafy and not clasping the corolla base Inflorescence of 10-50 flowers; corolla small, tube 7-16 m long, salverform, white; anthers exserted or inserted; ann mostly thick and clasping the corolla base. Tabernaemontana 12. te whorled (occasionally opposite); calyx without colleters; anther bases obtuse or prolonged into a fork (A//amanda), seeds without arils 14 — We — 202 Sipa 17(1) 14 (12). Ancther bases prolonged into a fork; corolla large, to 8 cm long, bright yellow; leaves without glands along the petiole; ovary 1; styles not cleft at base; fruit a spiny capsule; seeds winged; occasionally naturalized species amanda cathartica L. . Anther bases obtuse; corolla small, to 2. cm long, whi leaves with or witl v4 — yout glands along the petiole; ovaries 2; styles cleft at base; fruit not spiny; seeds not winged; nativ 15 (14). Petiole scattered with many colleters along its length; fruit a fleshy reddish-brown drupe; seeds glabrous us Rauvolfia iole without colleters; fruit a linear dry follicle; eds ciliate along the margin Anthers fused to pistil head, bases prolonged into a fork (obtuse in Ferna/dia) aestivation of corolla bud to the right; plants typically scandent (herba- ceous in eH shrubby in Nerivm, a lalhi Mandevilla karwinskii (Mill. Arg.) Hemsl.); leaves opposite (whor sy a dry follicle, seeds apically comose 6 (1). Erect herbaceous perennial to 0.9 m high; flowers small, long, campanulate-tubular; coroll pollen in tetrads; northern Mex 16. Shrubs Alstonia ed in Nerivm), never alternate; —12 mm a with internal scales at base - tube; Apocynum or lianas, Sea ee herbs (Telosiphonia), flowers pically large and showy, 10-70 mm long (3—9 mm in Echites subg Psendechites, Forsteronia and Thenardia), funnelform or salverform corolla without internal scales at base of tube; pollen granular; through out Mexico 17 (16). Pistil head pentagonal (Fig. 1b); leaves with 2—4 colleters at apex of petiole above, bases cordate (Fig. 2) or rounded; “Mandevilla” complex 18 (17). Anthers with pubescent en filamentous appendages; fila- ments long and pronounced; leaves ith domatia in axils of veins beneath; calyx large and s showy, ae burgundy margins Tintinabularia mortonii Woodson 18. Anthers without apical appendages; filaments reduced essentially sessile; leaves without domatia in axils of veins be low; calyx small and gre ee 19 (18). Inflorescence nad ee (Fig. 3b); leaf base rounded echites trifida (Jacq.) Mill. Arg. 19. Inflorescence simple and een Fig. 3a), or reduced to a oe flower (Telosiphonia), leaf base cordate (Fig. 2) or rounc 20 (19), Lianas or suffrutescent herbs; leaves with (subg. Exothostemon) or without (subg.Mandevilla) glands along midrib of the upper surface; inflorescences many ered; flowers diurnal, red, yellow, or white ........... Mandevilla . Suffrutescent herbs, never twining; leaves without glands along midrib of the upper surface; inflorescences reduced to a solitary flo bo ~ Cc wer; flowers vespertine, ICE 222.Hivecis Telosiphonia 17. Pistil head fusiform (Fig. 1a); leaves without colleters at apex of peti- WILLIAMS, Mexican genera of Apocynaceae 20: b. Fic. 1. Representative pistil heads: A. “Echites” complex. B. “Mandevilla” complex. ole above, or if colleters present then solitary and leaves with domatia in the axils of veins below (Forsteronia), bases rounded, “Echites” complex 21 (17). Anthers with apical Gianientons appendages 4—5 mm long; leaves opposite or whorle 22 22 (21). Anther appendages pubescent; shrubs; corolla with petaloid appendages within; leaves whorled (occasionally opposite); seed covered by short hairs, margins pubescent; cultivatec erium oleander L. 22. Anther appendages glabrous; lianas; corolla without peta- loid appendages; leaves opposite; seed glabrous, coma ros- trate; native and occasionally cultivated ocnseeens Pentalinon andrieuxii Muell. Arg.) Hansen & Wunderlin 21. Anthers without apical filamentous appendages; leaves opposite 2: 23 (21). Sepals ovate, 3-20 mm long, 2-10 mm wide (Fig. 4b,c,d) ie) aN 4 (23). Plants yellowish-villous or glabrous; corolla sal- verform; anthers exserted or included; corolla mouth with a thickened annulus; petioles subtended by pecti- nate glands Prestonia Sipa 17(1) NOG Ses EX t> pmol Ci -s * a oot ayo 2 Fic. 2. Representative leaf of “Mandevilla” complex, showing subcordate base and position of colleters. Bar equals 5 mm 24. Plants glabrous; corol a funnelform; anthers included; corolla mouth not thickened; petioles without pectinate glands 25 25 (24). Inflorescence of 1-4 flowers; sepals not imbri- cate (Fig. 4c); calycine colleters absent; anther body glabrous, the tip pubescent ............0...... Rhabdadenia biflora (Jacq.) Mull. Arg. 25. Inflorescence of 5—10 flowers; sepals imbricate (Fig. Ac jon ); calycine colleters present, alternate the sepals; anther body pubescent, the tip glabrous ............ Odontadenia caudigera Woodson 23. Sepals triangular to narrowly triangular, 0.5—5.0 mm long, 0.5-1.0 mm wide (Fig. 4a) 26 26 (23). Corolla 3-5 mm long, tube 0.5—2.0 mm long; an- ther tip exserted 27 27 (26). Inflorescence thyrsiform, leaves inconspicuously glandular at base of midrib above, domatia in axils of veins beneath; colleters alternate calyx lobes; fol- licles free Forsteronia 27. Inflorescence a subumbell . | lism dvlaw WCliale Cyl 1, S Cy above and without domatia below; colleters oppo- site calyx lobes; follicles fused throughout their length Thenardia WILLIAMS, Mexican genera of Apocynaceae 205 26. Corolla 7-65 mm long, tube 7-40 mm long (4-5 mm in Fernaldia ee Woodson); anthers exserted (Lanbertia) or i d 28 28 (26). eres een corolla tube twisted, mouth with a thickened annulus; sepals without colleters; ovary densely puberscent Laubertia contorta Mart. & Gal.) Woodson ( . Anthers included; corolla tube straight or twisted, 28 mouth not thickened; sepals with a solitary ay ae olleter; ovary glabrou 29 9 (28). Corolla Peanelor (Fig. oo 35-50 mm long, tube straight; corolla lobes villous ........... Fernaldia 29. Corolla salverform (Fig. 5a), 40-65 mm long (subg. Echites) or 7-9 mm long (subg. be: echites), tube straight (twisted in E. wmbellata Jacq.); corolla lobes glabrous (villous in E. woodsoniana Monac.) Echites ALPHABETICAL LISTING OF THE NEW WORLD MEXICAN GENERA OF APOCYNACEAE Allamanda L., Mant. 214. 1771. TYPE SPECIES: Allaah ieee LL. Represented by one species in Mexico. Fallen, M.E. 1985. The es development and systematic position of A//amanda eae eae Amer. 2:572-579. sa M.andG.J.Sh eperd: a Uma revisao do género Al/lamanda L. (Apocynaceae). ee Bot. 9:125—-149. {illus. and maps} — eben is native to South America and is represented in Mexico by the culti- ed species A. cathartica L. cae R. Br., Mem. on Asclepiad. 64. 1810. (nom. cons.). Type Species: Echites scholaris L Tonduzia Pittier, Contr. U.S. Natl. Herb. 12:103. 1908. Represented by two species in Mexico. entry, A. 1983. Adstonia (Apocynaceae): another pal America. Ann. Missouri Bot. Gard. 70:206—207. [reduction of Tonduzia to Alstonia)} Morales, J.F. 1995 Evaluacion del género A/stonia (Apocynaceae en Centro América) Phytologia 78:192-194 Amsonia Walt., Fl. Car. 98. 1788 Type Species: Amsonia tabernaemontana Walt. Represented by four species in Mexico McLaughlin, S.P. 1982. A revision of the southwestern species of Amsonia (Apocynaceae). Ann. Missouri Bot. Gard. 69:336—350. {illus. and maps] Apocynum L., Sp. Pl. 213. 1753. TYPE SPECIES: om cannabinum L. Represented by two species in Mexico. Woodson, R.E., Jr. 1930. Studies in t Apocymoidene( _ special reference to the genus Apocynum). Ann. Missouri Bot. 2. {maps Gard. 17 aeotropical genus in Central he Apocynaceae I. A critical study of the 206 Stipa 17(1) 3cm a Fic. 3. Inflorescence: A. Mandevilla subsagittata (R. & P.) Woodson. B. Mesechites trifida Jacq.) Mill. Arg., scanned from McGregor 930 (TEX), and Contreras 2303 (TEX), respectively. (4 Yh M, mM ay \ \ 1,7 Uy \ re ar (y iow ‘i zi ry! \,\ hen \\\ ‘ \ ff Y\ I Y aA ty, 10 mm \ YX Ys A | vX \ Wy AN h 7 AS . 1 NLS b a sS Fic. 4. Representative sepals: A. Echites yucatanensis Millsp. B. Prestonia mexicana Donn. Arg. D. Odontadenia caudigera Woodson. Smith. C. Rhabdadenta biflora (Jacq.) Mill. Aspidosperma C. Martius & Zucc., Flora 7(1) (Beil.) 135. 1824 (nom. 54): con TYPE SPECIES: A ee ma tomentosum C. Martius & Zucc. Cufodontia Woodson, t. Sist. 10:38. 1934. Represented by two species in Mede Marcondes-Ferreira, W/. 1989. Apidaporma Martius & Zucc. nom. cons. oo idos Taxonomicos. Doctoral Thesis, Universidade Estadual de Campina Woodson, R.E., Jr. 1951. Studies in the ome VII. An interim revision of the genus Aspidosperma C. Martius & Zucc. Ann. Missouri Bot. Gard. 38:119—206. [re- duction of Cufodontia; illus.} Cameraria L., Sp. Pl. 210. 1753. WiLLiAMS, Mexican genera of Apocynaceae 207 c d Fic. 5. Corolla and bud: A-B. Echites oereiaele Millsp. C-D. Fernaldia pandurata (A. DC.) Woodson. Scanned from Lundell 7455 (TEX), and Williams 95-90 (TEX), respec- tively. Note: After aes the flowers turn a ow (as seen here), however, in the field the they are in fact w a Type Species: Cameraria latifolia L. Represented by one species in Mexico. Woodson, R.E., Jr. 1938. is In: N. . —o* W.A. Murrill, and J.H. Barnhart, eds. Apocynaceae. N. Amer. Fl. 29: 1 20— Note: Cameraria was aoa a om Mexico at ae time of its last revision (Woodson, cited above). It has ees been ae in the Yucatan peninsula, where repre- sented by C. /atifolia L. Representative specimens. Campeche: 4 km W of Conhuas, 98 km along road to Escarcega, 18 Aug 1983, Cabrera 5353 (MEXU, MO). Tabasco: 2 km of La N-25, along La W-0 on the road to N-20, near Un Drene, Balancan, 13 Oct 1975, Menendez 296 (MEXU). Quintana Roo: Mpio. Carrillo Puerto, Sian Ka’an Biosphere Reserve, 15—20 km N of Carrillo Puerto (19° 50' N; 87° 40' W), 2 Nov 1984, Nez// 5752 (MO). Veracruz: Orizaba, 17 Aug 1940, Miranda 625 (MEXU). YUCATAN: 15 kn NW of Humucma, along road Merida-Sisal, 20 Jul 1985, Cabrera 9097 (MO). Cascavela Raf. = Thevetia Cufodontia Woodson = Aspidosperma Echites P. Browne, Civ. Nat. Hist. Jamaica 182. 1756. Type Species: Echites umbellata Jacq. Represented by five species in Mexico. Monachino, J. 1959. A new Echites from Mexico. Bull. Torrey Bot. Club 86:245—247. {description of Echites woodsoniana Monac.; illustration; Michoacan} Morales, J.F. 1996. A reevaluaion of the genus Echites (Apocynaceae). Brittonia (in press). Woodson, R.E., Jr. 1936. Studies in the Apocynaceae IV. Ann. Missouri Bot. Gard. 23:169-438. [Echites 217-252] Fernaldia Woodson, Ann. Missouri Bot. Gard. 19:48. 1932. Type Species: Echites pandurata A. DC. 208 Sipa 17(1) Represented by two species in Mexico. Morton, J. F., Alvarez, E., and Quianonez, C. 1990. Loroco, Fernaldia pandurata (A. oe \ Woodson (Apocynaceae): a popular edible flower of Central America. Econ. 4:301— asdon R. E. er 1932. New or eae noteworthy Apocynaceae of tropical America . Ann. Missouri Bot. Gard. 19:48—49. [erection of Fernaldia; illustration} Seats LO, mee es 1a the Rey IV. Ann. Missouri Bot. Gard. 23:169- 438. [Fernald ae 263] . 1939. New or otherwise note ey — of eee America . Ann. Missouri Bot. Gard. 26:96-97. f Fernaldia asper is Woodson; Guerrero} Note: Fernaldia is often confused with Echites subgenus Echites, it can be readily distin- guished from subg. Echites by its funnelform corolla (Fig. 5c) Forsteronia G. Mey., Prim. Fl. Esseq. 133. 1818. Type Species: Echites spicata Jacq. Represented by four species in Mexico. Hansen, B.F. 1985. A monographic revision of Forsteronia (Apocynaceae). Doctoral The- sis. University of South Florida. {illus. and maps} Haplophyton A. DC., Prodr. 8:412. 1844. Type SPECIES: Haplophyion “niin A. DC. Represented by two spe Nelson, C. god Haplophyto ton ae A. DC. versus Haplophyton cinereum (A. Rich.) Woodson (Apocynaceae). Fontqueria 40:49—52. Williams, J.K. 1995. Miscellaneous notes on Haplophyton (Apocynaceae: Plumerieae: Ai: iplophyeinae) Sida 16:469-475. ps} Laubertia A. DC., Prodr. 8:486. ren TYPE SPECIES: Liberte boissierii A. DC Streptotrachelus Greenman, Proc. Amer. Acad. Arts 32:298. 1897. Represented by one species in Mexi Woodson, R.E., Jr. 1936. ee: in eae Apocynaceae IV. Ann. Missouri Bot. Gard. 23:169-438. [Laubertia 370-375} Macrosiphonia Mill. Arg. subg. Telosiphonia Woodson = Telosiphonia £ Mandevilla Lindl., Edwards’s Bot. Reg. 26:t. 7. 1840. TYPE SPECIES: ie suaveolens Lindl. Represented by approximately Le in Mex sels C.L. 1942. Studies of Acs erice Se ines II. Contr. Univ. Mich. Herb. 47. - ae of Mandevilla a Lundell; Chiapas Wonks ., Jr. 1933. Studies in the Apocynaceae TV. Ann. Missouri Bot. Gard. 0:605— io [Mandev le 645-777] me Mandevilla is often confused with Echites, it can be readily distinguished from ichites by its colleters along the petiole apex (Fig. 2) and its pentagonal pistil head (Fig. 1b). Mesechites Miill. Arg., Fl. Bras. 6:150. 1860. DC Type Species: Mesechites mansoana A. eae by one species in Mex odson, R.E 1933. ae in aie Apocynaceae IV. Ann. Missourt Bot. Gard. on 605-790. [Mesechites 629-645} *3 WILLIAMS, Mexican genera of Apocynaceae 209 Note: Mesechites is often confused with Mandevilla, it can be readily distinguished from Mandevilla by its branching inflorescence (Fig. 3b) Odontadenia Benth., J. Bot. Goma 3:242. 1841. Type Species: Odontadenia oe Bentl ees me one species in Mex Woodson fie L955. 20 udies i in or Apocynaceae TV. Ann. Missouri Bot. Gard. 22: 52 on {Odontadenia 270-306 2192 tudies in the een ceae IV. Ann. Missouri Bor. Gard. 23:169- 438. (Odean 384-386, description of Odontadenia caudigera Woodson] Note: Odontadenia was unknown from Mexico at the time of its last revision (Woodson, cited above). It has recently been collected in the state of Chiapas, Mexico, where represented by the eee O. caudigera Woodson. Representative specimens. Chiapas: ge Ocosingo, in Ejdo El Pird, 15 km E of Chajul, n che road to Boca ie antum, 16 Apr 1986, Martinez 18212 (MEXU). Pentalinon Voigt, Hortus Suburb, Calcut. 523. 1845. Type Species: Echites erie Urechites Mill. Arg., Bot. Zei ene 18: 22. 1860. Represented — one oe in co. Hansen, B.F., and R.P. earns 196 Pentalinon Voigt, an earlier name for Urechites Mill. Arg. (Apocynaceae). Taxon 35:166-168. Peschiera A. DC. = Tabernaemontana Plumeria L., Sp. Pl. 209. 1753. Type SPECIES: Plumeria rubra L. Represented by apt ae two species in Mexic Bandyopadhyaya, M., and P. C. Dutta. 1986. ee anatomy of different species of Plumeria. Bull. Bot. Soc. Bengal 40:59- 6 Woodson, R.E., Jr. 1938. An Evaluation of the Genera Plumeria L. and Himatanthus Willd. Ann. Missouri Bot. Gard. 25:189—22 Plumeriopsis Rusby & Woodson = Thevetia Prestonia R. Br., on Asclepiad. 58. 1810 (nom. cons.). Type SPECIES: Prestonia tomentosa R. Br. Represented He approximately three species in sens Gentry, A. 1983. A new combination for a problematic Central American ie aia n. ee Bot. Gard. 70:205—206. a of Echites woodsontana Monac Prestonia Williams, L.O. 1968. Tropical American Plants, [X. Fieldiana, Bot. 31 -402—403. [de scription of Prewonia grandiflora L. O. Wms.; Chiapas] Woodson, R.E., Jr. 1936. Studies in the Apocynaceae IV. Ann. Missouri Bot. Gard. 23:169-438. [Prestonia 276-367] Note: Prestonia is subdivided into four sections. Two sections (Coa/itae and Acutifoliae) are not represented in Mexico, they are characterized by having small and incon- spicuous sepals similar to those of Echites (Fig. 4a). The other two sections (Annulares and Tomentosae) are represented in Mexico by one or two species each, they are charac- terized by large foliaceous sepals (Fig. 4b). Gentry (cited above) transfered Echites woodsoniana Monac. to Prestonia, relating it to members of sect. Coalitae. Because of I fusing nature of generic delimitations in the Apocynaceae, and the lack of a corolla annulus in E.woodsoniana, he was not confident of his transfer. 1am inclined to 210 Stipa 17(1) maintain the species in Echites given that at present no other Mexican species of Prestonia ave inconspicuous sepals and because of the taxon’s lack of a corolla annulus and pectinate glands at the base of the petiole. Rauvolfia L., Sp. Pl. 208. 1753. Type Species: Raavolfia tetraphylla L. Ranwolfia Gleditsch, Syst. 212 2. 1764 (orth. var.). Represe ie - two species in Mexico. Rao, A.S. | _ A revision of Rauvolfia with particular reference to the American spe- cies. a ee Bot. Gard. 43:253—355. [illus. and maps} Ranwolfia Gleditsch = Rauvolfia Rhabdadenia Miill. Arg., FI. ae % [7 os. 1860. Tyee Species: Rhabdadenia pohlit Mill. Represented by one species in Mexico. Woodson, R.E., Jr. 1936. SS in ag Apocynaceae IV. Ann. Missour: Bot. Gard. 23:169-438. [Rhali re 205— Stemmadenia Benth., Bot. Gan Sulphur 124. 1845. TyPe SPECIES: Senin eae elabra Benth. Represented by six species in Mexico. Allorge, L. 1985. Monographie des Apocynacées-Tabernaemontanotdées Américaines, Morphologies, Systématique, Chimio-taxonomie. Mém. Mus. Natn. Hist. Nat. Paris, vs. 30:1—216. {1 rae Leeuwenberg, A.J.M. 1994. Index of exsiccatae of Tabernaemontana, the New World species, and See Wi a Netherlands: Dept. of Picne een Wageningen eis ultural Univer i eeeeeeeEEES” _ A revision of 7 “se rnaemontana. Iwo. The New World species and ee ree Royal Botanic Gardens, Kew, Richmond. [illus. and maps] Streptotrachelus Greenman = Laubertia Tabernaemontana L., Sp. Pl. 210. 1753. Type Sprcies: Tabernaemontana citrifolia L Peschiera A. DC., Prodr. 8:360. 1844 Freee by three species in Mexico Allorge, L. 1985. Monographie des Apocynacées-Tabernaemontanotdées Américaines, Morphol, Sy BeSin igs Chimio-taxonomie. Mém. Mus. Natn. Hist. Nat. Paris —216. a Lecumenbery, A.J.N 1994, Index of exsiccatae of ‘ rnaemontana, the New World secies, and ie ches Wageningen, Netherlands: Dept. of Plant Taxonomy, nea Agricultural oo 1994. A Revision of Tahernaemon fan vo. The New World oo Coppa. The Royal Botanic Gardens, Kew, ene {illus. and map Telosiphonia (Woodson) Henrickson, Aliso 14:179—195, 1995 [1 oe Type Species: Echites hypolenca Benth Macrosiphonia Mill. Arg. subg. Telosiphonia Woodson, Ann. Missouri Bot. Gard. 20:778 1933 Represented by six species in Mexico. Henrickson, J. 1995 [1996]. Studies in Macrosiphonia (Apocynaceae): Generic recogni- tion of Telosiphonia. Aliso 14:179-195. {transfer of North American Macrosiphonia to Telosiphonia; illus. and maps] WILLIAMS, Mexican genera of Apocynaceae 211 Note: It has been recently suggested by Henrickson (cited above) that ee is at present polyphyletic: the North American species (subg. Te/osiphonia) having no relation to the South American ones (subg. Macrosiphonia). Henrickson has ene the transfer of the North American species to the newly erected genus Te/osiphonia. Thenardia Kunth, Nova Gen. Sp. 3:209. 1819. TYPE SPECIES: enna Horibunda Kunth Represented by three species in Mexico Woodson, R.E., Jr. 1936. Studies in ihe Apocynaceae [V. Ann. Missouri Bot. Gard. 23:169-438. [Thenardia 271-276] Williams, J.K. 1995. A new species of Thenardia with notes on the genus. Brittonia 47:403—407. {description of Thenardia chiapensis J. K. Williams; illus.; Chiapas} Note: Thenardia is the only apocynaceous genus endemic to 5 MERIC: Currently, there ve species recognized in Thenardia, however, (Williams, in prep.) suggests that two of these, T. gonolobies Woodson and T. ae Woodson, should be reduced to synonymy under T. ga/eottiana Baillon and T. floribunda Kunth, respec- tive Thevetia L., Opera Varia 212. 1758 (nom. cons.). TYPE SPECIES: ie ahouai L Cascavela Raf., Sylva Tell. 162. 1838. Plumer topsis nes & Woodson, Ann. Missouri Bot. Gard. 24:11. 1937. Represented by six species in Mexic Costa, E. de L. and C.G. Costa. 1980. Consens sobre o fruto . la ahouat (L.) Rusby & Woodson (Apocynaceae). Rodriguésia 32(55): 65-— Gensel, W.H. 1969 A revision of the genus Thevetia oe Masters Thesis. University of Connecticut. [illus. and maps Williams, J.K. 1996. A new combination in Thevetia (Apocynaceae). Sida 17:185-190. {elevation of T. oe (Pers.) K. Schum. var. pinifolia Standl. & Steyerm. to species; maps; Michoaca Woodson, R.E., Jr. 1937. New or otherwise noteworthy A pocynaceae of tropical America ; n. Missouri Bot. Gard. 24:11—12. [erection of P/wmeriopsis] Note: wn. Rusby and Woodson (Woodson 1937, cited above) de scribed the mono- S Plumertopsis (P. abonai (L.) Rusby & Woodson) they distinguished it from 7 hevetia of its baccate fruits (vs. drupaceous) and salverform corollas with reflexed iob es ( vs. funnelform and spreading or erect). They reported that both genera had “brilliant red” mesocarps. My observations in the field, however, indicate that ta has red fruits and that the fruits of Thevetia are green when immature and when mature. P/wmertopsis shares all other morphological ities stigma and oad features with Thevetia and I treat it is a synonym of T Tintinnabularia Woodson, Ann. Missouri Bot. Gard. ee Type SPECIES: Tintinnabularia mortonti “/oodson sea Y one species in Mexico. son, ., Jr. 1936. Studies in the Apocynaceae IV. Ann. Missouri Bot. Gard. 169— - . {Tintinnabularia 387-391; erection of a illustration] Note: Tintinnabularia is an extremely beautiful and elusive monotypic — which has been rarely collected. I took a field trip (Summer 1995) to the two most re -ollec- tion sites from Chiapas, Mexico (see below) in order to collect fruits, as are at present unknown. However, as a result of the area being converted into grazing land, the plant was not found. The native vegetation was replaced by weedy forbs and other adventive weeds. Sipa 17(1) vecimens. Chiapas: Municipio of Solosuchiapa, steeped walled canyon along a fast moving stream with seasonal Evergreen Forest, 2—4 km below Ixhuatan along road to Pichucalco, 8 May 1973, Breedlove 34900 (TEX); Municipio of La Trinitaria, slopes with Monrane Rain baer: E of Laguna Tzikaw, Monte Bello Na- Park, 1300 m, 13 May 1973, Breedlove 35191 (TEX). Representative s| tional Tonduzia Pittier = Alstonia Urechites Mill. Arg. = Pentalinon Vallesia Ruiz & Pavon, Fl. Peruv. Prodr. 28. 1794. Type Species: Rauvolfia Hebe Cav Represented by approximately a species in Mexico Woodson, R.E., Jr. 1938. Vallesia. In: N.L. Britton, W.A. Murrill, and J.H. Barnhart, eds. Apocynaceae. N. Amer. Fl. 29:138-141 Note: Woodson (cited above) recognized six species of Vallesia in Mexico. Preliminary research suggests, however, that V. en Woodson and V. conzatti7 Standley should reducing the number of species in be treated as synonyms of V. /anciniata Brand., Mexico to four. Further examination is ee, ae r, before a definitive treatment can be resolved. ADDITIONAL GENUS Trachelospermum Lemaire, Jard. Fleur. 1:pl. 61. 1851. Type Species: Rhynchospermum jasminoides Lindl. Note: eel ‘mum jasminoides (Lindl.) Lemaire, a native of Asia, is occasionally cul- tivated in Mexico. The genus ts een by one species in North America, 7. difforme ( Walt.) A. Gray, but this species has not been reported from Mexico. Trachelospermum is distinguished ia its scandent habit, opposite leaves, fragrant white salverform Be as ioe mm long), and connivent anthers. The genus has not become naturalized in 0. Below is the only specimen from Mexico that I have seen in any of the ene ae Representative specimens. Sonora: Hermosilo, cultivated, 6 (MEXU) Author’s note: Scanned images of representative herbarium specimens of selected Apocynaceae species in Mexico, with distribution maps, are avail able on the World Wide Web. These images can be found on the home page of the Plant Resources Center at the University of Texas (hetp:// www.utexas.edu/ftp/depts/pre/). May 1922, Ma/sallago 503 ACKNOWLEDGMENTS Tam obliged to Guy Nesom for providing me with bibliographical data and inspiring me to undertake this project. The illus. were drawn by Nancy Webber. Cooperation from the following institutions, BRIT, F, LL, MEXU, MO, TEX, and US, was also much appreciated. I also thank Billie Turner, Carol Todzia, Pierro Delprete, Mary Endress, Kurt Potgieter (Aspidosperma), and James Zurruchi for providing me with insightful comments. Finally, I acknowledge Jon Plum and Kathy Gould for their assistance in the field. Wittiams, Mexican genera of Apocynaceae REFERENCES Acosta, O.C. 1984. La familia Apocynaceae en el estado de Quintana Roo, Mexico. Mas- hesis. — Nacional Aut6noma de Mexico. [illus. and maps] DE CANDOLLE, A.L. 1 Apocynaceae. In: A.P. de ey agi een naturalis regni aa. a Masson & cie, Paris. 8:412— Gentry, A.H. 1993. A field guide to the families and genera sof woody plants of northwest South America (Colombia, Ecuador, Peru) with supplementary notes on herbaceous taxa. Conservation International. Washington, D.C. {illus.} Jupp, W.S., R.W. Sanpers, and M.J. DonoGHue. 1994. Angiosperm family pairs: prelimi- nary phylogenetic analysis. Harvard Papers in Botany 5:1—-51. [reduction of the Asclepiadaceae to the Apocynaceae LEEUWENBERG, A.J.M. 1994. Taxa of the Apocynaceae above the genus level. Agric. Univ Wageningen Papers 94-(3):45—60 Rosarti, T. 1989. The genera of suborder Apocynineae (Apocynaceae and Asclepiadaceae) in the southeastern United States, Apocynaceae. J. Arnold Arbor. 70:307—401. [illus.] STANDLEY, P.C. and L.O. WiLLiaMs. 1969 . Apocynaceae. In: Flora of Guatemala. Fieldiana, Bot. 24(8):335—407. [illus.] SrruWE, L., V.A. ALBERT, and B. Bremer. 1994. Cladistics and family level classification of the Gentianales. Cladistics 10:175—206. Woopson, R.E., JR. 1933. Studies in the Apocynaceae IV. The American genera of Echitoideae. han. Missouri Bot. Gard. 20:605—790. 1935. Studies in the er IV. The American genera of Echitoideae (contiaustion). Ann. Missouri Bot. Ga :153-—30 1936. Studies in the eeaee IV. The American genera of Echitoideae 169-438. Regachided). Ann. Missouri Bot. Gard. 23: «1938. Apocynaceae. In: N.L. Britton, W.A. Murrill, and J.H. Barnhart, eds. North American Flora. New York Botanical Garden, New Y vol. 29:103-192. «1: 940. The Apocynaceous Flora of the Vocacan Pe ae In: C.L. Lundell, ed. Botany Y the os Area. XV. Carnegie Inst., Washington Publ. 522:59-102. 214 Stipa 17(1) BOOK REVIEW states, JACK. S. 1990. Mushrooms and Truffles of the Southwest. ISBN 0-8165-1192-6. pbk). The University of Arizona Press, 1230 N Park Ave., Suite 102, Tucson, AZ 85719, Price not given, 234 pp, 156 color photos. This is the first publication that covers such a broad Eange of fungi for southwestern United States according to the author. Coverage also includes the Myxomycetes and li- chens, organisms usually not discussed or illustrated, along with traditional fungal groups such as mushrooms and their relatives. Southwestern geographically includes Arizona, New Mexico, and parts of Colorado, Utah, Nevada, California and northern Mexico. The natural history of mushrooms is emphasized by directing the reader to life zones and plant c ustrated by maps. Keys lead to communities where species can be found. Life zones are il descriptions and color illustrations for 156 of the major mushroom and truffle species. Tl Idi | descriptions for 155 other species. A unique feature of this book is the Mele Ale aCIdItiOllal inclusion of cross-references to assist in species identification. There are eight different cross-references for mushrooms that serve as field guides and one for lichens. There are no references for Myxomycetes. Introductory topics include an excellent discussion of life zones of the Southwest. There is a picture key to the major groups of fungi based on black and white illustrations of fruiting bodies and also a narrative dichotomous key. The Basidiomycetes represent the largest treatment of fungi, covering 129 pages with keys to the families based on macro- scopic characters, for example, spore print colors. Indeed, the book is written more for the amateur using nontechnical, easily understood characters to identify species. The format is very user friendly with color illustrations (4" x 2 1/2") grouped usually two to a page or sometimes one per page with the name of the species and species description on the same page. The habit color photographs are of excellent quality and oriented to show the neces- sary key characters. There is a comments section for each species that discusses the edibil- ity or toxic properties, ecology, and special morphological features. The Ascomycetes are represented by 32 pages and 18 pages are devoted to the tuberlike Ascomycetes and Ba- sidiomycetes (the truffles and false cruffles). This section of the book is especially valuable since this group of fungi is usually absent from field guides. However, a combination of the small size of these underground fungi, lack of bright coloration and distinctive mor- phology, and failure to stand out against a drab background of soil, litter, or man-made surfaces makes it difficult co clearly discern surface features and color. Eight pages are devoted to an assortment of lichens on rocks and branches of trees. There is a glossary of 65 terms defined. The bibliography is not extensive, consisting of the cross-references, six additional field guides, and suggested readings that mentions four books. Anyone inter- ested in the fungi of the southwestern region of the United States will find this book a welcomed addition to their bookshelf.—Harold W. Keller, Research Associate, BRIT. Sipa 17(1): 214. 1996 LAS COMUNIDADES VEGETALES EN LA ISLA SOCORRO, MEXICO JOSE LUIS LEON DE LA LUZ, AURORA BRECEDA and ROCIO CORIA BENET Division de Biologia Terrestre Centro de Investigaciones Bioldgicas del Noroeste, S.C. Apdo. postal 128 La Paz, Baja California Sur, MEXICO 23000 ABSTRACT In this work the different plant communities on Socorro Island are described and delim- ited in a map of vegetation; also, the distribution of the endemic species in the different plant communities is analysed. The identification and mapping of the different plant com- munities was based in four trips to the island which included plant collecting, walk arounds and overviews by airplane, aerial photos 1:70,000 were revised as guide. Most of the plant communities and nomeacae used in this work agree with those proposed by earlier ties (prairie, evergreen tropical forest, mixed scrub, Croton descriptions. Six scrubland, ten and coastal goupS) are described with data about their area and dominant species. The eroded area, induced by overgrazing of the domestic eee ee 1869 inhabits the island, is also considered in the mapping, it covers about 15% « a te surface. On 210 km?, Socorro Island counts with 116 species of native eek plants, plus 46 naturalized. RESUMEN En este trabajo se describen y delimitan los diferentes tipos de vegetacién de la isla Socorro, se presenta un mapa de las unidades de — se analizan la distribucién de las especies endémicas y de las formas de vida en los diferentes tipos de vegetaci6n, asi como la riqueza especifica en cada una de éstas. Para la identificacién y cartografiado de las diferentes agrupaciones vegetales se realizaron cuatro viajes a la isla, durante los cuales se hicieron recorridos por tierra y por aire, también se revisaron fotos aéreas a escala 1:70,000. La nomenclatura de los ae de vegetacién aqui presentados son compatibles con los meres por trabajos previos. En total se describen seis tipos de vegetacién (pradera, bosque tropical perennifolio, elon mixto, matorral de Croton, cae y agrupaciones costeras), proporcionandose la lista de las especies mas comunes a cada una de ellas y delimitando su area de distribucién. También se consideran las asics erosionadas, que cubren actualmente alrededor del 15% de la isla, las cuales han sido inducidas por el ir aredaien del borrego doméstico, introducido en 1869. La flora de la isla Socorro (210 km iperficie) esta formada por 116 especies de plantas vasculares nativas y 46 ce ee: recientemente. INTRODUCCION El archipiélago de las Islas Revillagigedo (Islas Socorro, San Benedicto, Roca Partida y Clarion, Fig. 1) posee una posicién geopolitica distintiva. El Stipa 17(1): 215-230. 1996 216 Sipa 17(1) archipiélago extiende el mar patrimonial mexicano en miles de km? en el pacifico occidental, lo que aumenta considerablemente el potencial de los recursos pesqueros dentro de la zona econémica exclusiva. Sin embargo, la importancia de este archipiélago y particularmente de la isla Socorro, la mayor de las cuatro del sistema, radica no sélo en el aspecto poliftico- economico, sino en su interesante biota, que es resultado de largos procesos evolutivos, caracterizados por un marcado aislamiento geografico. Sobre esta isla es posible hacer un historial en cuanto a las colectas de vegetales vasculares. La primera se ubica dentro de la segunda mitad de la década de 1830, a ésta siguieron muchas otras de naturalistas norteamericanos, apoyadas principalmente por la Academia de Ciencias de California, en especial resalca la exploracién de Johnston (1931). Avanzado el presente siglo, se realizan varias expediciones lideradas por bidlogos mexicanos. Entre las mas importantes se encuentran la de la Universidad de Guadalajara (Medina 1957) y la de la Universidad Nacional Auténoma de México (Adem et al. 1960). En esta Ultima el botanico a cargo, el Dr. Faustino Miranda, describio siete agrupaciones vegetales para la isla, siendo éste el primer intento al respecto. Del conjunto de colectores de plantas en la isla, el Dr. Reid V. Moran es quien la ha explorado mas intensivamente, a través de tres expediciones reunio alrededor de 400 ejemplares, hecho que le animé a estructurar la hasta ahora flora mas actualizada (Levin y Moran 1989). Esta obra conté con la colaboraci6n del personal del Centro de Investigaciones Biolégicas del Noroeste (CIBNOR, herbario HCIB), que aporté duplicados de 180 eyemplares, 12 de los cuales correspondieron a nuevos registros. En la presente contribucién se propone una nomenclatura para las unidades de vegetacion, se describen fisonémica y floristicamente los diferentes tipos de vegetacién existentes en la isla, que se muestan en un mapa. Se hace también un anilisis floristico y de formas de vida en relacién con su distribucién en las agrupaciones descritas. MATERIAL Y METODOS a) Descripcion de la Isla Socorro forma parte de las cuatro islas del archipiélago Revillagigedo, en la costa centro-occidental de México, se localiza a 110°59'N y 18°46'W, aproximadamente a 480 km al sur de la peninsula de Baja California y a 490 km al oeste de la costa de Colima (Adem 1960). Esta isla es la mas grande del archipiélago, con una extension aproximada de 210 km? y una _ alcitud maxima de 1050 m en la cima del volcdn Evermann, que establece una fisiografia concéntrica, cuyo eje es dicho volcan. Geoldgicamente se considera una isla oceanica de edad incierta. Bohrson LEON-DE LA Luz, ET AL., Vegetacién de la Isla Socorro, México 217 Océano ey Playa Norte x as Pacifico Playa blanca Cabo Henslow ) Cabo Pearce hin Punta tosca Pista Aerea i eee || 19°48' ——— FY Vegetacién de ambienles costeros SES Matorral mixto 1°00" \ wy SX Matorral de Psiaium Cabo Rule —— Matorral de Croton eee Bosque de Bumelia eo : | Bosque Tropical Perennifolio Bosque de Ficus 3 . Proporcién superficial WG Pastizal A Pradera — ao . 7[[I[I]] Areas erosionadas , 1026 489 3 127 6.1 4 51.3 244 5 3.1 1.6 6 1.3 0.7 7 38.5 18.3 Fic. 1. Distribucién de las 7 comunidades vegetales reconocidas en isla Socorro, Rev. La superficies relativas de cada una se citan en el text. La vegetacién de ambientes consteros bordea el litoral. Las Areas erosionadas contienen restos de la vegetacion primaria yen algunos sectores vegetaci6n adventicia. J 218 Sipa 17(1) (1994) le asigna un origen volcanico de pocos millones de afios. Blazquez (Adem et al. 1960) y Bryan (1949) la ubican en la transici6n mioceno- plioceno (unos 10—12 millones de afios). Johnston (1931) cita que el sustrato subyacente es de origen continental pre-miocénico, ademas de que la batimetria de la region sugiere un origen continental. De acuerdo con los datos climaticos disponibles en la Unica estacién climatol6gica, ubicada al sur de la isla (10 m de altitud), Coria (1994) describe el clima de ese sector como BS,(h)w(1); ésto es, drido, calido, con régimen de lluvias de verano, la precipitaci6n invernal aporta menos del 10% del cotal anual, siendo la oscilacién térmica poco significativa. La temperatura media anual esde 24.6°C, mientras que la precipitacié6n media anual es de 405 mm en ese sector. La zona norte de la isla es sensiblemente mas himeda; las superficies elevadas se encuentran frecuentemente cubiertas por neblina. Es factible que se presenten heladas en el invierno. Por su posicion geografica subtropical, la isla se encuentra en la trayectoria de los ciclones que azotan la costa occidental de México durante el verano. Los suelos de isla Socorro presentan caracteristicas asociadas a materiales igneos y se dividen en tres grandes areas de distribucién: en la zona norte, los suelos son j6venes formados a partir de las coladas de lava de erupciones recientes, en la zona central han derivado de cenizas volcdnicas y en la zona sur son suelos arcillosos y profundos (Maya et al. 1995). b) Agrupaciones floristicas Para la determinacion de los tipos de vegetacién, se realizaron cuatro estancias de trabajo de campo durante los meses de mayo, septiembre y diciembre de 1988 y febrero de 1990. Estas estancias comprendieron en total 26 dias de actividad, durante los cuales se efectuaron recorridos a pie que permitieron colectar intensivamente. La clasificacion de los tipos de vegetacién se hizo con base a un criterio fisondmico. El reconocimiento en avioneta y la observacién de fotografia aérea permitieron delimitar cada agrupaci6n designada; la fotografia usada fue cubierta por INEGI en 1976 escala 1:70,000. Visitas posteriores permitieron verificar y afinar las versiones previas de la delimitacién de cada agrupacion floristica. Cabe aclarar que los recorridos a pie cubrieron una amplia superficie de la porcién sur, pero la vertiente norte atin se encuentra inexplorada. En este trabajo se hace referencia a cada unidad fison6mico-floristica como un “tipo de vegetacion.” En el caso de la evidente dominancia fisonédmica de una 0 pocas especies se han empleado los nombres genéricos de los taxa caracteristicos de esa vegetacion. LEON-DE LA Luz, ET AL., Vegetacién de la Isla Socorro, México 219 RESULTADOS a) Flora De acuerdo con los registros de Levin y Moran (1989), la Isla Socorro cuenta con 116 especies nativas de plantas vasculares y 46 especies introducidas durante el presente siglo. Entre las nativas, se incluyen 5 especies atin no bien determinadas o no descritas, se trata de Opuntia sp., Psidium sp. (aff. P. sartorianum), Passiflora sp., Rubus P y una orquidea terrestre afin al género Habenaria (Levin, com. pers.). De las 116 especies nativas determinadas, 30 de ellas son endémicas a la isla a nivel especifico (27) o infraespecifico (3), lo que representa el 26.7% de la flora insular. Es posible que la determinacion de las 5 pendientes contribuya a elevar este nivel. En el cuadro | se muestra la distribucién en los diferentes tipos de vegetacion de las 30 especies endémicas. La mayoria de las especies endémicas se encuentran en dos o mas comunidades, pero ninguna de ellas se distribuye en todas, mientras que 13 parecen restringirse solo a una de éstas. En el cuadro 2 se muestra el espectro de las diferentes formas de crecimiento de las especies nativas que componen la flora de la isla. Es apreciable que las herbaceas congregan el mayor numero de especies tanto en el grupo de nativas como en las de introduccién reciente. b) Tipos de vegetacion La fisonomia de la vegetacion es en términos generales “tropical”; sin embargo, las diferencias altitudinales, topograficas, climaticas, de exposici6n y de suelo, han permitido el desarrollo de diferentes tipos de vegetaci6n. De acuerdo con los recorridos por la isla y la fotointerpretaci6n, se distinguen seis tipos, mds una superficie considerable gravemente erosionada. A continuacion se describen cada uno de éstos, en la Fig. 1 se delimitan estas mismas comunidades y se anota la superficie relativa. 1. Agrupaciones costeras Esta corresponde a lo que Miranda (Adem et al. 1960) denomina “Agrupacion de haldfitos costeros” y a lo que Levin y Moran (1989) sefialan como “Shore.” Se estima que éstas ocupan una superficie de 0.5 km? lo que representan un 0.2% del area insular. Sus agrupaciones se encuentran distribuidas irregularmente a lo largo de la franja litoral rocosa, acantilados, playas y dunas costeras. Algunas de las especies representativas son: Canavalia rosea (Sw.) D Cyperus howell1i ] Neill et Benedict Ayers Chamaesyce incerta me . Brandegee) Cyperus ligularis Millsp. Heliotropinm eee Le Conocarpus erecta L. Hibiscus pernambucensis Arruda Cressa truxillensits Kunth Hippomane mancinella L. 220 Sipa 17(1) Cuapro 1. Distribucién de las 30 espécies endémicas de Isla Socorro dentro de las comumidades vegetales consideradas. Comunidad Compartidas (17) Restringidas (13) Vegetacion Costera 8 | Matorral de Croton 5 | Pastizal 3 | Matorral Mixto 16 2 Bosque Tropical 13 4 Pradera 11 4 Cuabro 2. Especcro de las formas de crecimiento consideradas en este trabajo para la flora de Isla Socorro, Rev. Numero total % Flora Nativa % Flora Adventicia % de especies Arboles 12 7.40 11 9.46 | 2.17 Arbustos 22 13.59 19 16.36 3) 6.52 Herbaceas 104 64.20 65 56.01 10 86.97 Trepadoras 8 1.94 7 6,02 | 2 Epifitas 6 5.7] 6 5.17 0 0 Hemiuparasitas 2 1.23 2 71 0 0 Rastreras 7 4.32 6 5.17 | ZAy Acuaticas | 0.61 l 0.86 0 0 Total 162 100% 116 100% 4G 100% Ipomoea pes-caprae (L.) Sweet subsp. Scaevola plumieri (L.) Vahl brasiliensis (L.) Ooststr Sida nesogena 1.M. Johnston Jouvea pilosa (C. Presl.) Scribner Sorghastrum nutans (L.) Nash in Small Physalis mimulus Waterf. 2. Matorral de Croton Corresponde al “matorral de Croton” de Miranda (Adem et al. 1960), y al “scrub” de Levin y Moran (1989). El componente dominante es Croton masonit I.M. Johnston, especie que durante la temporada de sequia, febrero-junio, pierde la totalidad de sus hojas. Esta comunidad ocupa alrededor de 12.6 km? (6.1%), encontréndose bien representada en la parte sur de la isla, desde el contorno de la franja costera hasta unos 250 m de altitud, se desarrolla sobre una extensa capa basdltica, con pendiente de leve a moderada. Existen manchones reducidos de esta vegetacién hacia el norte y el oeste de la isla, en la misma franja altitudinal. Entre las especies allf existentes es posible distinguir: Abutilon californicum Benth. Cordia curassavica (Jacqg.) Roemer et Bursera eppinata (Rose) Engl. Schultes LEON-DE LA Luz, ET AL., Vegetacién de la Isla Socorro, México 221 Cynanchum californicum (Benth.) Moran Rhamnus sharpii M. et L.A. Johnston Ipomoea triloba L. Tournefortia har ee Steudel Malvastrum americanum (L.) Torrey in cae ae E. Greene Emory Walther Melochia pyramidata L. Zapoteca ae a H. Hern. subsp. Opuntia sp. socorrensis (ILM. Johnst.) Levin, H. Pavonia hastata Cav. ernandez et Moran 3. Pastizal Corresponde parcialmente a lo que Miranda (Adem et al. 1960) integré en la agrupacion de hal6fitos costeros y Levin y Moran (1989) al “grass- land.” Ocupa una extensi6n de unos 3.1 km? (1.6%), se distribuye en el extremo sur de la isla (Cabo Regla, Fig. 1), en lugares de pendiente moderada y lomerios de escasa elevacion. Si bien el terreno puede considerarse rocoso, ha logrado acumularse gran cantidad de suelo entre las mismas rocas permitiendo asi el establecimiento de estos vegetales. En esta vegetaci6n dominan las gramineas, tales como: Eragrostis tenella (L.) P. Beauv. ex Roemer & Schultes, Heteropogon contortus P. Beauv. ex Roemer & Schultes, Paspalum longum Chase in I.M. Johnston, Rhynchelytrum repens (Willd.) Hubb., y Schizachyrium sanguineum var. sanguineum (Retz.) Alston Cabe sefialar que el sector naval fue establecido permanentemente hacia finales de la década de los afios cincuenta sobre esta agrupacion floristica; en consecuencia, una importante superficie de la misma, ubicada dentro del Area de influencia de las instalaciones militares, ha sido alterada. En décadas pasadas se introdujo el “zacate buffel,” Cenchrus ciliaris L. y C. myosuroides Kunth in Humb. Especies acompanantes son: Amaranthus palmeri S. Watson Sie ais hirta (L.) Millsp. var. Arrta stida adscensionis Waltheria ae myosuroides Kunth in Humb. Ler es ares H. Hern. subsp. Chamaesyce anthonyi (T.S. Brandegee) socorrensis (I.M. Johnst.) Levin, H. Levin Hernandez et Moran 4, Matorral mixto El matorral mixto, ne al “matorral de Dodonaea viscosa Jacq.” de Miranda (Adem et al. 1960) y al “scrub” de Levin y Moran (1989). Esta comunidad ocupa la mayor superficie en la isla, cubre aproximadamente 102.7 km? (48.9%) desde el nivel del mar hasta 950 m de elevacién en algunos sitios; sin embargo, la mayor parte de esta vegetacién se distribuye por las partes baja y media de las zonas centro y norte. El matorral mixto se desarrolla sobre suelos pedregosos con pendientes de moderadas a pronunciadas. Esta comunidad vegetal se compone principalmente de herbaceas perennes, de tallos semi-postrados y estoloniferos; aparecen también algunas especies arbustivas en esta comunidad, como D. viscosa, que en Ciertos sitios son fisondmicamente dominantes, asi como elementos 222 SipA 17(1) achaparrados de especies propiamente arb6reas, como Gwettarda insularis T.S. Brandegee, y Prunus sevotina Ehrh. La altura media de estas especies varia entre 1 m y 2.5 m. El helecho Preridinm caudatum (L.) Maxon es muy abundante y en ocasiones forma densos manchones, puede considerarse como una especie pionera o integrante de las primeras etapas sucesionales, dada su elevada demanda de radiacién solar, alta produccién de estructuras reproductivas (esporas), y requerimientos relativos bajos de calidad edafica. Si bien esta comunidad guarda cierta unidad fisonémica en el drea considerada, su composicion floristica cambia dependiendo de las condiciones topograficas, altitudinales, sucesionales y de exposicién. Algunas especies representativas son: Acalypha umbrosa T.S. Brandegee Galium mexicanum Kunth in Humb. Bidens socorrensis Moran & Levin subsp. mexicanum Brickellia peninsularis T.S. Brandegee var. Hibiscus diversifolius Jacq. amphithalassa Robinson ex I.M. Johnst. Lepechinia hastata (A. Gray) Epling var. Cardtospermum halicacabum LL. socorrensis Moran Caesalpinia bonduc (L.) Roxb. Phaseolus lunatus L. Coreocarpus tnsularis (T S. Brandegee) E.B. Rubus sp. Smith Salvia psendomisella Moran & Levin Enpatorium pacificum Robinson ex I.M. Spermacoce nesiotica (Robinson) Levin Johnsc Triumfetta socorrensis T.S. Brandegee Waltheria indica .. En el mapa de la Fig. 1 se sefiala el matorral de Psidinm (socorrense). Este se encuentra ya casi desaparecido por la erosi6n de su sue consideramos que d james o, los autores e manera natural se encontraba integrado en el matorral mixto (ver mas adelante el apartado de Areas Erosionadas). 5. Bosque Tropical Perennifolio Bajo esta denominacién se incluyen “selva de higueras,” con o sin guayabillo, y “selva de zapotillo-guayabillo” de acuerdo a la clasificacién empleada por Miranda (Adem et al. 1960). Levin y Moran (1989) usan la denominacion “forest.” En su conjunto los bosques ocupan una superficie aproximada de 51.3 km? (24.4%, ver Fig. 1). Las formas bioldgicas caracteristicas de los bosques son arboles, con densidades relativamente bajas pero con un amplio dosel. La dominancia relativa de las especies parece variar en funcion de los niveles altitudinales, de acuerdo con floristicos y estructurales (no presentados en este trabajo) boscosa se puede subdividir en tres agrupaciones. los datos la comunidad Bosquetes de Ficus Entre los 250 y S00 m de altitud, se localizan bosquetes de Ficus cotinifolia Kunth in Humb. (amate, higuera o higuerilla), ocupando mesetas con pendientes leves y suelos aparentemente profundos. Estos bosquetes se distribuyen discontinuamente formando manchones LEON-DE LA Luz, ET AL., Vegetacion de la Isla Socorro, México 223 sobre los diferentes flancos de la isla, ocupando mayor superficie en el sudeste, donde existen evidencias de un fuerte proceso de deterioro debido a la continua pérdida de suelo como efecto del sobrepastoreo del borrego Ficus cotinifolia extiende su dosel hasta en algunos centenares de m7 de cobertura, su densidad es dificil de calcular, ya que es diffcil distinguir a cada individuo por la forma de propagacion vegetativa de la especie. Las otras especies lefiosas presentes en este tipo de bosques, tienen escasa aportacion a la dominancia. Tanto Bumelia socorrensis como Guettarda insularis forman, junto con F. cotinifolia, el estrato arb6reo superior con una altura de 5 a 7 m. El estrato arbéreo bajo (de 2 a 3 m de altura) esté compuesto principalmente por individuos abundantes de Psidium aff. sartorianum (O. Berg.) Niedenzu. El estrato herbdceo de estos bosques es poco conspicuo, quiza debido a la densa cubierta vegetal de los Arboles, la baja incidencia de luz, y al sobrepastoreo por borregos. Sdlo en los margenes de cada bosquete se ubican algunas especies herbaceas y arbustivas, entre las que se encuentran Nicotiana stocktonii T.S. Brandegee, Opuntia sp., y Argemone ochroleuca Sweet, mismas que parecen ser favorecidas por la actividad selectiva del borrego. Bosque de Bumelia-Psidium Este tipo de agrupacion se distribuye en el gradiente altitudinal de 500 a 700 m, sobre laderas y cafiadas del volcan Everman, en sitios de pendientes moderadas. Esta agrupaci6n presenta mayor numero de especies arbéreas que los bosquetes de F. cotinifolia, entre las principales se encuentran: Bumelia socorrensis T.S. Brandegee Psidium socorrense 1.M. eae Guettarda insularis T.S. Brandegee Psidium aff. sartorianum (O. Berg Ilex socorroensis T.S. eee Niedenzu Bajo el dosel arboreo se encuentran algunas especies trepadoras y arbustos decumbentes como: Chiococca alba (L.) A. Hitchce., Vernonia littoralis TS. Brandegee, Forestiera rhamnifolia Griseb., y Zanthoxylum insulare Rose, asi como algunas epifitas como: Cattleya aurantiaca (Bateman) P. N. Don, Peperomia tetraphylla (G. Forster) Hook & Arn., P. socorronis Trel. in I.M. Johnston, Polypodinm polypodioides (L.) Watt var. aciculare Wreath. Bosque de Oreopanax-Ilex Este tipo de agrupaciéon se localiza en la parte noroeste de la isla, en altitudes superiores a los 700 m, es la mas densa de todas y presenta el mayor numero de especies. Oreopanax xalapense (Kunth) Decne. & Planchon e [lex socorroensis parecen ser las especies con mayor dominancia; como especies acompafiantes se encuentran Gwettarda insularis, Bumelia socorrensis y Meliosma nesites 1.M. Johnston, las cuales pueden rebasar los 11 m de altura. La mayoria de los individuos presentan una altura maxima entre 5 y 8 m. Los componentes exhiben coberturas con ramificaci6n densa y abundante; Sipa 17(1) ie) ~ posiblemente ocupan la isolinea que delimita el régimen térmico de otro con mayores fluctuaciones diurnas on relativamente estable, tropical, y/o estacionales, ya que subitamente desaparecen los escasos individuos de Ficus y Psidium, mientras que M. nesites, O. xalapense y Prunus serotina aparecen con dominancia. Estas tltimas especies son propias de algunos bosques mes6filos de montana en la superficie continental del pats La franja altitudinal de 700 a 900 m, parece ser la que mayor humedad recibe gracias a la constante neblina, que al ser condensada sobre los Arboles, facilita que éstos se cubran densamente con Ifquenes. En el sotobosque on aparecen varios helechos como: Adiantopsis radiata (L.) Fée Polypodium alfredii Rosenstock Asplenium sessilifolium Desvaux olystichum muricatum (1...) Fée Asplentum formosum Willd. 6. Pradera Corresponde a la “pradera” de Miranda (Adem et al. 1960) y a la “prairie” de Levin y Moran (1989). El bosque desaparece repentinamente en niveles altitudinales situados entre los 850 y 950 m de altitud, para dar lugar a esta comunidad que se desarrolla sobre suelos pedregosos y rocosos, con pronunciada pendiente. La pradera ocupa la parte central de la isla, sobre el cono del volcan Evermann, con una superficie de 1.3 km? (0.7%). Ademas de la carencia de suelo, otros factores limitantes para el crecimiento de especies lefiosas son el viento fuerte y las temperaturas relativamente bajas que alli inciden. Son especies caracteristicas: Aegopogon solisti Levin Heterotoma cordifolia (Hook. & Arn.) Aristida vaginata A. Hitche. McVaugh Castilleja socorrensis Moran Hypericum eastwoodianum 1.M. Johnston Centanrium a Broome Linaria canadensis (L.) Dum.-Cours. var. C. wigginst Bre texana (Scheele) Pennell Cestrum wacificum' 76. Brandegee Setaria genicnlata (Lam.) P. Beauv. Cleame viscosa L. Spermacoce nesiotica (Robinson) Levin Daucus montanus Humb. & Bonpl. Sporobolus purpurascens (Sw.) W. Ham. Expatorium pacificur Triodanis perfoliata (L.) Nieuwl. var. aie Gnaphalium attenuatum DC. (Ruiz Lopez et Pavén) Bradle Gnaphatinm sphactlatum Kunth in Humb. 7. Superficies Erosionadas La presencia del borrego doméstico en la isla ha ocasionado una grave y acelerada erosion, especialmente en la vertiente sur-centro. Este 6vido fue introducido hacia 1869 (Gentry 1949), su poblacién ha prosperado principalmente en la region sur, y actualmente se calcula una poblacién de 2 mil animales en la mitad sur de la isla (Cardenas et al. 1994 El efecto de sobrepastoreo y pérdida de suelo, se va ere en los ultimos 30 afios, ya que Miranda (Adem et al. 1960) en 1958, advierte el LEON-DE LA Luz, ET AL., Wegetacién de la Isla Socorro, México 225 efecto del borrego sobre la vegetaci6n, particularmente en las areas correspondientes a la comunidad que denomin6é como “matorral de guayabillo” (Psidium socorrense). Esta especie endémica ha disminuido considerablemente sus poblaciones naturales, ya que ocupé las escasas areas planas del sur, este y oeste de la isla, las mismas que actualmente son deterioradas por las manadas de borrego, la accién edlica y la erosi6n pluvial. En la actualidad cerca de 38.5 km? (18.3%) se encuentran severamente afectados y solamente quedan dos superficies reducidas del matorral de guayabillo, que cubren alrededor de 4.5 km? (Fig. 1). Las superficies erosionadas mantienen una flora muy pobre en cuanto a individuos y ntimero de especies, las fases mas graves se localizan sobre pendientes fuertes, en donde las carcavas proporcionan un panorama impresionante en el paisaje. Sobre algunas mesetas erosionadas, ubicadas entre los 260 y 400 m de altitud, se desarrolla una comunidad secundaria, originada a partir de la desaparicién de superficies posiblemente correspondientes a bosquetes de Ficus cotinifolia, Psidium sp., y Guettarda insularis y del matorral deciduo. Sobre estas mesetas se desarrollan algunos arbustos y especies anuales, éstas iltimas aparecen después de la temporada de Iluvias. La especie con mayor densidad de individuos es Mitracarpus hirtus (L.) DC., que puede considerarse la invasora mas agresiva de la isla, pues a pesar de su introducci6n relativamente reciente ingreso (Levin y Moran 1989) se encuentra presente en practicamente todas las comunidades. Otras especies presentes en estos sitios son: Aristida adscensionis L. Corchorus aestuans L. Aristida vaginata A. Witch. une dactylon (L.) Pers. Boerhavia coccinea Miller Dactyloctenium aegyptinm (L.) Willd. Cenchrus echinatus L. Desmodium procumbens (Miller) A. Hitche. Cenchrus myosuroides Kunth in Humb. Eragrostis pectinacea (Michaux) Nees Chamaesyce anthony? (T.S. Brandegee) Malvastrum woman (L.) Garcke evin Paspalum long Chamaesyce hirta (1..) Millsp. var. hirta Rhynchely les Sees Willd. ) Hubb. Chamaesyce hyssoptfolia (L.) Small Verbena sphaerocarpa Perry En las cdrcavas profundas que surcan pendientes moderadas y pronunciadas, originadas por aguas broncas provenientes de mayores altitudes, se ponen al descubierto suelos rojos, volcdnicos, arcillosos. En la mayor parte de estas zonas no existe vegetacién; sin embargo, en algunos sitios logran instalarse unas pocas especies en estas condiciones, correspondiendo a malezas de introducci6n reciente a la isla como: Argemone ochrolenca Sweet. var. ochroleuca Portulaca oleracea Ii, Cyperus rotundus L. Salvia riparia H.BK Glinus radiatus (Ruiz et Pav6n) Rohrb. Sida Hee Ls Mollugo verticillata L. 226 Sipa 17(1) c) Especies de amplia distribucion Es posible encontrar ciertas especies en varios tipos de vegetacién, por lo que aparentan ser taxa de amplia tolerancia ambiental. A excepcién de Mitracarpus hirtus, ninguna de éstas Ilega a dominar en una superficie dada y todas son especies herbaceas. A continuaci6n se listan algunas de estas especies, indicandose si son nativas o naturalizadas (1), o bien si son de introduccion reciente (2), ésto es, si han aparecido en las Ultimas colectas. Aspleninm formosum Willd. Willd. (1) Eragrostis ciliaris (L.) R. Br. (2) Bulbostylis nesiotica 1.M. Johnsc.) Fern. (1) Matracarpus birtus (L.) DC.(2) Cyperus duripes 1.M. Johnsct.(1) Schizachyrium sanguineum (Retz) Alston Chamaesyce anthony: (T.S. Brandegee) var. sanguinenm (1) Levin (1) Sorghastrum nutans (L.) Nash (1) Cheilanthes peninsularis Maxon vat. Verbena sphaerocarpa Perry (1) msularis Weath (1 Waltheria indica 1. (1) Elytraria imbricata (Vahl.) Pers. (1) d) Distribucion de la fora por comunidad El cuadro 3 muestra la riqueza floristica de cada una de las comunidades vegetales consideradas (el matorral de Pszdivm senalado en el mapa esta contenido aqui en el matorral mixto). Si bien es posible sefalar que el matorral mixto, el bosque y las areas erosionadas presentan una relativa “riqueza” de especies, la abundancia de cada taxon podria mostrar otro panorama. De esta manera, las areas erosionadas las ocupan especies herbaceas (anuales y perennes) de abundancia local y temporal. Otro aspecto que llama la atencién es que practicamente todas las especies de introducci6n reciente (42/46) se ubican en las areas alteradas (areas erosionadas + pastizales). DISCUSION En el cuadro 4 se hace una comparaci6n de las diferentes clasificaciones de la vegetacion de la Isla Socorro. En este cuadro se advierte que la clasificaci6n propuesta por Levin y Moran (1989) es mds general que las otras dos, no hace diferencia en los subtipos de matorrales y bosques. La clasificaci6n propuesta por Miranda (Adem et al. 1960) comprende siete tipos de vegetacion, en el —_— a describe el matorral de guayabillo entre los 250 y 400 m, sobre suelos rojos, también advierte la presencia del borrego sobre esta comunidad vegetal. En la actualidad este matorral se encuentra fuertemente afectado ya que ocupaba las Areas que hoy presentan la erosion mas severa, quedando solamente algunos manchones de esta comunidad. En la clasificaci6n de Miranda (Adem et al. 1960) no se contempla e pastiza — como un tipo de vegetacion, en su noe: menciona solamente el establecimiento de especies herbaceas en los claros del matorral de Croton, pero cabe sefalar que en el momento de esta observacién el impacto de la LEON-DE LA Luz, ET AL., Vegetacién de la Isla Socorro, México 227 Cuapro 3. Riqueza especifica por comunidad vegetal en isla socorro, Rev. (flora nativa + adventicia). Agrupacién taxa Matorral mixto (incluye matorral de Psidium) 53 Matorral de Croton 39 Bosque tropical perenifolio (incluye bosquetes de Ficus) 50 Pastizal 49 Agrupaciones costeras 30 Pradera 35 Superficies erosionadas 48 Cuapro 4. Sinopsis de las clasificaciones de la vegetacion empleadas para Isla Socorro, Rev. Miranda (1960) Levin y Moran (1989) Esta propuesta Agrupaciones de hald Shore Agrupaciones costeras ficos costeros (en parte) Matorral de Croton Scrub Matorral de Croton Matorral de Guayabillo Matorral mixto (Matorral de Ps1dinm casi desaparecido) Selva de amates con o sin guayavillo Forest Bosque tropical perennifolio (Bosque de T7CUS Selva zapotillo-guayabillo Forest Bosque tropical perennifolio (Bosque « ‘ le Bumelia-Psidium y Bosque de Oreopanax-Ilex) Matorral de Dodonea Scrub Matorral mixto Pradera Summit Pradera Agrupaciones de halo Grassland Pastizal (agrupacion fitos costeros (en parte) secundaria) _ — Superficies erosionadas base naval (Cabo Regla) debié ser apenas incipiente, en la actualidad este tipo de vegetacién se ha desarrollado, principalmente, como producto de las perturbaciones antropogénicas, que convierten partes del matorral de Croton en pastizales. En la clasificaci6n propuesta en este trabajo, se propone el término “matorral mixto” obedeciendo a la heterogeneidad en la composici6n florfstica de esta unidad. Un ajuste a la clasificaci6n de Miranda (Adem et al. 1960) es la subdivisi6n del Bosque Tropical Perenifolio, donde aquel autor propuso un tipo caracterizado por Me/zosma y Oreopanax, mientras que en el presente trabajo se consideré como una subunidad, dado que estas especies no se apartan de la fisonomia propiamente boscosa. 228 Sipa 17(1) La nomenclatura propuesta en este trabajo, no se corresponde con los nombres empleados para designar a los grandes tipos de vegetacié6n en México (Miranda y Hernandez X 1963; Rzedowski y McVaugh 1966; Rzedowski 1978), ya que son escalas geograficas dispares y en este caso se considera una descripci6n de la vegetacion en escala local. Los resultados de este trabajo, asf como lo sefialado en estudios anteriores muestran que proporcionalmente a la superficie de la Isla Socorro (210 km’), existe cierta diversidad de tipos de vegetacién que corresponden a diferentes factores ambientales. El gradiente altitudinal y la exposicién de la isla influyen en cambios climaticos que permiten el establecimiento de diferentes comunidades vegetales a lo largo del gradiente altitudinal. Son también notorios los variados tipos de suelos presentes en esta pequefia isla (Maya et al. 1995) como producto de la intensa actividad volcanica de la region. Esta combinacion de factores permiten una relativa diversidad de ambientes y el establecimiento de diferentes tipos de vegetaci6n. La relativa diversidad de comunidades vegetales en la isla contrasta con lo planteado por Levin y Moran (1989) en torno a la flora; de acuerdo con estos autores el endemismo de la flora puede catalogarse como “moderado” (26.5% de la flora nativa, comparado con las islas Hawaianas que presentan niveles superiores al 90%), pues consideran que una de las principales causas que lo explican son las reducidas dimensiones de la isla y los escasos microambientes particulares que promueven la radiacién adaptativa de los taxa existentes. Sin embargo, la distinci6n de varias comunidades vegetales es un reflejo de ambientes diversos, en cada uno de los cuales se presentan ciertos grados de presion selectiva. Probablemente, otros factores que mencionan Levin y Moran (1989), como el relativo pequefio tamano del archipiélago Revillagigedo y la cercanfa de isla Socorro al macizo continen- tal, sean aspectos de mayor influencia sobre el moderado ntiimero de especies endémicas de la isla. De igual forma este fen6meno puede ademas encontrar una respuesta en el origen de la misma isla, posiblemente de unos pocos millones de afios (Bohrson 1994). No deben descartarse, tampoco, la extincion masiva de la biota por erupciones volcdnicas durante el pasado, tal y como sucedi6é en la década de los afios 50° en la cercana isla San Benedicto, tras lo cual, el proceso de sucesi6n primaria ha tomado reiteradamente su lugar. Cabria senalar que la alta proporci6n de especies herbaceas sobre la isla (mas de la mitad de las especies, ver cuadro 2), y particularmente de gramineas y compuestas, formas de crecimiento y grupos taxonémicos que exhiben una alta capacidad de dispersi6n, puede interpretarse como un hecho que supone una colonizacion relativamente reciente. Tal capacidad obedece probablemente a sus estrategias reproductivas y sindromes de diseminacién, LEON-DE LA Luz, ET AL., Vegetacion de la Isla Socorro, México 229 entre los que cabe mencionar el bajo concurso de polinizadores y su predisposicién a la autogamia y la agamoespermia, la baja proporci6n biomasa total/nimero de semillas, la tendencia a la anemocoria, la aparente alta viabilidad de semillas y la elevada tasa de crecimiento en el estadio de pldntula (Carlquist 1974 y Grant 1971). Todas estas caracteristicas representan condiciones para alcanzar y prosperar en comunidades insulares. CONCLUSIONES El conjunto de condiciones ambientales propias de la posicién geografica y la accién antropogénica, han favorecido el desarrollo de diferentes tipos de vegetacion en la isla. Algunas comunidades, como los bosques, parecen establecerse en lugares condicionados por las variaciones ambientales impuestas por el gradiente altitudinal; en tanto que otras, como el matorral de Croton masonii que se establece primordialmente sobre las capas basalticas, deben su presencia al tipo de sustrato. Los pastizales actuales deben considerarse comunidades secundarias, generadas a partir de la accion humana sobre dreas del matorral de C. masonii con cierta acumulacion de suelo. Los bosques sustentan el mayor numero de las especies del biota de vertebrados terrestres presentes en la isla, particularmente de aves (Brattstrom 1990, Rodrfguez-Estrella et al. 1995), que ejercen una activa funcion ecolégica. El bosque ubicado en la parte norte de la isla (bosque de Oreopanax-Ilex) es el de mayor diversidad floristica y posiblemente la presencia de esta comunidad en esta parte del area se debe a su exposici6n a las corrientes himedas, a la neblina, y consecuentemente a una menor incidencia de radiaci6n. Analizando el conjunto de comunidades boscosas, destaca la presencia continua de Bumelia socorrensis y Guettarda insularis, en tanto que Psidium aff. sartorianum sOlo se registra en el sector sur; también, Oreopanax xalapense y Meliosma nesites presentan una distribucioén restringida a las mayores elevaciones. Del estudio actual de la vegetacién de isla Socorro destaca la severa erosi6n a la que se encuentra sujeta, asf como a los drasticos cambios sufridos du- rante los Ultimos treinta afios, como producto del sobrepastoreo de las poblaciones de borrego. Urgen medidas de control de la erosi6n en el area sefialada en la Fig. 1 de este trabajo. Es dificil pensar en la instalacién de infraestructura que contenga la avenida de los arroyos y torrenteras, como presas de gaviones y otras, debido a los elevados costos, mano de obra y problemas logisticos que implicarfan. Los autores consideran que el repoblamiento con especies nativas, como Opuntia sp., Dodonaea viscosa e incluso Preridium caudatum, pueden coadyuvar a la atenuaci6n de la erosién, 230 Sipa 17(1) ya que presentan caracteristicas propias para desarrollarse bajo esas condiciones. Son altamente recomendables investigaciones que atiendan este problema. REFERENCIAS Abe, J., E. Copo, L. BLAZQUEZ PF. MIRANDA, - VittaALtopos, T. HERRERA, B. VILLA, y L. VAzQuez. 1960. La Isla Socorro, archipiélago de las Revillagigedo. Universidad Nacional oe Autonoma de México. nar del fasten de Geoffsica/2 Bonurson, W.A. 1994. Geology and geochronology of Socorro Island. Memorias de la Reunién Internacional - Investigadores del Archipiélago Revillagigedo, México. Nov. —18, 1994. IPN, CIB, SM, CONACYT y UNAM. P. 6. Bra TSTROM, B.H. oo, of the Islas Revillagigedo, Mexico. J. Biogeogr. 17:177— 183 — Bryan, W.B. 1949. High silica alkaline lavas of Clarion and Socorro Island, Mexico: their genesis and regional significance. Ph.D. thesis. Univ. Wisconsin. (Dissertation Abstracts 20089. 1959) CARDENAS, S., A. CASTELLANOS, P. GALINA, A. ORTEGA y G. ARNAUD. 1994, Aspectos de la poblacion y el habitat del borrego doméstico (Ovzs aries) en la Isla Socorro. Memorias de la Reunion Internacional . Investigadores del Archipiélago eats rca México. ana A 18, 1994. IPN, CIB, SM, CONACYT y UNAM. CARLQUIST, S. 1974. Island he Columbia University Press. — a Cora, R. . C eens In: OrrrGA, A. y A. CASTELLANOS VERA, eds. La Isla Socorro, Reserva de la Bidsfera Archipiélago de Revillagigedo, México. Pub. 8. Centro de Investigaciones Biolégicas del Noroeste. La Paz, México. Pp. 55-62. Gentry, H.S. 1949. Land plants collected by the Velero HI, Allan Hancock Pacific Expe- Bren 1937-1941. Allan Hancock Pacific Expeditions, University of California Press. Los eee Pp. 63-73. GRANT, V. 1971. Plant speciation. Columbia University Press. 2nd. Ed. New York. JOHNSTON, ILM. 1931. The flora of Revillagigedo Islands. Proc. Calif. Acad. Sci., Ser. 4, 20(2):9-104 Levin, G.A. y R. as 1989. The vascular flora of Isla Socorro, Mexico. Mem. San Diego Soc. Nat. Hist. 16 Maya, Y., F. Satinas y E. Troyo. Estado actual del suelo y propuestas para su conservacién. In: Ortega, A. y A. Castellanos Vera, eds. La Isla Socorro, Reserva de la Bidésfera Archipiélago de Revillagigedo, México. Pub. 8. Centro de Investigaciones Biolégicas del See La Paz, México. Pp. 66-76 MEDINA, G 1957. Memoria de la expedicién cientifica a las islas Revillagigedo. Imprenta Sens Guadalaja Miranba, EF. y E, HERNANDEZ 2X. i Los tipos de vegetacion de México y su clasificaci6n. Bol. Soc. Bot. Mex. 28:29-17 ee R., J.L. LEON be ta Luz, A. BREcEDA, A. CASTELLANOS, J. CANCINO y J.Luinas. 1996. eli minary assessment of density and habitat oe of the terrestrial birds i cae Island, Mexico. Conservation Biol. 76:1 RzepowskKl, J. 1978. Vegetacién de México. Editorial Limusa. México : F Rzrpowskl, J.y R. oe VauGH. 1966. La vegetaci6n de Nueva Galicia. Contr. Univ. Michi- gan Herb. 9:1—12 CATALPA (CATALPA BIGNONIOIDES, BIGNONIACEAE) AND BOIS D’ARC (MACLURA POMIFERA, MORACEAE) IN EARLY TEXAS RECORDS DEL WENIGER Professor Emritus Our Lady of the Lake University 411 8.W. 24th St. San Antonio, TX 78207, U.S.A. ABSTRACT Field notes of pre-1860 Texas surveys in the Texas General Land Office were read and the included data on witness trees collected. This data were analyzed for evidence concern- ing the indigenous range of three trees: Catalpa speciosa Warder, oe eS Walt. and Maclura pomifera (Raf.) Schneid. Catalpas were named as witne 1860 in nly two east Texas counties, Jasper and Newton. Their presence in oe surveyors’ reports from only these two contiguous counties, with their absence from the thousands of reports from the rest of the state, is advanced as evidence that these two counties make up most, if not all of any Catal pa’s indigenous range in Texas. The location of these two countie at the southeast Texas border seems to indicate that only C, eer re i Catalpa) should be Se oo in this state. Bois d’ arcs appear as witness trees before 1860 in onl northeast Texas counties. This is presented as evidence that the native range of ae ae in Texas was probably lmited to those 12 counties. RESUMEN e leyeron las notas de campo de los informes sobe Texas, anteriores a 1860, de la Texas ees Land O fice y se recogieron los datos existentes acerca de los arboles testigo. Estos datos fueron analizados para buscar evidencias acerca del drea autéctona de tres arboles: Catalpa speciosa Warder, Catalpa bignontoides Walt., y Maclura ne (Raf.) Schneid. An- tes de 1860 las catalpas fueron designadas como arboles testigo en sdlo dos condados del este de Texas, Jasper y Newton. Su presencia en los informes de los topdgrafos en sdlo estos dos condados contiguos, junto con su ausencia de miles de informes del resto del estado, se toma como evidencia de que estos dos condados forman la mayor parte, si no el total del drea autéctona de Catalpa en Texas. La localizaci6n de estos dos condados en la frontera sudeste de Texas parece indicar que sélo C. bignonioides (1a catalpa del sur) deba ser considerada autéctona en este estado. Antes de 1860 Maclura pomtfera aparece, como arbol testigo, en Ww s6lo 12 condados del nordeste de Texas. Esto se presenta como evidencia de que el area nativa de Maclura pomifera en Texas estaba limitada probablemente a estos 12 condados. More than 50 years ago it came to the attention of some who were study- ing the history of plant communities that there was a wealth of informa- tion for them in accounts of witness trees included in the field notes of Sipa 17(1): 231-242. 1996 232 Sipa 17(1) early surveyors. Those surveyors did not have the luxury of leaving perma- nent markers indicating the points on their lines or of having sophisticated instruments with which to relocate the points, so it was important for them to leave directions enabling those coming after them to relocate the points by using the natural surroundings. To help make that possible it was the early surveyor’s responsibility to follow enn instructions of the survey- ors-general. Those procedures were outlined by Dodds et al. (1943). Those directions mandated that the surveyors, after establishing points, were to locate the trees nearest to the points. Those trees were called “witness trees.” And it was the surveyor’s duty to record in his field notes the direc- tion by compass reading and the measured distance from each point to at least one witness tree, as well as to note the species of that tree and its trunk diameter. Although almost none of those witness trees still survive, the original records of where and what species literally millions of them were are care- fully preserved in official files of the governmental land offices of the U.S. Lutz (1930) was among the first to make use of this material to help determine the original vegetation of an area. Other early studies based to a large extent on the witness tree records were those of Mcintosh (1962), Oosting (1942), and Stearns (1949), There soon was so much reliance on these witness tree records that it became necessary to deal with any possible bias or negligence on the part of the surveyors. Several ways in which they might have skewed their tree records were pointed out. It is possible that they might have bypassed small trees and chosen only large specimens to mark and record. However, it does not seem that this happened in any significant amount, since I find many witness trees re- corded as only 6 inches in diameter; the smallest one in early Texas records was a sapling only 4 inches in diameter. In other places even smaller trees were sometimes used. The 1832 survey of Sumter County, Alabama, in- cludes witness trees down to 2 inches in diameter (Jones & Patton 1966), and it has been stated that, “in exceptional cases, 2 and 3 inch stems were blazed” (Bourdo 1956). It could be surmised that surveyors might discriminate against short- lived and cull trees, but, since willows, birches, and understory witness trees appear regularly in environments appropriate for them, this does not seem a significant factor. Bourdo (1956) pointed out that the trees selected may be expected to present a complete qualitative list of what was present because “there is a tendency to use an uncommon species . . . because relo- cation of the corner will be easier. Some have thought that surveyors, by just not bothering to record trees, might have left an incomplete record of them, but the urge to minimize WENIGER, Catalpas and bois d’arc in early Texas records 233 labor seems to have kept this from being a significant factor. In most cases any surveyor not finding a tree within reasonable distance from any point was expected to erect a cairn of stones on that point. The time and work involved in doing this would not have been relished, and the fact that I find some witness trees described as being many yards away from their points shows that the surveyors chose to measure quite long distances to trees rather than to laboriously gather stones and erect monuments. And the paucity of cairn records in any but what are known to have been very open deserts or grasslands emphasizes that if there was any tree in the vi- cinity of a point it can be presumed that it was recorded. There was sometimes malpractice and even fraud in surveying, resulting in points and lines being misplaced. However, this does not need to con- cern us, because even though such incorrectly located points introduced errors into land plats, they were still arrived at without regard for existing trees. Bourdo (1956) concluded that “errors of this sort do not necessarily affect the usefulness of the field notes.” Studies based on Texas witness trees came late and have been few, but some workers, such as Harcombe and Marks (1977) and Schafale and Harcombe (1983), used them in reconstructing some of the original forest communities of southeastern Texas. All these studies have dealt with plant communities on a more or less local and quantitative level. It occurred to me that the witness tree evi- dence could show a different kind of qualitative reality when viewed from a broader perspective. I therefore embarked upon an attempt to get a gen- eral view of the whole of early Texas in its original state as shown by the witness trees the surveyors said were present before significant settler in- terference with them. Because of the knowledge of on-going processes that caused significant changes in the natural ecology after 1860, I chose 1860 as my cut-off date and used only surveys made up to that date to establish the original scene. In this study I was able to go through all field notes of the pre-1860 surveys in the general file in the Records Division of the Texas General Land Office, Austin. I compiled data on 153,030 witness trees as described in 22,879 early Texas land surveys. Each survey was located by the Land Office down to the county in which it is presently included. The surveys are on file as official land office records and are available for study in the Texas General Land Office. From these | have therefore been able to make lists of the trees officially declared to have been part of the original vegeta- tion of each present county, and also to make range maps showing the counties in which each species was found growing before 1860 and in which each species can therefore be presumed indigenous. 4 Stipa 17(1) @ Counties having pre-1860 witness trees called Catalpas Fic. 1. Texas counties having pre-1860 witness trees called “catalpas.” WITNESS TREE DATA APPLIED TO THE ENIGMA OF TEXAS CATALPAS Catalpa is the American Indian name for some trees found in scattered locations almost throughout the eastern half of the U.S. That such a word has entered our language as the accepted common name for some trees and that the word has even become the name for the genus indicates that these are indigenous American trees. It is also generally agreed that there are two species of this genus in the U.S. One, Cata/lpa speciosa Warder, the northern catalpa, is indigenous in the central Mississippi River basin, with general agreement that it grew native from at least Indiana and Illinois southwest into Missouri and Arkansas. The other, Cata/pa bignonioides Walt., the south- ern catalpa, is thought to have been native from Florida west into at least Mississippi and, according to some, into Louisiana. But today we find catalpas in scattered locations almost throughout Texas, sometimes planted, sometimes growing wild. So what about the catalpas in Texas today? Are they Texas natives or not? If native, where in Texas did oO ie) WN A) WENIGER, Catalpas and bois d’arc in early Texas records ice L aa a Pe ie ee Oe a eee Seeeen, aac pei ae —e---o rte rae A Oe OR Ne ae AO | ar oe he a eo Cc ene See ees eet ae ‘. eee iee ' _— as i eae wet, . en % a Se an a Wag. : gee. Weta y \ See mies a F Ne ieee ~ i non a ~~ \ @ Counties having pre-1860 witness trees called bois d’ arcs -) Counties having pre-1860 witness trees called yellow woods Fic. 2. Texas counties having pre-1860 witness trees called “bois d’arcs” and “yellow woods.” they originally grow? Botanists dealing specifically with Texas leave us with rather uncertain and sometimes even contradictory answers to these questions. Concerning the northern catalpa, Sargent (1905), writing early, stated flatly that this tree was not native anywhere in southern Arkansas, western Louisiana, or eastern Texas. Harrar and Harrar (1946) gave its range as from “Illinois and Indiana south to Arkansas,” thus leaving out Texas all together. A guide to the trees of Texas published by the Texas Forestry Association (1928) says C. speciosa “is found naturalized in east Texas,” so denying it a Texas origin. Vines (1960) and Correll and Johnston (1970) were noncommittal, dealing only with the tree’s present distribution. Simpson (1988), by not even listing it in his guide to Texas trees, makes it clear that he does not consider this a Texas tree. But Shinners (1958), in his study of northern Texas plants, says the tree is “rare as a native tree in pine belt [of Texas],” although he attributes the statement to Smith (1970). 236 Sipa 17(1) There 1s equal disagreement about C. bignonioides in Texas. Small (1903) early stated that the tree is native in southeastern Texas. Reeves and Bain (1946) also considered it to be a Texas tree. Vines (1960), however, said it is “thought to be native from Florida and Georgia, westward into Louisiana. Doubtful whether native in Texas.” Correll and Johnston (1970) did not commit themselves on the point. Harrar and Harrar (1946) made the definite statement that the tree ranges west only as far as Mississippi, and neither the Texas Forestry Association (1928) nor Simpson (1988) even mentioned this species, these latter three at least implying that the tree is not indig- enous to Texas. So we have the enigma of Texas catalpas. They may both, the northern and the southern, be here today. But is either one indigenous here? Is there any way to determine that from any actual evidence? The accompanying map shows the present counties of Texas in which catalpas were found and recorded as witness trees prior to 1860. They ap- pear only in surveys of Jasper and Newton counties of that early era. This is significant in several ways. First, the fact that there was not one cata of trees used as witness trees in all of northeastern Texas, which was appar- ently the first region of the state to be widely surveyed, makes it highly unlikely that there were any indigenous catalpas in that region of Texas. Second, catalpas recorded as growing in two counties just within the southeastern corner of Texas at such an early time provide evidence hard to deny that some catalpa is indigenous to that limited part of the state. This pas had to have been already _ pa among all of the thousands — is especially crue because some of those catal mature and grand in size when the pioneers first entered Texas. So early as 1838, in listing the forest trees found in Texas at that time, von Wrede (1970) described them in the following words: “Next would be the catalpa tree which is forty to fifty feet high, with a trunk up to three feet in diam- eter and foot-long leaves which are eight to ten inches wide. Its flower, a pyramid-shaped cluster similar to the horse chestnut, is a magnificent sight.” Third, the absence of any early-day catalpas from all of northeastern Texas makes it highly unlikely that che northern catalpa was native to the state. If those found down in Jasper and Newton counties were C. speciosa they would have been a relict population isolated by several hundred miles and avery different environment from the nearest definitely known native stands of that species. And fourth, those catalpas found growing just within the southeastern border of Texas were separated by no large distance or major environmental factors from the known range of C. bignonioides. It would seem to follow, then, that these catalpas must have been the southern catalpa, extending that species’ known range just into Texas. > WENIGER, Catalpas and bois d’arc in early Texas records 257 I submit that this is clear and adequate new evidence allowing us to put C. bignonioides into the list of indigenous Texas trees but also indicating that we should delete C. speciosa from that list. THE BOIS D’ ARC IN EARLY TEXAS RECORDS The bois d’arc or Osage-orange, Maclura pomifera (Raf.) Schneid., has been called “perhaps one of the classic examples of an endemic species in North America” (Smith and Perino 1981). Endemic it certainly is, and as Texan as any, but that does not mean there has been consensus about the tree's natural range in Texas. Some authorities, for instance Correll and Johnston (1970), have merely made general comments about the tree’s being found growing in at least Oklahoma, Texas, and Arkansas but avoid being more specific. Beyond this, statements of its natural range in Texas are widely divergent. Collingwood (1939) claimed the tree was native from beyond the Red and Sabine rivers southwest to at least the Rio Grande and beyond the lower Pecos River. Betts (1953) claimed it was native over all of eastern Texas but the coastal plain and all of central Texas to beyond San Antonio. Burton (1963) drew its natural range as a narrower belt from the northeastern corner of the state down to an arc from about San Antonio on the west to the coastal plain near Wharton. Such ambiguity concerning an endemic species is remarkable, and there must be a reason for it. The reason for this uncertainty concerning the original range of this tree is that even before the time of first botanizing in this region bois d’arc had already been introduced into new areas far outside its original range. How very early this process was going on is attested to by the remarkable state- ment of Custis (1984). In his catalog of plants he found in 1806 on the Red River he wrote, “Bois d’arc, of this tree you have already had a descrip- tion.—it is probably a new Genus . . . it is said first to make its appearance about the 2nd Little river and is very care: on acreek called Bois d’ arc. The tree which I saw was one growing within a mile of Nachitoches [Loui- siana}] & was probably transplanted.—This is about 30 ft. high; its trunk 7 to 8 ft. in circumference & about 6 to 7 ft. to where it begins to ramify.—— its general aspect is that of an apple tree.—its fruit is about the size of the large sour Oranges and of a greenish yellow.” Any tree having attained by 1806 the size described by Custis (1984) must have been transplanted long before, perhaps when there was no one present at that site to do the job except Indians. This may mean ascribing much more in the way of horticulture to the Indians than we would have suspected, but Custis’ observation may require it. His editor notes that, “Scattered trees found farther east, in the vicinity of the Caddo settlements, quite possibly were transplanted by the Indians.” Simpson (1988) recently 238 Sipa 17(1) noted that “Bois d’Arc shows up in many places in the Trans-Pecos—al- ways at Indian campgrounds or caves . . . Today, great thickets of Bois d’Arc are found in these areas, seeded from the horse apples carried by these tribes.” The tree was also carried about by explorers and settlers. In 1804, Lewis (1962) wrote ina letter to Thomas Jefferson that he had sent him cuttings of this plant. Lewis stated that he had obtained the cuttings in St. Louis “from the Garden of Mr. Pierre Choteau .. . [who} informed me, that he obtained the young plant at the great Osage Village from an Indian of that nation, who said he procured them about three hundred miles west of the place.” A huge individual of the species still stands on the grounds of the American Horticultural Association, within sight of the Potomac River. These grounds were once George Washington’s River Farm, and since it is known that Jefferson gave Washington other trees, this individual may be either one of Lewis’s cuttings or a descendant of them. There is also a single equally huge, old bois d’arc in the front yard of the Oatlands Plantation near Leesburg, Virginia. Since Jefferson is known to have visited this home as well, this may be another legacy of Lewis, through the agency of Jefferson. This chain of events emphasizes the remarkably early introduction of this species across the land. The tree proved so useful for living fences that John A. Wright, editor of the Prairie Farmer, and others were promoting it ac- tively from the 1850s on. Such widespread introductions of the tree have erased for all time any possibility of field work rediscovering the limits of the species’ natural range. All we have now is some indication of where each author found it growing in his time. So how, at this late date, can we ever arrive at the true original range of this important endemic tree? I think there is a simple — — answer, but one involving much tedious research. It is simply to check explorers’ written accounts and field notes from their surveying projects for their testimony concerning where they found these trees before the settlers came in and broadcast them. I have carried out such a lengthy records search for the state of Texas. To this end I have read all of the surviving material written by people who were in Texas before 1860 that I have been able to locate. This in- cludes eyewitness accounts of many aspects of the region in books, letters, itineraries, reports official and unofficial of exploring and military expedi- tions, diaries, etc., written by 290 explorers, soldiers, adventurers, settlers, and other pioneers who, up to 1860, experienced and observed the Texas wilderness and took the trouble to write about it. I have not used any reminiscences written at a distance from the observations and so subject to possible errors of memory or the prideful embellishments of old-timers. In — WENIGER, Catalpas and bois d’arc in early Texas records 259 these accounts there are thousands of reports of different trees found grow- ing in those early days in Texas. I have not found any distinct tree species generally recognized as native to Texas that 1s not reported by at least some of these explorers. And the bois d’arc is there. Among all those tree descriptions I have found only 14 pre-1860, con- temporarily written records of bois d’arcs growing in Texas. The small num- ber of these early bois d’arc reports seems very significant in itself, when compared to the more than 600 separate written records of mesquites ap- pearing in those pioneers’ accounts. This alone would seem to promote a healthy skepticism about the hypothesis that the bois d’arc was native to any wide area of the state. I have also tabulated the data on 153,030 witness trees located and named in early Texas surveys. Of all these thousands of Texas witness trees, only 123 were bois d’arcs. This hardly portrays the species as of any general significance in the original tree cover of the state. And the locations where those naturally growing bois d’arcs were observed further reduces them from state-wide importance. Map 2 shows the present counties of Texas in which bois d’arcs were named as witness trees up to 1860. The testimony of the early surveyors is that the species was then growing wild in what have become 12 counties: Bowie, Collin, Dallas, Delta, Fannin, Grayson, Hopkins, Hunt, Kaufman, Lamar, Red River, and Rockwall. Because in each of these counties some of these trees were stated to have been large, mature individuals, they must have been indigenous. And although it is impossible to prove a negative, the fact that, in more than 150,000 pre-1860 witness trees from all of the rest of Texas, there appears not one bois d’arc makes for a very strong pre- sumption that this 12-county area was the total original range of this spe- cies in the state. This presumption is strengthened by the 14 explorers’ accounts of the tree all in the same 12 counties, by the fact that no explorer mentioned the species in any other location 1n Texas, as well as by the very early naming of two creeks, one draining the upper part of this same area into the Red River and the other running out of the lower part of it into the Trinity River, both as Bois d’Arc creeks. I therefore submit the conclu- sion that this 12-county area and this alone was the natural range of this species in Texas. The early records make it clear that the pioneers did not use the name “yellow wood” for bois d’arc but for some other. (What “yellow wood” is cannot be determined from the common name alone. In pertinent litera- ture several species of trees are referred to by that name.) We can see this because, first, no tree found growing within the original range of bois d’arc as outlined above was called yellow wood in pre-1860 writings, and sec- 240 Sipa 17(1) ond, because all of the yellow woods named previously to 1860 were found over 200 miles from this species’ otherwise known range. Map 2 shows the discrepancy in the ranges of the pioneers’ bois d’arc and yellow wood. After reading the explorers’ reports I am convinced that we have little conception of how profusely the bois d’arc grew in the center of its native haunts along the creeks named for it, so Ladd some of their descriptions of this. In 1805, Sibley (1931) wrote of “a beautiful bayou situated on the left side {of the Red River}” that “the French call ... Bayou del Palo de Arco, Bois d’Arc, or Bow-wood Creek, from the large quantity of that wood that grows upon it, of which the Indians make their bows.” Moore (1965) wrote of the “forests of Bois d’Arc” in the same area. Several of the surveyors described what they encountered in these areas as “thickets” of bois d’arc. There is an especially interesting passage showing incidentally the large amount of bois d’arc that must have been in the area of Collin and Rockwall counties and at the same time showing a strange way in which these trees could have been a danger to travelers. The correspondent of the Clarksville {Texas} Standard, who signed himself merely as C De M, was traveling from McKinney down the East Fork of the Trinity in 1853 (C De M 1853). When at about the present boundary of Collin and Rockwall counties, his party had to stop for the night because “the creeks upon both sides of the river were overflown, and although there was a ferry flat at the river, there were none at the creeks, and so we concluded to wait a few hours, as we were told that swimming the creeks might be dangerous from Bois d’Arc brush in them full of thorns—so we waited till morning, and the water rose during the night.” That the thorny branches of this species could be a real hazard and, at least upon Bois d’Arc Creek itself, could form impenetrable thickets is further emphasized by the description of one curious episode. Marryat (1970) wrote that he was struggling to get from the western wilderness to the comparative civilization of Louisiana in 1843, when he experienced the situation which he described as follows: “... after traveling some six or eight miles, we found our further progress cut off by a deep and precipitous chasm, lined with impassable briars. To return was our only alternative, and at noon we again found ourselves near to the point from whence we had started in the morning. A consultation was now held as to our future course ... On the morning of the preceding day we had passed a large, though Shallow, sandy stream. Gabriel and I thought it more advisable to return to it. This stream was evidently one of the tributaries of the Red River, and was running in an easterly direction, and we were persuaded that it must flow through the chasm, and enter into the forest ... The next day at noon, we encamped on the stream ... The next morning after break- WENIGER, Catalpas and bois d’arc in early Texas records 241 fast, we filled our saddle-bags with the remainder of our provisions, and followed the stream for ten miles, with water to our horses’ shoulders, as both sides of the river were covered with briars ... For nine miles more we continued wading down the river, till at last the prickly pears and briars receding from the banks, allowed us once more to regain the dry ground.” Anyone who has ever stumbled into and had to extricate himself from a patch of bois d’arc sprouts surely can understand why thickets of these would be virtually impassable and also why they might be called briars. And Nuttall (1980), who first named this species, attested to its growing in just such thickets as described in Texas by writing that when almost directly across the Red River from Bois d’Arc Creek in Choctaw County or western McCurtain County, Oklahoma: “... along the margin of all the rivulets we met with abundance of the Bow-wood (Maclura), here famil- iarly employed as a yellow dye.” So I think that we have to make the effort to imagine at least the center of the 12-county range already outlined as a vicious, thorny, almost impassable thicket of bois d’arc instead of the open expanse it is today. Smith (1970) required this of us, having written in 1849, that “he thinks very highly of Reily’s lands on the Bois d’arc, al- though they are covered with the dense thickets peculiar to this creek.” But this plant was not only a thorny shrub. While writing of this same general area in 1840, Moore (1965) gave something of the size of the par- ent trees: “The Bois d’ arc trees attain a remarkable size and are often found four feet in diameter and eighty feet in height.” We may class this as among the grandest of Texas’ original forest trees, but it seems we should conclude that its natural range in the state was not large. ACKNOWLEDGMENTS I thank the Ewing Halsell Foundation and the Robert J. Kleberg, Jr. and Helen C. Kleberg Foundation for their support, which made the re- search and writing of this report possible. REFERENCES Berts, H.S. 1953. American woods: Osage-orange. U.S.D.A., Forest Service. Bourpbo, E.A., JR. 1956. A review of the General Land Oifce Survey and of its use in ’ e quantitative studies of former forests. Ecology ee 768. Burton, J.D. 1963. aie pelea U.S.D.A. Publ. 0101-00315 C = M [Dr Morsg, C.}. . Letter from a ae Caesal le {Texas} Standard, 4 May ne oe G.H. 1939. Osage-orange. Amer. For. 45:508. CorreLl, D.S. and M.C. Jounsron. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. P. 498. Custis, P. 1984. The Custis accounts of the Red River ... in the year 1806. In: Dan L. Flores, ed. Jeffe oe Exploration. Univ. Oklahoma Press, Norman, OK. Pp. 260-261, note 2 242 Sipa 17(1) Dopps, J.S., J.P. McKean, L.O. Srewarr, and G.E. Ticces. 1943. pie instructions governing public land surveys of lowa. lowa Engineering Society, Ame Harcompe, P.A. and P.L. Marks. 1977. Understory structure of a mesic ae in southeast Texas. Ecology 58:1144-115 Harrar, ELS. and J.G. Eran: 1946. Guide to southern trees. McGraw-Hill Book Co. New York. P. 656 Jones, A.S. and E.G. Parton. 1966. Forest, Sed and soils in the Black Belt of Sumter County, Alabama, in 1832. Ecology 47:75— Lewis, M. 1962. Letter to Jefferson, March a 1804 . In: D. Jackson, ed. Letters of t Lewis and Clark Expedition, with related documents, 1783-1854. Univ. Illinois es Urbana. Pp. 170-171. Lutz, H.J. 1930. Original forest composition in northwestern oo as indicated by early land survey notes. J. Forest. (Washington) 28:1098—1103 Marryart, F. 1970. The travels and romantic adventures of Mme viclet Gregg Press, Upper Saddle River, NJ. (First published in 1843.) Pp. 283-2 McinrosuH, R.P. 1962. The forest cover of the Catskill ee Re gion, New York, as indicated by Land Survey Records. Amer. Midl. Naturalist 58:409—423, Moore, F., Jk. 1965. Map and description of Texas. Texian Press, Waco, TX. (First pub- lished in 1840.) P. 59 Nutra, T. 1980. A ouraal of travels into the Arkansas Territory during the year 1819, Univ. Oklahoma Press, Norman. (First published in 1821.) OostING, J.J. 1942. An ecological analysis of the plant communities of Piedmont North Carolina. Amer. Midl. Naturalist 28:1—126. Reeves, R.G. and D.C. Bain. 1946. A flora of south central Texas. A&M College of Texas, College Station, TX. SARGENT, C.S. 1905. Manual of the trees of North America. Houghton Mifflin, Boston, ScHAFALE, M.P. and P.A. HarcomBe. 1983, tet vegetation of Hardin County, Texas. Amer. Midl. Naturalist 109:355— SHINNERS, L.H. 1958. Spring flora of the Se Worth area, Texas. Published by the author, Dallas. Siptey, J. 1931. Untitled. Volume 2, p. 71 in Pichardo’s Treatise on the limits of Louisiana and Texas, by Father Jose Antonio Pichardo, tr. Charles Wilson Hackett, Austin: Univ. Simeson. B.J. 1988. Field guide to Texas trees. Texas Monthly Press, Aus SMALL, J.K. 1903. Flora of the southeastern United States. Published by amos New York SMITH, E. 1970. Account of a journey through north-eastern Texas undertaken in 1849, embodied in a report to which are oe nded letters and verbal communications from eminent individuals ... third part. East Texas Hist. Assoc. J. 8:72. Smitu, J.L. and J.V. Perino. 1981. Osage orange (Maclura pomifera): history and economic uses. Econ. Bor. 35:24-4l. STEARNS, FW. 1949. Ninety years change in a northern hardwood forest in Wisconsin. Ecology 30:350—-358 TEXAS FORESTRY Assoc IATION. 1928. Forest trees of Texas. Texas Forestry Association, Col- lege Station Vines, R.A. 1960. Trees, shrubs and woody vines of the Southwest. Univ. Texas Press, Austin VON Wrebe, EW. 1970. Sketches of life in the United States of North America and Texas. Translated by C.W. Geue. Texian Press, Waco, TX. (First published in 1844.) P. 100. EMANUEL D. RUDOLPH’S ROLE FOR BIOLOGICAL ABSTRACTS WILLIAM R. BURK Biology Library University of North Carolina Chapel Hill, NC 27599-3280, U.S.A. ABSTRACT The life of Emanuel Rudolph (1927-1992), noted polar lichenologist and historian of botany, was well documented by Ronald Stuckey. Rudolph’s scholarly endeavors is further revealed through his devotion to Biological Abstracts. He accepted the intellectual chal- lenge Serre eee in 1952 when a graduate student in the Henry Shaw School of Botany at Washington University. From then as a volunteer, he prepared — abstracts through 1977, a fe exacting and time consuming task, perhaps an art. Some back- ground on procedures at Biological noe is reviewed and Rudolph’s role as a ie aie abstractor for Biological Abstracts/BIOSIS is assessed. RESUMEN La vida de Emanuel Rudolph . eum notable cela ae e historiador de la botanica, que fue bien documentada por | ald Stuckey. Rudolph, se desvela en un grado mayor a través de su dedicacién a = ee ee El acepto el reto intelectual de preparatr resumenes en 1952 cuan idi en la Escuela de Botdnica Henry Shaw de la Universidad de Washington. Desde entonces como voluntario, preparé 382 resimenes durante 1977, una tarea muy exigente y laboriosa, quizds un arte. Se revisan algunos antecedentes de los procedimientos de los Resimenes Biolégicos, y se evaltia el papel de Rudolph como voluntario en la elaboracién de resimenes para los Bio- logical Abstracts/BIOSIS BACKGROUND ON ABSTRACTING AT BIOLOGICAL ABSTRACTS The practice of abstracting emerged and developed after 1790, but its roots precede the advent of printing when annotations were placed in the margins of manuscripts. Scientific journal abstracts summarize articles ina succinct manner so that ee have a clear idea of their content, and therefore value to them. Biological Abstracts, one of the pre-eminent abstract- ing journals in the field of biology, was first published in December 1926, formed by the merger of two existing abstract indexes, Botanical Abstracts and Abstracts of Bacteriology (Manzer 1977). Some general background in the production of abstracts is explained in the preliminary pages of the January 1955 volume of Biological Abstracts. The distinction between Biological Abstracts, the serial, and Biological Abstracts, the corporate name, was con- Siva 17(1): 243-249. 1996 Q44 Stipa 17(1) fusing until the corporation title was changed to BioSciences Information Service of Biological Abstracts (BIOSIS) in October 1964 (Steere 1976). In 1928 the editors of Biological Abstracts relied on 3,000 scientists throughout the world to provide abstracts on a voluntary basis (Biological Abstracts 1957). At that time such a large number of volunteers was needed because the inclusion of abstracts in journals was the exception rather than the rule. Due to financial difficulties, the publication of Biological Abstracts nearly ended in 1936. Following the reorganization of Biological Abstracts early in 1938, appeals to the biologists of the United States for volunteer support of the publishing venture were enthusiastically met. Although a new policy had been instituted in 1937 to compensate abstractors for their work in order to speed the preparation of abstracts, collaborators were now returning their monetary compensation. Nearly a year later collaborators “advocated a policy of gratis abstracting, and the payments for abstracting were therefore discontinued” (Flynn 1951). By 1956 the number of volun- teer abstractors declined to 656 people who provided 50% of the needed abstracts, while 30% of the abstracts were written by the authors of articles and forwarded by the journal editor in batches, and the remaining 20% of the abstracts were provided by office staff of Biological Abstracts or copied from published abstracts or summaries (Biological Abstracts 1957). In the late 1960s through the mid 1970s announcements appeared in the preliminary pages of Biological Abstracts for the solicitation of abstrac- tors, especially for those willing to write abstracts for taxonomic publica- tions. One can ascertain the number of volunteer abstractors from 1959 to the present from the lists of volunteer abstractors which were published in the introductory pages of issues of Biological Abstracts for these years. Only 45 volunteer abstractors were listed in 1995. John Schnepp, Chief, Litera- ture Analysis Section, Bibliographic Control Department, BIOSIS, related that this list of abstractors may be eliminated since the inclusion of author abstracts by publishers is now standard practice. (phone conversation of 7 July 1995). The value and impact of volunteer abstractors to the success of Biological Abstracts did not go unsung. Appreciative of che contributions of volunteer abstractors, the administration of Biological Abstracts maintained com- munication with these volunteers and extended its gratitude in various ways. For many years volunteers received a Christmas greeting which sum- marized the year’s accomplishments and progress at Biological Abstracts and also thanked the volunteers for their work. Ann Farren, Associate Bib- liographer, Literature Acquisition Department, stated in her greeting of 15 December 1961 that “Our personal contacts with you have been rare, usually confined to an occasional visit to BA’s booth at national and inter- national meetings. Sometimes you may even think that we have forgotten Burk, Emanuel D. Rudolph 245 you. Believe me this is never true. We are reminded and say a quiet “thank you’ each time a group of your abstracts are received.” Among the senti- ments of thanks that were echoed on numerous occasions by BIOSIS ad- ministrators, those of Phyllis V. Parkins, Director of BIOSIS, were espe- cially touching when she stated that “on behalf of all biologists, [we] declare our deep gratitude and appreciation to our Abstracters for the contribution they continue to make in the communication of biological information. May the sun never set on BA Abstracters!” (Parkins 1965). When volun- teers received updated, revised instructions for the preparation of abstracts or were given new or additional assignments, expressions of gratitude were usually made. Through the years volunteers were offered publications as a token of appreciation, such as: (1) retaining issues of journals for which abstracts were being prepared, and (2) receiving a complimentary subscrip- tion to different publications produced by Biological Abstracts, through a choice of either: a sectional edition of Biological Abstracts (BA); B.A.S.1.C., the subject index to BA; one topic of the STANDARD PROFILE Services, one member of the BIORESEARCH TODAY series; or the annual BIOSIS List of Serials. EMANUEL D. RUDOLPH AS A VOLUNTEER ABSTRACTOR While a graduate student in the Henry Shaw School of Botany at Wash- ington University and the affiliated Missouri Botanical Garden 1n St. Louis, Rudolph first volunteered as an abstractor for Biological Abstracts in 1952. Perhaps his major professor, Carroll W. Dodge, who was editor of the Biog- raphy, History and Bibliography section of Biological Abstracts since its be- ginning, influenced Rudolph’s decision to write abstracts. Because of the broad scope of most of the journals in the 1950s and 1960s, many abstrac- tors were assigned their work on a journal basis rather than by subject specialties. Emanuel Rudolph, however, was given abstract assignments for articles in botanical and antarctic research journals. His first assign- ment was to write abstracts for the Spanish ee botanical journal, Lilloa, most likely because he could read Spanish, was a student of botany, and had access to this journal in the Library at the Missouri Botanical Gar- den. In a letter to Rudolph (14 May 1952), John E. Flynn, Editor-in-Chief of Biological Abstracts, commended him for his fine abstracts. In October 1952, Dr. Flynn again wrote to Rudolph, complimenting him for his ex- cellent abstracts and stating that such superb work was helping to build Biological Abstracts into an instrument of greater utility and better scholar- ship. He prepared abstracts for articles in volumes 16(1949)—31(1962) of Lilloa. Coinciding with the completion of his first assignment, Rudolph was offered, as a token of appreciation, a complimentary subscription to a Sectional Edition of Biological Abstracts. Whether or not Rudolph accepted 246 SIDA 17(1) one of these sections is unknown. An itemization of the number of ab- stracts prepared by Rudolph for L7//oa and other serials is listed in Table 1 by serial title and volume. Upon accepting a teaching position in the Department of Botany at Wellesley College, Massachusetts in the fall 1955, Rudolph wrote the edi- tor of Biological Abstracts to relate that he no longer had access to Li/loa. After repeated correspondence to the publisher of Li//oa, the editor of Bio- logical Abstracts succeeded in obtaining issues so Rudolph could continue to write abstracts for this journal. In fact, Rudolph was allowed to keep any issues sent to him. In June 1958 Rudolph became concerned that he had not received any copies of Li//oa for a long time. Apparently this was a turning point for him to accept a new abstracting assignment, for yet an- other Spanish language botany journal, Anales del Instituto Botanico A.J. Cavanilles, commencing with volume 17 (1959) and later concluding with volume 21 (1963). In the 1960s significant changes occurred in Rudolph’s professional life and in Biological Abstracts’ reorganization. In the Spring of 1961 Rudolph started a new career as an Assistant Professor of Botany at The Ohio State University, Columbus, where rigorous research and an active teaching sched- ule occupied much of his time. These academic responsibilities would even- tually diminish available time for writing abstracts. Considerable reorga- nization occurred at the offices of Biological Abstracts in the 1960s. Its Literature Acquisition Department saw expanded abstracting services due to the exponential increase in scientific journal publications and the com- puterization of operations. Ann Farren asked Rudolph in December 1961 if he would accept an additional assignment. Documentation is lacking whether Rudolph accepted another journal for abstracting when he returned from his Antarctic research expedition at the end of February 1962. Ex- actly three years later, Rudolph took on additional serials for abstracting. The new assignments began, and subsequently ended, with the following volumes of serials: British Antarctic Survey Bulletin, nambers 3 & 4(1964)— 13(1967); Commonwealth Mycological Institute. Miscellaneous Papers, numbers 17 & 18(1964); and Kirkia, volumes 4(1963—G64) —G, Part I(1967). Soon after these new assignments were made, another serial, Commonwealth My- cological Institute, Mycological Papers, which was a successor to the CMI Maiscellanous Papers, was added, commencing with number 99(1965) and concluding with number 141(1977). Rudolph’s busy teaching schedule and research work contributed to de- lays in his preparation of abstracts in the late 1960s. In fact he was unable to complete abstracts for the two Spanish language journals, A nales del Instituto Botanico A.J. Cavanilles and Lilloa because the extra time required to read these foreign language journals was not available. Rudolph reluctantly wrote Burk, Emanuel D. Rudolph 247 Taste 1. Number of abstracts prepared by Emanuel D. Rudolph for Biological Abstracts, arranged by journal title. Volume Year No. of Abstracts Volume Year No. of Abstracts Anales del Instituto Botanico A.J. Cavanilles = 119 1969 | 17 1959 14 120 1970 ] 18 1960 14 121 1970 l 19 1961 1 125 1971 l 20 1962 5 126 1971 1 21 1963 8 127 1971 1 22 1964 9 128 1972 1 subrocal 51 129 1972 2 972 British Antarctic Survey Bulletin oe bene ; Z F 13] 1972 l : a = 939 1973 4 1964 0 | ; ; mane . | - 1974 6 1965 4 ae - 136 1974 l 7 1965 3 ia 1974 166 > 138 oe | 9 1966 3 ss es 139 1975 1 10 1966 2 141 1977 l U1 1967 0 ; 12 1967 1 subtotal 40 13 1967 4 Kirkia subrotal 27 1963/64 19 c 4 Commonwealth heron Institute. 2 oe ve 6(Part I) 1967 6 oo Publication coy 1 subtotal 39 : 1964 1 Lilloa subtotal 2 16 1949 12 7 94C 8 Commonwealth aie Institute. a. oe Mycological Paper : i . 19 1949 23 99 oe l 5 20 1949 24 100 1965 ] 1 17 a 2 194911951} 103 1965 iL =e Mit : A 23 1950 24 104 1966 1 ? 24 1950 2 105 1966 l a ava te 25 L9OS1{1952} 8 106 1966 l : . 107 1967 1 26 1953 12 Di, 1953{1955} 7 108 1967 l 08 1957 g 109 1967 l a . 29 1959 4 110 1968 , 2 30 1960 14 111 1967 : 31 1962 14 112 1967 1 113 1968 subtotal 222 114 1968 l Opera Lilloana 145 1968 l 4 1962 ] 116 1969 l subtotal 117 1969 2 Grand total 382 — = ee) \O ion \o 248 Sipa 17(1) to Robert Sutcliffe, Associate Bibliographer at BIOSIS (25 February 1969), that he had to return any outstanding issues of these two journals. Accord- ing to Rudolph’s own admission, his desire to complete abstracts was greater than his ability to write them at that time. During this same time more journals were beginning to provide author abstracts with articles, a prac- tice becoming more commonplace. Because of this emerging practice Rudolph’s abstracting assignments were cancelled for British Antarctic Sur- vey Bulletin July 1968) and Kirkia (April 1969). Little documentation on Rudolph’s abstracting role after 1977 is avail- able. John Schnepp wrote Rudolph (22 September 1982) to ask if he was “still available to provide fabstracting] service in the future either for che same or for another journal.” Rudolph’s answer must have been affirmative as his name continued to appear in lists of volunteer abstractors in the preliminary pages of each issue of Biological Abstracts for 1982-1992. Even though he may not have been actively writing abstracts from 1978-1992, he was available if needed. In the preparation of abstracts Rudolph was required to follow high standards established by the editorial staff of Biological A th — ystracts. Through ne years guidelines, along with rules for abbreviating and acceptable ab- breviations, were issued to aid the volunteer abstractor. Except for a few unsigned abstracts written in the late 1960s, Rudolph’s abstracts were signed, E.D. Rudolph, and variec column of text. eon in length from one sentence to a whole Emanuel Rudolph’s role as an abstractor provided essential volunteer assistance at a time when Biological Abstracts was still seeking stability as a self-sustaining organization. A search in Biological Abstracts for articles appearing in the serials for which Rudolph prepared abstracts located 382 abstracts written by him. Through these abstracts Rudolph served his fel- low biologists and the scientific community. ARCHIVAL RESOURCES CITED AND CONSULTED Emanuel Rudolph’s archives dealing with his association with Biologi- cal Abstracts/BIOSIS are housed in the Herbarium, Museum of Biological Diversity, The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212-1192 and curated by Dr. Ronald L. Stuckey, Professor Emeritus and Curator Emeritus, Department of Plane Biology, The Ohio State Univer- sity, Columbus, Ohio. ol ACKNOWLEDGMENTS [ thank John W. Schnepp for reviewing and providing pertinent changes to the manuscript. Special gratitude is extended to Ronald L. Stuckey w provided access to Emanuel Rudolph’s archives and reviewed the paper. —_— 10 Burk, Emanuel D. Rudolph N aX \o REFERENCES BioLocicaL Apsrracts. 1957. 30th Anniversary. Biological Abstracts and the literature of biology, a symposium in celebration of the thirtieth anniversary of the founding of the Biological Abstracts, held at the University Museum, Baas of Pennsylvania, on Feb- ruary 17, 1956. [Biological Abstracts, Philadelphia, PA]. FLYNN, - E. 1951. A history of Biological Abstracts. ihe Author}, Philadelphia, PA. (Quote, Pp. ve B.M. 1977. The een aes 1790-1920: Origin, development and diffu- sion. Scarecrow Press, Inc., Metuchen, NJ and London. ParKINS, P.V. 1965. The sun never sets on BA abstracters. Biol. Abstr. 46(20):x1i STEERE, W.C., with the collaboration of PV. Parkins and H.A. Philson. 1976. Biological Abstracts/BIOSIS, the first fifty years, the evolution of a major science information ser- e. Plenum Press, New York and London. ae cKEY, R.L. 1994. fee He Budell sh (1927-1992): Polar lichenologist and his- torian of botany. Bryologist 97:437—-446. aie Publications and reports of Emanuel D. Rudolph ee Serie 443-446, with William R. Burk}. ee eee . Emanuel avin ahi (1927- on Book collector and li friend. ara Bot. 34:4—2° OTary 250 Sipa 17(1 BOOK REVIEW Hosss, CHRISTOPHER. 1995. Medicinal Mushrooms, An Exploration of Tradition, Healing, and Culture. (Library of Congress Catalog Card Number: 95-60381, pbk, 2nd ed.). Botanica Press, 10226 Empire Grade, Santa Cruz, CA 95060, $16.95, 251 pp, 30 color photos. This book is carefully documented through literature sources that are discussed at the very beginning ina section entitled Sources, Historical and Scientific. This lays the ground- work for the 39 page Fungi Bibliography (900 literature sources) that is a comprehensive array of literature that traces medicinal mushrooms from the ancients to presentday. Hobbs notes that the text is divided into two parts: the first focuses on a general overview of the fungi, summarizes the history and use of medicinal fungi and gives practical instructions on how to use them. The second part is detailed and more technical, covering botany, chemistry, pharmacology, clinical studies, indications, and dosages of medicinal mags com- monly available in the United States, Asia and ae and other parts of the w History of Use is a chapter packed with detailed information of medicinal uses sof fungi in Western Medicine (ancient Greek writers, the herbalists, native American Indians) and uses in Asia. There is also a discussion of the origin of the words mushroom, fungus, and agaric that merit careful perusal. The chapters on Modern Uses Of Medicinal Fungt, Adaptogens and Immune Stimulants, and a Guide To Using Medicinal Fungi contain valuable information on certain species of fungi that can enhance a healthful state A Guide to Using Medicinal Fungi tells how to gather medicinal mushrooms, how to store them, how to prepare powdered extracts, powders, and pills, how to make teas and soups, and how to prepare tinctures and doses. Nutritional Value of Fungi (5 pages) and recipes (4 pages) discusses the nutrient content of edible and medicinal fungi. Medicinal Fungi Monographs is devoted to 27 species of fungi. These species are dis- cussed under the following headings: species name and author, common name, synonyms, other common names, description and habitat, range, history, chemistry, pharmacology, human clinical studies, toxicity, uses in traditional medicine, medical uses, preparation and dosage, related species, and procurement. There is a handy table entitled Overview of all Medicinal Mushroom Species that collates much of the previous information into a more simplified and retrievable tabular form. There is s short glossary of 29 mostly medi- cal terms. Resources gives the company name and address of mushroom products. The index is incomplete for many species and common names that appear in a number of places in the text. The modest price, wealth of information carefully gleaned from a variety of excellent sources, and nontechnical narrative directed to a lay audience, make this book a valuable addition to the professional and amateur mushroom 7 One of the best bargains avail- able today.—Harold W. Keller, Research Associate, BRIT jon SIDA 17(1): 250. 1996 DOCUMENTED CHROMOSOME NUMBERS 1996:1. CHROMOSOME NUMBERS IN CAREX SECTION OVALES (CYPERACEAE) FROM EASTERN NORTH AMERICA PE. ROTHROCK patie Environmental Center Lor University Upland, IN 46989-1001 US: A.A. REZNICEK University of Michigan Herbarium North University Building Ann Arbor, MI 48109 U.S.A. ABSTRACT Chromosome counts for seventeen species of Carex section Ovales are reported. T hey range from ”=26 + IV tox 40. Counts for Carex alata, C. hyalina, C. longii, C. ozarkana, C. reniformis, C. suberecta, c tincta, and G, vexans were determined for the first time and included ; ineaploid (sensu lato) series. Four species (C. albolutescens, C. hormathodes, C. straminea, and C. tribulotdes) yielded counts similar to previously published results. On the other hand, two species (C. cumiulata and C. silicea) differed from previous reports and represent possible aneuploidy, while aneuploidy was demonstrated for C. festucacea. RESUMEN Se realizan recuentos cromos6micos de diez y siete especies de Carex seccién Ovales. Su numero varia de 7=26 + IV hasta »=40. Se realizan por primera vez recuentos de Carex alata, C. hyalina, C. longii, C. ozarkana, C. reniformis, C. asia C. t ncluyen tres series gees es — lato). C C Straminea y C. t 1en neta y C. vexans que ee fe hormathodes, tos similares a resultados publicados previamente. Por otro lado, dos especies (C. ae y C. silicea) difieren de los recuentos previos y representan aneuploidia posible, mientras que se demostro6 la existencia de aneuplodia en C. festucaced. INTRODUCTION Carex L. section Ova/es Kunth includes approximately 40 species in east- ern North America with a deserved reputation for being taxonomically difficult. In contrast with other sections of Carex, Gleason and Cronquist (1991) describe species of section Ovales as narrowly defined and more or less confluent. In spite of there being only subtle morphological differences Sipa 17(1): 251-258. 1996 252 Sipa 17(1) between many species of this section, the karyotypes among them vary widely and reinforce the established species concepts. Eastern North American species have haploid numbers ranging from 26 to 40 (Whitkus 1991). For some species in which multiple plants have been studied cytologically, the counts are constant. Such is the case for C. crawfordii Fernald (plants from 9 populations) C. normalis Mack. (4 plants) and C. molesta Mack. ex Bright (3 plants) as well as C. cristatel/a Britton, C. projecta Mack., and C. tribuloides Wahlenb. with 2 plants each (Wahl 1940, Whitkus 1991). On the other hand, other species (C. bebbiz (L.H. Bailey) Fernald, C. bicknellii Britton, C. merritt-fernaldii Mack., and C. tenera Dewey) reportedly have narrow aneuploid, or more specifically, agmatoploid series. One species, the polymorphic C. scoparia Willd., may have a broader aneu- ploid series (Whitkus 1991). Two species studied thus far, Carex festucacea Willd. and C. ene exhibited multivalents or univalents at meiosis metaphase I (Wahl 1940; Tanaka 1942). The frequent aneuploidy in Carex is thought to be a consequence of polycentric chromosomes with diffuse centromeres (Grant 1981). Ifa poly- centric chromosome fragments, even small chromosomal pieces have the — potential of completing normal movements during mitosis and meiosis. Thus, in this particular kind of aneuploidy known as agmatoploidy, changes in chromosome number may be readily preserved. Furthermore, unlike taxa with monocentric chromosomes, these variations in Carex chromosome number do not effect fertility within a taxon (Whitkus 1988). In spite of numerous chromosome studies of these species, many species of Carex section Ovales have yet to be examined cytologically. At the same time, given the known potential for variation in chromosome number within species of this section (Heilborn 1932; Wahl 1940; Tanaka 1942; Moore & Calder 1964; Packer & Whitkus 1982; Whitkus & Packer 1984; Whitkus 1991), additional counts are necessary for those species with only single or few counts. This study contributes to our understanding of karyology within section Ova/les by reexamining 7 taxa and reporting the chromosome counts for 10 taxa for the first time. MATERIALS AND METHODS Living plants were collected during the flowering or fruiting season and cultivated in a sand-peat-perlite potting medium in 1O—15 cm pots under greenhouse conditions. During the growing season, dilute liquid fertilizer was applied approximately monthly. Plants were cold treated (O—5 C) for 3—4 months to stimulate the production of new inflorescences. Immature spikes were collected and preserved in methanol, chloroform, and propionic acid (6:3:2). Within the subsequent 72 hours, anthers were dissected from the spikes and squashed in 2% lactic-acetic-orcein ROTHROCK AND REZNICEK, Chromosome number reports in Carex 253 (Cooperrider & Morrison 1967). Meiosis I chromosome figures were exam- ined from five or more pollen mother cells. Drawings and photographs were made with a Nikon Labophot-2 microscope using phase contrast at LOOOx magnification. Voucher herbarium specimens were typically prepared at the time of field collection. In some cases, where material was being supplied by corre- spondence or field material was not fruiting, voucher specimens were later prepared from greenhouse material. Vouchers, on deposit at the University of Michigan Herbarium (MICH), include drawings and photomicrographs of countable figures. RESULTS AND DISCUSSION The chromosome counts for the 17 taxa reported in this study ranged from 7=26 + IV to »=40 (Table 1), approximating the known range of n= 26 to 45 for the section as a whole (Whitkus 1991). Of the 11 species for which multiple individuals were investigated, three new aneuploid (sensu lato) series were discovered. Of the species being reexamined, four had the same numbers as previous reports, while three others were different, indi- cating possible aneuploidy. Carex alata Torr. The three counts of 7=37 are the first reported for this species. Although the plants were all collected in southwestern Michigan, the habitats sampled were diverse. The Berrien County site was a wet prat- rie opening, the first Kalamazoo County site (#2522) was a Sphagnum bog, and the second Kalamazoo County site (#2527) was a peaty Acer rubrum swamp. The plants varied widely in robustness with those of the Acer rubrum swamp having the most gracile form. Morphologically this species has the obovate perigynia of C. albolutescens Schwein., C. cumulata (L.H. Bailey) Fernald, C. /ongii Mack., and C. ozarkana P. Rothr. & Reznicek, all of which have lower, sometimes markedly lower, chromosome numbers. The 2=37 is identical with that observed for C. straminea Willd. and C. hormathodes Fernald, species that also have the awned pistillate scales and stipitate achenes seen in C. alata. Carex albolutescens Schwein. Both of our plants, from widely differ- ing parts of the range, had an 7=33. Wahl (1940) reported m=33 for an unidentified plant from section Ovales which he listed as C. sp. 15168. Examination of his voucher specimen (at PAC) from Centre County, Penn- sylvania confirmed that it is C. a/bolutescens. This species has often been confused with C. /ongii and C. festucacea (Rothrock 1991). The distinctive chromosome counts support the mor- phological analysis which indicated that these taxa are best recognized as distinct species. Carex bicknellii Britton var. opaca F.J. Herm. An 7=33 was observed 254 Stpa 17(1) TabLe 1. Chromosome number and location of species of Carex examined. Species n Location: Voucher Carex alata Torr. 37 Berrien Co., MI: PE.R. 2531 37 Kalamazoo a MI: PE. is 2522 37 Kalamazoo Co., MI: PE.R. 2527 C. albolutescens Schwein. 43 Berrien Co., MI: PE.R. 2532 33 Columbus Co., NC: A.A. R. 8972 C. bicknellti Britt. var. opaca FJ. Herm. 33 LeFlore Co., OK: PE.R. 2916 C. cf. brevior (Dewey) Mack. 37 Garland Co., AR: PE.R. 2904 C. cumulata (LH. Bailey) Mackenzie 29 Newton Co., IN: PE. R. 2538 29 Hancock Co, ME: A. Dibble s.n. C. festucacea Willd. 34 Tallahatchie ] o., MS: PE.R. 2944 45 Lonoke Co., >: PELR. 2890 C. hormathodes Fernald 37 Hancock Co., ME: A.A.R. 9162 37 Knox Co., ME: A. Dibble s.n, C, hyalina Boott 37 Tunica Co., MS: P.E.R, 2947 C. longti Mack. 28+1V Berkeley Co., SC: R. By ees 52789 3] Flagler Co., PL: PE 2 3] Indian River Co., - aN 2375B 31 Kalamazoo Co., MI: PE.R. 2525 C. ozarkana P. Rothr. & Reznicek 264+1V LeFlore Co., OK: PE.R. 2917 28+I111 Garland Co., AR: PE.R. 2973 31 Perry Co., AR: PE.R. 2914 C. rentformis (L.H. Bailey) Small 40 Clark Co., AR: PE.R. 2909 40 Quitman Co., MS: PER. 2945 40 Stoddard Co. “MO: P.E.R, 2936 C. silicea Olney 37 Hancock Co., ME: A. Dibble s.n. C. stramtnea Willd. 37 Newton Co., IN: P.E.R. 2537 37 Kalamazoo Co., MI: PE.R, 252. 37 Shannon Co., MO: PE.R. 2928 C. suberecta (Olney) Britton 36 Madison Co., IN: PE.R. 2 36 Kalamazoo Co., MI: P.E.R 2528 ROTHROCK AND REZNICEK, Chromosome number reports in Carex 255 TABLE 1. (continued.) Species n Location: Voucher C. tincta Fernald 36+1V Penobscot Co., ME: A.A.R. 9122 C. tribuloides Wahlenb. 35 Lonoke Co., AR: PE.R. 2897 C. vexans EJ. Herm, 33411 Pasco Co., FL: PE.R. 2379 34 Hendry Co., FL: PE.R. 2376 35 Hillsborough Co., FL: PE.R. 2378 35 Indian River Co., FL: PE.R. 2375 for this variety from the south central U.S. Previous counts of 7=37 (Love & Love 1981) and m=38 (Tanaka 1942) were for the typical, more northerly variety. Given the strikingly different chromosomal condition, a reevalua- tion of the taxonomic status of this variety is being undertaken. Carex cf. brevior (Dewey) Mack. This plant from the Ozark Mountains has an 7=37. It is morphologically close to C. brevior and C. molesta but has a notably higher chromosome number. Love and Léve (1981) reported z= 34 for C. brevior and C. molesta. Wahl (1940) also observed v= 34 for C. molesta. Our plant has the wide achenes of C. brevior but the rounded spike bases and ventrally nerved perigynia of C. molesta. Preliminary morphological evaluation suggests that this is an undescribed taxon. Carex cumulata (L.H. Bailey) Mack. The only published count for this species (Love & Love 1981) is m= 28. Our report of 7=29 is based upon two plants from Newton County, Indiana and one from Hancock County, Maine. Carex cumulata is an uncommon species from barren sandy habitats with affinities to C. longiz. Carex festucacea Schkuhr ex Willd. Wahl (1940) reported that this species had a chromosome count of #=33 + HI. Our material, with 2=34 and 7=35, showed no trivalents but does demonstrate an aneuploid series. Carex hormathodes Fernald. Our counts of #=37 from plants of Maine confirmed that reported by Wahl (1940) for a plant from Rehobeth, Dela- ware. Carex hormathodes has sometimes been treated as a variety of C. straminea which also has 2=37. The two seem, however, to consistently differ in habitat and in the shape of the spikes and the beak of the perigynium. Carex hyalina Boott. The count of #=37 is the first for this rare species of northeast Texas, Arkansas, Oklahoma, and Mississippi. Correll and Johnson (1970) suggest that C. brittoniana L.H. Bailey most resembles this species. Unfortunately, no chromosome count exists for that species. Carex hyalina also bears some notable similarity to C. reniformis (L.H. Bailey) 256 Sipa 17(1) Small. Both occur in floodplain woods and have short-creeping rootstocks, blunt pistillate scales, and perigynia with green beak and shoulders and fine papillae. On the other hand, the width of their perigynia and achenes are strikingly different. Carex longit Mack. This widespread member of section Ovales repre- sents another aneuploid series, v=28 + IV and w=31. Our collections in- clude individuals from the heart of its range on the Atlantic Coastal Plain as well as from the Great Lakes region. The comparatively low chromo- some number compares favorably to that of C. a/bolutescens and C. cumulata, species with close morphological affinities to C. /ongiz. Carex ozarkana P. Rothr. & Reznicek. This newly described species, with a narrow geographic range centered on the Ozark and Ouachita Moun- tains of Arkansas and Oklahoma, displays limited morphological variabil- ity. Nevertheless, it has a variable chromosome condition, 7= 26 + IV, 7=28 + I, and #=31. Based upon morphological considerations, Rothrock and Reznicek (1995) concluded that Carex ozarkana is most closely allied with C. longit. Carex reniformis (L.H. Bailey) Small. This relatively distinctive mem- ber of section Ovales gave a constant count of 7=40. The Clark County, Arkansas site is in the eastern Ozark Mountains, while the other two sites are within the Mississippi River embayment. Carex silicea Olney. Two plants from a site in Hancock County, Maine gave n=37. Moore and Calder (1964) have previously reported 7=38 for this species, suggesting possible aneuploidy. Mackenzie (1931) aligned C. siicea with other species having obovate perigynia, including C. a/bolutescens, C. cumulata, and C. longi. Compared to these species, however, Carex silicea has a much higher chromosome number, frequent auricula at the base of its leaf blades, moniliform inflorescences, long pistillate scales, and somewhat papillose perigynia. The relationship of this distinctive coastal sand dune species to other Ova/les species needs further investigation. Carex straminea Willd. in Schkuhr. All plants examined had 7=37, including those from disjunct populations in Shannon County, Missouri and Kalamazoo County, Michigan. These counts confirm that reported by Wahl (1940) under the synonymous name C. richii Mack. Carex suberecta (Olney) Britton. The two counts of 7=36 are the first reported for this fen species. Among those species with proximate chromo- some counts, Carex hormathodes and C. straminea (n= 37) are likely the clos- est relatives of C. swberecta. Carex tincta Fernald. This species is locally frequent in Maine and New Brunswick and is known from scattered localities as far west as northern Hlinois, Michigan, Wisconsin, and Ontario. Its affinicy with other species is unknown. The chromosome count of 7=36 + IV is the first for this species. ROTHROCK AND REZNICEK, Chromosome number reports in Carex 257 Carex tribuloides Wahlenb. Wahl (1940) and Moore and Calder (1964) reported n=35 for northern plants of this species. Our plant, from the south- ern part of the range, also has an »=35 but has the shorter perigynia and smaller spikes of variety sangamonensis Clokey. Carex vexans EJ. Herm. The counts for this Florida endemic revealed another aneuploid series, 7=33 + HI, 7=34, and »=35. Carex vexans, like C. ozarkana, links a narrow geographic distribution and relatively limited morphological variation with variable karyotypes. Carex vexans is probably most closely related to C. alata, with which it was often confused before being described as a distinctive species. Unlike C. alata, C. vexans lacks awns on the pistillate scales and its perigynia have longer, more gradually tapered beaks. ACKNOWLEDGMENTS Our thanks are given to Alison Dibble for providing living material from Maine, Paul Lightfoot for his invaluable assistance in the greenhouse, and to the Indiana Academy of Science and Taylor University Fund for Faculty Scholarship for their financial support of this wor REFERENCES CoopE RRIDER, T.S. and J. ee aaa 1967. Lactic-acetic-orcein as a chromosome stain. ichigan Bot. 6:176- a D.S. and M.C. re 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner Gieason, H.A. and A. Cronguist. 1991. Manual of vascular plants of northeastern United States and et Canada (second edition). New York Botanical Garden, Bronx. Grant, V. 1981. Plant speciation (second edition). Columbia University Press, New York. HEILBorN, O. ie Aneuploidy and polyploidy in Carex. Svensk Bot. Tidskr. 26:137— 146. Love, A. and D. Love. ee Chromosome number reports LX XII. Taxon 30:829-861. Mackenzik, K.K. 1931. Cyperaceae-Cariceae. North Amer. Flora 18(1—3):1—168. Moore, ‘ J. and J.A. CALDER. a cae ncn numbers of Carex species of Canada and Alaska. Canad. J. Bot. 42:1387-1391 Pac KER i: G.and R. Wuirkus. 1982. aa number reports LX XV. Taxon 31:342- 368. Rorurock, PE. 1991. The identity of Carex albolutescens, C. festucacea, and C. longti (Cyperaceae). Rhodora 93:5 1— Rorurock, PE. and A.A. Reznicek. 1996. A new species of Carex section Ovales (Cy peraceae) occurring in the Ozark Mountain region. Brittonia 48:104—110. ANAKA, N. 1942. Chromosome studies in Cyperaceae. XX. Chromosome numbers of Carex (Vignea - II). Med. & Biol. 2:220-224. Wau, H.A. 1940. Chromosome numbers and meiosis in the genus Carex. Amer. J. Bot. Wuirkus, R. 1988. Experimental hybridization among chromosome races of Carex pachystachya and the related species C. macloviana and C. pres/ii (Cyperaceae). Syst. Bot. 13:146-153. 258 Sipa 17(1) «991, Chromosome counts of Carex section Ovales. Bot. Gaz. (Crawfordsville) 152:224-230. Wauirkus, R. and J.G. Packer. 1984. A contribution to the taxonomy of the nite Cadi aggregate (Cyperaceae) in western Canada and Alaska. Canad. J. Bot. 62:1592- DOCUMENTED CHROMOSOME NUMBERS 1996:2. MISCELLANEOUS U.S.A. AND MEXICAN SPECIES, MOSTLY ASTERACEAE ZAIMING ZHAO Department of Botany University of Texas Austin, TX 78713, U.S.A. ABSTRACT Chromosome counts are reported for over eighty collections of iene plants from the U.S.A. and Mexico, representing 59 species in 39 genera of 10 families RESUMEN e presentan recuentos cromosémicos de mas de ochenta recolecciones de plantas con flores de U.S. México, pertenecientes a 59 especies, de 39 géneros, de 10 familias. g The following chromosome counts (Table 1) are documented by speci- mens deposited at the herbarium of the University of Texas, Austin (TEX). Previously uncounted taxa are marked by an asterisk(*). ouble aster- isk(**) indicates a new number for the species. Fedorov (1969) and the standard indices of plant chromosome numbers published since that opus (through 1991 :cf.introduction, Goldblatt & Johnson 1994) were consulted to ascertain previous counts for the taxa concerned. METHODS Chromosome counts were made from pollen parent cells from floral bud material collected in the field and fixed in a modified Carnoy’s solution(4:3: 1; chloroform, absolute ethanol, glacial acetic acid), using standard squash procedures. DISCUSSION Astranthium.—DeJong (1965) reported the base chromosome num- bers of this genus to be x=3, 4, and 5. His counts for A. splendens (2n=18) are hexaploid. My counts of this species are 27=12 and 2n=24, the first such counts for the species. The two counts were obtained from one head of a single plant and displayed 6 bivalents and 12 bivalents respectively. Tradescantia.—Chromosome numbers for the five species listed in Table 1 are consistent with numbers for species of Tradescantia generally. Both diploidy and tetraploidy are well documented in various species of Stipa 17(1): 259-263. 1996 260 Stipa 17(1) TasLe 1. Chromosome numbers of miscellaneous U.S.A. and Mexican species, mostly Asteraceae. var. daxacana (DC.) Grashoff Family/Species Voucher! Chromosome number(27) ACEAE ~~ Sphaer ocephalus UT: Washington T 95-127 18 Gra . Gra phan riddellti Torr. & A. Gray i ee P 1561 10 ranthium splendens De rik .N 7468 12,24 a tmultiradiata Harv . Gray oe ae Me T 95-129 64 Chaetopappa bellioides (A. jee Shinners = MEX:N.L 16 Chaetopappa bellioides (A. Gray) Shinners = TX:Val Verde T 95-13 16 Chaetopappa tmberbis (A. Gray) Nesom TX: Wilson T 94-4 16 Conoclinium betonicifolium (Miller) King MEX:Tamp. PA 7269 20 Robinson var. betonicifolium Miller Conoclininm betonicifolium (Miller) MEX:N.L. T.F. PA 7404 20 King & Robinson var. integrifolinm ae *Conoclininum sp. nov. T.F. Patterson MEX:Tamp. PA 7 20 (in prep.) MEX:Dur. PA vee 20 Coreopsis wrightii A, Gray TX:Llano Z 944 24 Dyssodia penta haeta (DC.) B.L. Rob. MEX:Coa. N 7650 ca.26 r. pentachaet TX:Dimmic T 94-10 ca.26 Dyn venuilebe (DC.) B.L. Rob. TX:Fayette M 1946 26 ar. tenutloba ee tenuis Yorr. & A. Gray TX:Liberty M 1959 7-9 Il + 22-181 *Erigeron veracruzensis Neso MEX:Coa. N 7486 18 — aestivalis (Wale.) = Rock TX:Austin T 94-86 34 TX:Lee T 95-69 34 Gaillardia anton F. Gay TX: aoe T 95-88 34 Gaillardia pulchella Fouger. MEX:Tamp. M 1790 34 var. drumondii Hook *Grindelta obvatifolia S.F. Blake MEX:N.L. N 7485 12 Grindelia tenella Steyerm. MEX:Tamp. N 7451 & 7452 12 EK Heterotheca mucronata Harms ex MEX:N.L. N 7469 36 urner , and Global Stat Taxa Distr. KY ™ Nat. Global Aster paludosus Ait. ssp. bemisphericus (Alex.) Crong. T E - - G 2 ROUND-HEAD ASTER (Asteraceae) Carex alata Torrey C T - - G 5 INGED-FRUITED SEDGE (Cyperaceae) Carex decomposita Muhl. M T - - G > YPRESS-KNEE SEDGE (Cyperaceae) Gymnopogon ambiguus (Michx.) BSP. C,M S - - G 5) EARDGRASS (Poaceae) Hieracium longipilum Torrey M,S,T OT 5 - G4/GS ONG-HAIRED HAWKWEED (Asteraceae) Hydrastis canadensis L. M,S = Tr = G 4 GOLDENSEAL (Ranunculaceae) ee cinerea L.. S S T C2 G3/G4 peace Malus sti (Aiton) Mich C S S = G 5 N CRABAPPLE ee ‘od ene Old alli Scribn C,T S S S G 4 MOOTH MUHLY (Poaceae) Ocnothera linifolia N utt. C,M,S,T E = - G 5 THREAD-LEAVED SUNDROPS (Onagraceae) Panax quinquefolius L. M,S - T - G 4 INSENG ia Phacelia ranunculacea (Nutt.) Const M AS S — G 4 LUE SCORPIONWEED ae silineeasy Platanthera peramoena (Gray) Gray M = T a G 5 URPLE FRINGELESS ORCHID (Orchidaceae) Populus talons Michx M - S - G 5 BIG-TOOTH AS er Galicacene) Prenanthes aspera Michx, E E E - G 4 OUGH RATTLESNAKE ROOT (Asteraceae) Prenanthes barbata (T.&G.) Mil. M E S C2 G2/G3 ATTLESNAKE ROOT (Asteraceae) Rudbeckia ae Pursh C,M,S,T E T = G 5 SWEET CONEHOWER (Asteraceae) Scleria He Nica: T E = - G 5 NutrusH (Cyperaceae) Silphium laciniatum L. M,S,C E T ~ G 5 OMPASS-PLANT (Asteraceae) Silphium pinnatifidum Elliott M - T a G 3 RAIRIE DOCK (Asteraceae) Tomanthera auriculata (Michx.) Raf. M =? E C2 G 2 SARLEAF FOXGLOVE (Scrophulariaceae) 'Known county distribution at FCMR given as: C = Christian Co., KY; M = Montgomery Co., TN; S = Stewart Co., TN; T = Trigg Y. 7E = endangered, T = thr esiened: S = eel: concern. 32 = a candidate for federal listing. Sipa 17(1): 271. 1996 272 Stipa 17(1) ‘Global rankings: G2 2 = imperiled globally because of pie or because of some factors making it very vulnerable to extinction throughout 1 Its range, G3 ‘ither very rare and local throughouc i its ranpe or found locally (even abundantly) in a restricted range, or because of some other factors making it vulnerable to extinction throughout its range; G4 = apparently secure globally, though it may be quite rare in parts of its range, especially at the ee G5 = demonstrably secure globally, though t may be quite rare in parts of its range, especially at the periphery. Tomanthera auriculata is not known from Kentuc ky and thus not listed there. and 1s infrequent to rare throughout its range. Carex decomposita (numerous clumps at one site) and Platanthera peramoena (several small populations) range over much of eastern U.S., but are infrequent to rare in occurrence. Juglans cinerea (five trees and numerous juveniles at one site) also is wide- ranging in eastern U.S., but is in general decline and now extirpated from several areas (Anderson & LaMadeleine 1978; Rink 1990). Lastly, Hydrastis canadensis (several large populations) and Panax guinguefolius (several small populations), are wide-ranging but rare, primarily because of loss of habi- tat and commercial exploitation. State Records Two non-native taxa new to the known flora of Tennessee were found. Nymphoides peltata (Gmel.) Kuntze (YELLOW FLOATING-HEART, Meny- anthaceae) is a rooted, floating-leaved aquatic native to southern Europe and Asia Minor. It was introduced into this country for cultivation in gar- den pools, but has become sporadically naturalized over much of eastern U.S. However, it has not been reported for states south of Virginia or east of Louisiana, Arkansas, and Missouri (Godfrey & Wooten 1981). This report, based on an extensive population in a Reservation pond, adds not only the species but also the genus and family to che known Tennessee flora. oucher specimens: TENNESSEE. Montgomery Co.: pond by FCMR golf course, 19 Jun 1993, Wofford 93-23 (TENN); 6 Jun 1994, Chester 13040 (APSC). Richardia brasiliensis (Moq.) Gomez (without vernacular, Rubiaceae) is a diffuse, pilose annual or perennial native to South America. It has be- come a naturalized weed, primarily on the Coastal Plain from Texas to Florida and northward to Virginia. FCMR plants were in sandy soil of a disturbed field where rareness indicates that it is either a recent introduc- tion or is slow to become naturalized in the area. Voucher specimen: TENNESSEE. Montgomery Co.: FCMR, field on N side of Orien- tal Village Road at junction with Hellcat Road, 20 Oct 1994, Wofford & Chester 13150 (TENN) Other Noteworthy Taxa Two FCMR taxa are noteworthy because of their limited distribution in Tennessee. Sipa 17(1): 272. 1996 Notes 273 Phragmites australis (Cav.) Trin. ex Steud. (REED-GRAsS, Poaceae) 1s often weedy and occurs over much of the U.S. except for several southern states. Previous Tennessee reports are from Henry County, northern west Tennessee (DeSelm et al. 1994) Voucher specimen: TENNESSEE. Montgomery Co.: beaver-formed marsh at jct of ordan Springs Road and Oriental Village Road, FCMR, 28 Aug 1993, Wofford G Chester 12996 (APSC, TENN, VDB Psoralea onobrychis (Nutt.) Rydb. (Scurr-pea, Fabaceae) ranges from Ohio and Kentucky to Iowa and Missouri, and also is found in West Vir- ginia, Tennessee, and South Carolina. Isely (1990) reported it from one county each in North Carolina, South Carolina, and Virginia, and from two counties in Tennessee, noting that “material seen for reports from NC and TN is represented only by collections from the last century.” The FCMR collection is from a dense stand of several hundred plants along a roadside- barren. Voucher specimen: TENNESSEE. Montgomery Co.: barrens on N side of Ghost Corp Trail one mile E of Palmyra Road, FCMR, 2 Jul 1993, Chester 12985 (APSC, TENN, VDB). ACKNOWLEDGMENTS This research (1993-1994) was funded by the Tennessee Chapter of the Nature Conservancy through the Legacy Resource Management Program. Numerous people at FCMR provided significant assistance, especially Eu- gene Zirkle, FCMR Coordinator for Land Condition Trend Analysis, who made studies possible in 1995. —Edward W. Chester, Department of Biology, Austin Peay State University, Clarksville, TN 37044, U.S.A.; B. Eugene Wofford, Department of Botany, The University of Tennessee, Knoxville, TN, 37996, U.S.A.; Landon E. McKinney, Kentucky State Nature Preserves Commission, 501 Schenkel Lane, Frankfort, KY 6 S.A.; David Campbell, Tennessee Field Office, The Nature Conser- vancy, 50 Vantage Way, Suite 250, Nashville, TN 37215, U.S.A. REFERENCES ANDERSON, _and L.A. LAMApELEINE. 1978. The distribution of butternut decline in the eastern a States. Forest Survey ee S-3-78. U.S. Dept. Agric., Forest Ser- vice, Washington, D.C. Cuuane, T.I. and L. Constance. 1977. Cytogeography of Phacelia ranunculacea (Hydrophyllaceae). Rhodora 79:11 22, DeSeLm, H.R., B.E. Worrorp, R. Kra, and E.W. CHesTer. 1994 . An annotated list of grasses (Poaceae, aa of Tennessee. Castanea 59:338— — ria Stpa 17(1): 273. 1996 274 Sipa 17(1) FisHer, TR. and J.M. Speer. 1978. Systematic studies in the genus Si/phinm: possible ori- gin of S. pinnatifidum Ell. (Compositae). Environmental Physiology and Ecology of Plants 1978:451— GLEASON HA. and A. CRonquisr. 1991. Manual of vascular plants of northeastern United Scates and adjacent Canada. New York Botanical Garden, Bronx. Gopekey, R.K. and J.W. Wooren. 1981. Aquatic and wetland plants of southeastern United States. Volume 1, Dicotyledons. University of Georgia Press, Athens. Isety, D. 1990. Vascular flora of the southeastern United S . Volume 3, Part 2, Leguminosae (F sei: University of North Carolina Press, C Chapel 1 Hall. KENTUCKY STATE NATURE PRESERVES COMMISSION. 1992. Endangered, threatened, and spe- cial concern cei and animals of Kentucky. Unpublished lise. ee Kentucky. Nickens, E. 1993. ae conservation. Nat. Conservancy 43(2):24—29. QUARTERMAN, E. and R.L. Powerit. 1978. Potential sieges natural landmarks on the pe Low anon . S. Dept. Interior, Washington, D.C. RINK, G. 1990. Juglans cinerea 1.., ou tternut. In: Burns, R. M. and B. H. Honka cords.). ae of North America, 2: hardwoods. Agriculture Hand Agric., Forest Service, Washington, D.C. Pp. 386-390. TENNESSEE DEPARTMENT OF ENVIRONMENT AND CONSERVATION. = a (tech. s00k 654. U.S. Dept ary 1994. Rare plants of the State of Tennessee, 17 May 1994. Ecological Services Division, Nashville, Tennessee JS. Fish AND WILDLIFE Servicr. 1993. Plant taxa for listing as endangered or threatened species: Notice of review. Federal Register 58(188):5 1144-51190 Worrorb, B.E. and R. Krat. 1993. Checklist of the vascular pl Bot. Misc. 10. _— ants of Tennessee. Sida, Sipa 17(1): 274. 1996 NOTES 275 A REPORT OF FOUR EXOTIC CYPERUS (CYPERACEAE) SPECIES NEW TO FLORIDA, U.S.A. Cyperus alopecuroides Rottb., C. distans L.f., C. prolifer Lam., and C. sphacelatus Rottb. are reported new to Florida. All four taxa are exotic weeds and ap- pear to be naturalized. Cyperus alopecuroides, C. distans, C. sphacelatus are newly introduced, and C. prolifer is apparently recently naturalized. This report of C. alopecuroides is the second for the western hemisphere. General distributional data and pest potential are discussed. Each taxon 1s com- pared with similar species, and collection data are given. In the past decade a number of Cyperus species have been reported new to Florida: C. difformis L., C. echinatus (L.) Wood, C. entrerzanus Bockeler, C. hystricinus Fernald, C. pilosus Vahl, and C. reflexus Vahl (Burkhalter 1984, 1985; Carter 1988, 1990; Carter & Jones 1991; Anderson 1991). Recently, specimens from central and southern Florida have been identified as Cyperus alopecuroides Rottb., C. distans L.£., C. prolifer Lam., and C. sphacelatus Rottb. Cyperus alopecuroides, C. distans, and C. sphacelatus are apparently casual in- troductions, and C. prolifer is likely an escape from cultivation. All four species are weeds (Reed 1977; Haffliger et al. 1982) with potential to be- come pests in other subtropical areas of the southeastern United States, and all appear to be naturalized in Florida. None has previously been reported from Florida (Ward 1968; Long & Lakela 1971; Godfrey & Wooten 1979, Wunderlin 1982; Clewell 1985), and only C. distans (Small 1933; Kiikenchal 1935: Radford, Ahles & Bell 1968; Beal 1977; Kartesz 1994) and C. sphacelatus (Mohr 1901; McGivney 1938) have previously been reported from the United States. Cyperus alopecuroides Rottb., Descr. Pl. Rar. Progr. 20. 1772. Cyperus alopecuroides is widely distributed in tropical and subtropical re- gions of the Old World, e.g., northern and tropical Africa, Madagascar, India, Ceylon, Indo-China, Malaysia and northern Australia; in the New World it is known only from Guadeloupe in the West Indies (Kiikenthal 1936; Koyama 1985). In habit and general inflorescence pattern, C. alopecuorides resembles the tropical species C. imbricatus Retz., and both taxa were placed in section Exaltati by Kiikenthal (1935). Its foliage and stout reddish base somewhat resemble those of C. erythrorhizos Muhl. Cyperus alopecuroides is a robust aquatic to 1.5 m high. Its size in combination with other characters make it a striking plant in the field: broad bracts and leaf blades (to 15 mm wide) with contrasting surfaces (adaxial light-green, abaxial glaucous), and a branched inflorescence with spikes of densely clustered golden-brown spike- Sioa 17(1): 275. 1996 276 Sipa 17(1) lets. Although its affinities are clearly with subgenus Cyperus (Kiikenthal 1935; Koyama 1985), it has characteristics that seem to defy placement there: namely, a bicarpellate gynoecium with two stigmas and a lenticular achene with face adjacent to rachilla. When taken alone, the gynoecium and fruit characteristics seem to indicate a relationship with subgenus Juncellus (Clarke 1908). However, Koyama (1985) has observed both bi- and trigynous pistils in the same inflorescence, and this, in combination with other characteristics, supports inclusion of C. alopecuroides in subge- nus Cyperus. Illustrations of C. a/opecuroides are in Hiffliger et al. (1982, p. A6: as Juncellus alopecuroides) and Haines & Lye (1983, p. 181, Fig. 348). In Florida, C. alopecuroides was growing as an emergent in a reclamation wetland in an abandoned phosphate pit, where it was locally abundant in — floating mats and was less common in shallow water along the wetland edges. Associated taxa include Eleocharis interstincta (Vahl) R. & S., Sagittaria sp., and Scirpus californicus (Meyer) Steud. This exotic Cyperus may have been introduced as a contaminant in nursery stock used for revegetation and has probably been in place at least two years (Tom Nix, IHS Environ- mental Consulrants, pers. comm. 1994). Several hundred plants were ob- served in an area of about six ha. The plant’s ability to form extensive stands and floating mats here suggests invasive potential should it escape from the reclamation site. Close monitoring is called for in the area. The follow- ing data are for first collections from Florida and the United States. Voucher specimens: U.S.A. FLORIDA. Polk Co.: ca. 3 mi W of Fort Meade, N from county road 630 on ae sa by Tiger Bay, reclamation wetland #SP-6 on U.S. Agrichem property, NW 1/4 Sec T31S R25E, 15 Sep 1993, T. Nix with M. Phillips s.n. (FSU, age and Para at ¥VSC); 11 Feb 1994, Carter 11605 with Mears & Phillips (FSU, VSC, to be Ae wee 24 May 1995, Burks 1018 with Nix, Phillips & ee (FSU, V S Cyperus distans L.f., Suppl. pl. 103. 1781. Cyperus distans is a pantropical weed inhabiting marshes, canal banks and ditches in Africa, India, Sri Lanka, southeastern Asia, Malaysia, south- ern China, the Philippines, the Caribbean islands, central America, Mexico, and tropical South America (Clarke 1900; Uittien 1932; Kiikenthal 1935; Koyama 1985; Adams 1994; Tucker 1994). Cyperus distans is rare in south- eastern United States, where it has been reported from coastal North Caro- lina (Kiikenthal 1935; McGivney 1938; Radford, Ahles & Bell 1968: Beal 1977) and from Georgia (Small 1933; Kiikenthal 1935; Beal 1977). The citations of C. distans from Georgia are doubtful; neither have we seen speci- mens nor was it reported by Jones & Coile (1988). Cyperus distans has trifid styles and trigonous achenes and, in habit, re- Stipa 17(1): 276. 1996 Notes 077 sembles C. odoratus L. or C. stigosus L., from which it can be distinguished by its looser spikes and more delicate spikelets. The following combina- tion of characteristics distinguishes C. distans from all other North Ameri- can Cyperus: scales ascending, remote, with 3-5 nerved greenish keels, sanguineus to reddish brown nerveless sides, and scarious emarginate tips. Cyperus distans is illustrated in Haffliger et al. (1982, p. 17) and in Beal (1977, p. 117). The following data provide the first documentation of C. distans from Florida. Voucher specimens: U.S.A. FLORIDA. St. Lucie Co.: ca. 8 mi E of Okeechobee Co. line on FL 68, ca. 9 mi W of Ft. Pierce, T35S R38E Sec. 9, 26 Sep 1985, Wunderlin, Hansen & van Hoek 101 38 (TENN, pers. herb. C.T. Bryson, fragment and photocopy at VSC). Cyperus prolifer Lam., Ill. 1.147. 1791. Cyperus prolifer inhabits marshes, marshy shores, and swampy stream banks in eastern Africa (Kiikenthal 1935; Bailey & Bailey 1976; Haines & Lye 1983). Cyperus prolifer is commonly known as dwarf papyrus or miniature papyrus and is sold as an ornamental for use in water gardens (Bailey & Bailey 1976). It has been variously listed as Cyperus haspan viviparus (Watkins & Sheehan 1975; Graf 1985), C. papyrus cv. “nanus” (Bailey & Bailey 1976), and C. isocladus Kunth (Bailey & Bailey 1976; Everett 1981). Hluscrations of C. prolifer are in Kiikenthal (1935, p. 11, Fig. 2A) and Haines and Lye (1983, p. 171, Fig. 327). In addition to the typical variety of C. prolifer, Kiikenthal (1935) recog- nized C. prolifer var. isocladus (Kunth) Kiikenthal (=C. ssocladus Kunth). According to Kiikenthal (1935), in var. ésocladus the culm is trigonous and apically triquetrous and scabrid, while in the typical variety che culm is terete and apically obtusely trigonous and smooth. Our Floridian plants are consistent with Kiikenthal’s concept of C. prolifer var. prolifer. In habit C. prolifer superficially resembles C. baspan L., and Kiikenthal (1936) classified both species in section Haspani. Cyperus prolifer can be readily distinguished from C. haspan by its thick rhizome and inflorescence of 50— 100 rays, with the rays of more-or-less uniform length. In central Florida C. prolifer has apparently escaped from cultivation and is found growing in floating mats and occasionally along margins of natu- ral limesink lakes, where it is associated with Oxycaryum cubense (Poepp. & Kunth in Kunth) Lye {=Scirpus cubensis Poepp. & Kunth in Kunth}. The landowner of lakefront property adjacent to an infested area of Lake Hunt- ley indicated “dwarf papyrus” had been cultivated there in a water garden for about eight years and that during that period it had spread into the lake where it had become an invasive pest (M. Hout, pers. comm. 1994). In Sipa 17(1); 277. 1996 278 SIDA 17(1) 1991 the landowner sought and obtained a control permit from the Florida Department of Environmental Protection (FDEP), which allowed hand- removal or treatment with glyphosate (trademark Rodeo). After two years of growing-season treatment, the original Lake Huntley population was greatly reduced in size. However, several other colonies have been found on the lake (David Demmi, FDEP, pers. comm. 1994). This population and others found more recently in nearby Lake Francis and Lake Lotela and in Lake Howard should be monitored, and the species sought in systematic routine surveys in central Florida lakes. If C. prolifer continues to spread from cultivation, it should probably be placed on the state list of prohib- ited aquatic plants (Ramey 1990). The following data provide documenta- tion for C. profifer in Florida. Voucher specimens: U.S.A. FLORIDA. Highlands Co.: city of Lake Placid, W and E sides of southernmost shore of Lake Huntley, juste W of jct of ee Avenue and county road 29, SE 1/4 Sec. 8, T36S R29E, 6 Jul 1993, D. Demmi s.n. (FSU, VSC); 5 Oct 1993, D. Demmi s.n. (FSU, VSC); 13 Feb 1994, Carter 11640 & Mears (FSU, SWSL, VDB, VSC, others to be distributed); shore of Lake Francis, W of Hwy. US 27, ca. 3 mi N of city of Lake Placid, N 1/2 Sec 22, T36S R29E, 14 Sep 1993, D. Demmi s.n. (FSU, VSC); along NW shore of Lake Lotela near boat ramp near Ci of hwys SR 17 and CR 17A, in city of Avon Park, NW 1/4 Sec. 28 T33S R28E, 2 Nov 1994, D. Demmi s.n. (FSU). Polk Co.: ter Haven, N shore of Lake Howard ae Lake —— Drive NW and shore of lake. locally common between 8th St NW and Mirror Terrace, 4 May 1994, Mears 94-30 VSC). Cyperus sphacelatus Rottb., Descr. Pl. Rar. Progr. 21. 1772. Cyperus sphacelatus is a widely distributed tropical and subtropical taxon, known from eastern Africa, Ceylon, Malaysia, northern Australia (Queensland), Tahiti, South America, Central America, and the Caribbean (Clarke 1900; Britton 1907; Uietien 1932; Kiikenthal 1936; Haines & Lye 1983; Tucker 1983; Koyama 1985). Mohr (1901) reported C. sphacelatus from ballast heaps in Mobile, Alabama, and McGivney (1938, p. 51) cited it among excluded taxa. Reed (1977) listed C. sphacelatus as an economi- cally important foreign weed with potential for becoming a problem in the United States. Additional field work is needed to determine its total range in southern Florida and its pest potential there. Although C. sphacelatus is a distinctive taxon, it might be confused with C. esculentus L., C. filicinus Vahl, or C. rotundus L., with which it bears a superficial similarity. The following combination of characteristics readily distinguishes C. sphacelatus from other North American Cyperus spp.: an- nual cespitose habit; triquetrous achene; diffuse inflorescence with flattened spikelets; and variegated floral scales pale, nearly white, each with two conspicuous reddish patches. Ilustrations of C. sphacelatus are in Hiiffliger SIDA 17(1): 278. 1996 Notes 279 TABLE 1. Geography and variation in scale length in Cyperus sphacelatus. Location Scale length Specimen or reference citation Florida, U.S.A. 2.0—2. m (mean= 2.48, N=50) Carter 11627 & Mears eee an Republic 2.25—2.6 mm (mean=2.43, N=25) Carter 5190 & Garcia Cameroon 2.4-2.8 mm Goetghebeur 4908 ‘Tanzania 2. 4—2.8 mm Haines & Lye (1983) Ceylon 2.0—2.8 mm Koyama (1985) Costa Rica and Panama (2.2-)3.0—4.0(—4.4) mm Tucker (1983) Central America 2.5—-4.0 mm Adams (1994) et al. (1982, p. 25), Haines & Lye (1983, p. 195, Fig. 383), and Reed LOTT: 180). The floral scales are longer in Central American specimens than in West Indian ones (Tucker 1983). Scale length in our Floridian specimens ranges from 2.0 to 2.8 mm, well within the range seen in West Indian specimens and mostly shorter than reported from Central America (see Table 1). Thus, C. sphacelatus was likely introduced into Florida from the West Indies. The habitat of C. sphacelatus was described by Reed (1977) as “disturbed ground, in damp, grassy places” and by Tucker (1983) as “beaches, shores of rivers, moist thickets, fields, and disturbed sunny sites.” In southern Florida, C. sphacelatus is a locally common heliophyte in moderately well drained soil along an open, sloping road berm adjacent to a swamp, where it is associated with Amaranthus spinosa L., Ambrosia artemistifolia L., Argemone mexicana L., Bidens alba (L.) DC., Catharanthus rosens (L.) G. Don, Chamaesyce hyssopifolia (L.) Small, Chromolaena odorata (L.) King & Robins., C onoclinium coelestinum (L.) DC., Cyperus filicinus Vahl, C. polystachyos Rottb., C. surinamensis Rottb., Dactyloctenium aegyptium (L.) Beauv., Eclipta alba (L.) , Emilia fosbergii Nicols., Erechtites hieracifolia (L.) Raf., KylMinga brevifolia L., Macroptilium lathyroides (L.) Urban, Parthenium hysterophorus L. Pennisetum sp., Physalis sp., Pluchea odorata (L.) Cass, Poinsettia cyathophora (Murr.) Kl. & Gke., P. eas dla (L.) Kl. & Gke., Ricinus communis L., Scoparia dulcis L., Sida acuta Burm. f., Sorghum halepense (L.) Pers., Stachytarpheta jamaivensis (L.) Vahl, Trema micrantha (L.) Blume, Tridens flavus (L.) Hitchc., and Urena lobata L. The following data provide documenta- tion for C. sphacelatus in Florida. Voucher specimens: U.S.A. FLORIDA. Dade Co.: 0.58 mi S of jct hwys US 41 and FL oe shoulder of hwy FL edge of swamp, locally common, 12 Feb 1994, Carter 1627 & Mears (FSU, SWSL, VSC, others to * distanced): 1.58 mi S of jct hwys US 41 - FL 997, shoulder of hwy FL 997, edge of swamp, locally common, 25 Dec 1993, R.L. Mears s.n. (VSC); 12 Feb 1994, Carter 11634 & Mears (FSU, SWSL, VSC, others to be distributed). Hillsborough Co.: E of jct Dale Mabry Drive and Lakeview Drive, 0.55 mi jen Sipa 17(1): 279. 1996 280 Sipa 17(1) N ject Lakeview Drive and Lake Heather Drive by Lake Heather Drive, along E side of Lake Heather Drive, 10 Dec 1994, K. eee R.L. Mears s.n. (VSC). Other specimens examined: DOM CAN REPUBLIC. Distrito Santo Domingo: a orilla del Rio Manoguayabo, en el ob ee de Manoguayabo, 12 Dec 1986, Carter 5190 & Garcia (VSC). CAMEROON. Litt. Prov.: Loum Chantiers, Moungo River beds, near large bridge, 5 km NW of Ebonji, 27 oe 1983, Goetehebenr 4908 (VSC). ACKNOWLEDGMENTS Matt Phillips and David Demmi, Florida Department of Environmental Protection (FDEP), Bartow, and Vern McNeilus (TENN) are gratefully acknowledged for providing specimens of Cyperus alopecuroides, C. prolifer and C. distans for determination. Matt Phillips and the Florida Depart- ment of Environmental Protection provided logistical support for the au- thors’ field work, and the Faculty Research Fund of Valdosta State Univer- sity provided funding for Carter’s field work. Richard Carter, Her se, Department of Biology, Valdosta State University, Valdosta, GA 31698, U.S.A.; Randy L. Mears, Department of Biology, 235 Moore Building, Eastern eae University, Richmond, KY 40475, U.S.A.; Kathleen Craddock Burks, Bureau of Aquatic Plant Management, Florida De- partment of Environmental Protection, 3917 Commonwealth Blud., MS 710, Tal- lahassee, FL 32399, U.S.A.; Charles T. Bryson, USDA, ARS, Southern Weed Science ae P.O. Box 350, Stoneville, MS 38776, U.S.A. REFERENCES ei C.D. 1994. Cyperus. In: Flora Mesoamericana, Vol. 6: Alismataceae a Cyperaceae. . Davidse, M. Sousa S., and A. O. Chater, eds. Universidad Nacional Auténoma de ee Ciudad Universitaria. Pp. 423—44( ANDERSON, L.C. 1991. Noteworthy plants from north Florida. V. Sida 14:467—474 Battery, L.H. and E.Z. Bailey. 1976. Hortus third. Macmillan Publishing Co., Inc., New k. Brat, E.O. 1977. A manual of marsh and aquatic vascular plants of North Carolina with ania data. Tech. Bul. 247. North C ae, Agriculcural | Research Service, Raleigh. Brirron, N.L. 1907 ee of Se aica. Dep. Agric. Bull. Vol. V, Suppl. 1, W. Fawcett, ed. (Contr. N.Y. Bot. Gard. No. 97). Kingston. Jamaics BURKHALTER, J.R. 1984. Additions to the vascular flora of Florida. Castanea 49:180—186. BURKHALTER, J.R. 1985. Aletris farinosa, Cyperus difformis, and Cyperus pilosus new tor Florida. Sida 11:247-248. Carter, R. sea C ela piles (Cyperaceae) new to Florida. Sida 13:118—-119. Carter, R. 1990. ‘anus (Cyperaceae), an overlooked species in temperate North America. tie 14: 69-7 Carver, R. and Jones, SD. 1991 . Additional records of Cyperus entrerianus (Cyperaceae) in the United States. Sida 14:615-616. CLARKE, C.B. 1900. Cyperaceae. In: Symbollae Antillanae, Vol. 2, I. Urban, ed., Fratres Borntraeger, Leipzig. Pp. 8—160. Sipa 17(1): 280. 1996 NOTES 281 Ciarke, C.B. 1908. New genera and species of Cyperaceae. Bull. Misc. Inform. 8:1—196 Ciewett, A.E. 1985. Guide to the vascular plants of the Florida panhandle. University Presses of ae Tallahassee Everett, T.H. 1981. The New York Botanical Garden illustrated encyclopedia of horti- culture. Vol. 3. pe Publishing, Inc., New Goprrey, R.K. and J.W. Wooten. 1979. Aquatic aa wetland nies of southeastern United States, a ane es of Georgia ce Athe Grat, A.B . Exotica series 4. 12th Ed. Vols. 1 & 2. Roehrs Company Inc., E. Ruth- erford, New ae HaArflicer, E., U. Ktun, L. Hamer, C.D.K. Cook, R. FADE weeds 3. Ciba-Geigy ce Basle, Switzerland. N, and F. Speta. 1982. Monocot Haines, R.W. and K.A. Lye. 1983. The sedges and rushes of east Africa. Ease African Natural His sce Nairobi, Ken Jones, $.B. and N.C. Corte. 1988. The disceibution of the vascular flora of Georgia. De- partment ei me University of Georgia, Athe Kartesz, J.T. 1994. A a checklist of a feces flora of the United States, Canada, ah Greenland. Vol. I. 2nd Ed. Timber Press, Bay Orego Koyama, T. 1985. Cyperaceae. In: Flora of Ceylon, Vol. Fosberg, eds. Amerind Publishing Co. Pvt. L td., New Ae Pp. 125- KUKENTHAL, G. 1935-1936. Cyperaceae-Scirpoideae-Cypereae. In: Das oe 20 (Heft 101):1-671. L. Diels, ed. Verlag von Wilhelm Engelmann, Leipzi Lone, R.W. and O. Laketa. 1971. A flora of tropical Florida. University of en Press, Coral Gables, Florida. McGivnry, M.V. 1938. A revision of the subgenus Excyperus found in the United States. Cath. Univ. Biol. Ser. 26.1— Mou in, C. 1901. Plant life of oer Contr. U. g Office, Washington, D.C. aes A.E., H.E. Antes and C.R. Brit. 1968. Manual of the vascular flora of the Caro- linas. The Upiversity of North Carolina Press, Chapel Hill. Ramey, V. (ed). ae Florida Prouibitee aquatic plants. Florida Department of Natural Resources, Bureau of Aquatic Plant Management, Tallahass Reep, C.F. 1977 rene important foreign weeds. nena Handbook No. 498. United States Department of Agriculture. U.S. Government Printing Office, Washing- DC R. Passa and E.R. h IV. S. National Herb. 6. Government Print- ton, SMALL, J.K Chapel Hill. TUCKER, _ 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. 85 1933. Manual of the southeastern flora. University of North Carolina Press, Syst. Bot. Monog. 2:1—85. Tucker, G.C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monog. 43:1—213. Urrrien, H. 1932 (1966 reprinc). Cyperaceae. In: Plow of aie Vol. Pulle, ed. Royal Tropical Institute, Amsterdam. Pp. 72-14 Warp, D.B. 1968. Checklist of vascular flora of Florida. Paik I. Florida Agric. Exp. Sta. Inst. Food Agr. Sci. Bull. 726:1—72. ATKINS, J.V. and T.J. SHEEHAN. i 5. Florida landscape plants. The University Presses of I, Part 1. A. orida, Gainesville aches as R.P. 1982. Guide to the vascular plants of central Florida. University Presses of Florida, Tampa. Sipa 17(1): 281. 1996 282 Sipa 17(1) CYPHOMERIS GYPSOPHILOIDES VAR. STEWARTIL A SYNONYM OF A VARIABLE C. GYPSOPHILOIDES (NYCTAGINACEAE) In a recent paper, Mahre and Spellenberg (1995) reported upon multivari- ate analyses of the small genus Cyphomeris, and provided a taxonomy that recognized only two variable species, C. crassifolia (Standl.) Stand. and C. gypsophiloides (Marc. & Gal.) Standl. One of the goals of that study was to search for geographically coherent series of populations that might be wor- thy of taxonomic recognition. Overall, such structure in the genus was determined to be weak at best. In that paper they noted that both taxa were variable, particularly the latter, and that C. eypsophiloides var. stewartii 1.M. Johnston was statistically somewhat peripheral to groups of populations or individuals produced by c luster analyses or principal components analysis, but was not clearly distinct nor any more distinct in these analyses than other isolated and moderately differentiated populations. By innuendo it is apparent that they believed that C. eypsophiloides var. stewartii was not wor- thy of infraspecific recognition, but unfortunately in the taxonomy pro- vided, this name was omitted from synonymy. Although cumbersome by doing now, I wish to explicitly indicate to future workers that we do not recognize this variety to be distinct from C. gypsophiloides when recognized as a variable taxon composed of numerous semi-isolated populations differ- entiated from one another to varying degrees. The cpg R.M. Stewart 1943, Chihuahua, large canyon near northeast end of Sierra Diablo, 29 Jul 1941, GH! J. Arnold Arbor. 25:173. 1944) was used in the analyses.— Richard Spellenberg, Department of Biology, New Mexico State University, Las Cruces, NM 88003-8001, U.S.A. REFERENCI Maurer, M. and R. SPELLENBERG. 1995, Taxonomy of Cyphomeris eae) ) based on multivariate analyses of geographic variation. Sida 16:679—697 Sipa 17(1): 282. 1996 NOTES 283 RESULTS OF A FIELD SURVEY FOR CYPERUS GRAY OIDES (CYPERACEAE) IN ARKANSAS A field survey for Cyperus grayoides Mohlenbrock was conducted in Arkan- sas in 1995. In 138 locations surveyed, seven populations were found in two counties. Six populations are new discoveries. All populations were found on Agala loamy sand soils. Cyperus grayoides Mohlenbrock (Cyperaceae) is an obscure member of Cyperus section Laxiglumi. It has been reported previously from Illinois, Missouri, Texas, and Louisiana, and more recently from Arkansas. In Texas and Louisiana, C. grayoides is found on communities variously described as sandhill woodland, pine barrens, xeric riparian sandhills, and deep sand savanna. In Missouri the habitat is sand prairie on the Scotco sandridges of the southeastern lowlands. The original Arkansas location is a barrens area in loose eroding sand of the Agala soil association (Bridges & Orzell 1989, Carter & Bryson 1991; Logan 1994). The purpose of this paper is to document additional Arkansas locations of C. grayoides, to assess its habitat preferences in Arkansas, and to suggest where future survey work may reveal still more populations of the species. METHODS A total of 138 sites were surveyed in the field during August and Sep- tember of 1995. Survey sites were in northeastern Arkansas near the Mis- souri populations of C. grayoides, including Crowley’s Ridge and Missis- sippi Embayment; southwestern Arkansas near the Texas and Louisiana populations; and south-central Arkansas with extensive sand barren com- munities and the original Arkansas location of C. grayoides. Additionally, it was suggested (Paul McKenzie, pers. comm.) that dune soils of the West- ern Lowlands of the Mississippi Embayment (between Crowley’s Ridge and the Ozark Escarpment) might contain populations of this sedge (Table 1). Because of habitat requirements of C. grayoides, only sites with sandy soils were surveyed. In Miller and Lafayette counties Briley loamy fine sand was the only sandy soil (Laurent 1984). In northeastern Lafayette County, areas of Bowie fine sandy loam were examined due to reports of sandhill communities in that area. No appropriate soils were found in Ashley, Bra- dley, Calhoun, Columbia, or Union counties of southern Arkansas. In Ouachita and Nevada counties in south-central Arkansas, Agala loamy sand was surveyed because sand barren communities are known on this soil type. No appropriate soils were found in Clark County. In Mississippi County of northeastern Arkansas, the only appropriate soil is Steele loamy sand (Ferguson & Gray 1971). However, examination Sipa 17(1): 283. 1996 284 Stipa 17] TABLE 1. Summary of areas, Counties, site types, and soil types surveyed for Cyperus grayoides. Area/county Types of sites/Communities Soils I. Mississippi E i es Co Mississippi fields Steele loamy sand II. Crowley’s Ridge Greene seeps & associated sites no particular type Clay Co. seeps & associated sites no particular type Craighead Co seeps & associated sites no particular type III. Western Lowlands of Mississippi Embayment Clay ¢ Monioe fe 0. Randolph C 0. cemeteries dune/swale areas dune/swale areas dune/swale areas dune/swale areas : dune/swale areas Woodrutf Co. cemeteries dune/swale areas lV. eye est Arkansas Miller ee os cemeteries, forests, sand Briley loamy fine cemeteries, forests, sand Briley loamy fin Bowie fine ae loam V. South- central Arkansas Nevada C€ cemeteries, forests, sand Agala loamy sand Ouachita c Oo. cemeteries, forests, sand Agala loamy sand of soil maps showed this soil only in small, narrow deposits, all of which were included in agricultural fields. Thus, it was concluded that no native vegetation remained, and Mississippi County was eliminated from further consideration. In the Crowley's Ridge counties of Greene, Clay, and Craighead, no sandy soils are mapped (Robertson 1969; Fielder et al. 1978: Ferguson 1979). However, eroded areas of the ridge have small outcrops of sand and gravel. Since the sandhill communities of south-central Arkansas are often associ- ated with springs and seeps, known seep locations were used to identify potentially sandy areas on Crowley's Ridge. Intensive agriculture in the Western Lowlands of the oe Embayment has replaced native vegetation on the dune-associated soils and where native flora remains the soils are generally too wet for ae ture, €.g., wooded sandpond areas. Thus, cemeteries were the primary sur- vey sites in the Western Lowlands. Sipa 17(1): 284. 1996 NOTES 285 RESULTS Seven populations of C. grayvides, including the original state record re- ported by Logan (1994), were found in Nevada and Ouachita counties of south-central Arkansas. All populations occur on Agala soils, most with enough slope to produce active erosion. In addition, 131 other locations, ranging in size from fractions of a hectare to several sections, were surveyed with negative results. Although occasionally the sandhill indicator Froelichia floridana (Nutt.) Mog. can be seen along roadsides, very little dry sand community vegeta- tion remains in the Western Lowlands. Most cemeteries tended to be too highly alcered for native species to persist, and Bermuda grass (Cynodon dactylon (L.) Pers.) the most common species in them. The few abandoned cemeteries surveyed either were densely wooded or were surrounded by agricultural fields and had become dominated by weeds such as Setaria spp., Polygonum spp., and Johnson grass (Sorghum halepense (L.) Pers.). Crowley's Ridge proved no better for sandhill species. One location with Aristida lanosa Muhl. ex EIlL., a sandhill species, was inspected; however, this site was densely wooded except for occasional highly disturbed areas. In southwestern Arkansas, most sites with Bowie and Briley soils were dominated by dense, brushy secondary successional forests. Such habitats with dense canopies proved unsuitable for C. grayoides. Additionally, few cemeteries were found on these soils and no sandhill species were found in those surveyed. Also an inspection of Miller County Sandhills Natural Area and surrounding lands with Briley soils produced no C. grayordes. Voucher specimens: ARKANSAS. Nevada Co.: Ebenezer Cemetery, T12SR20W, sec- tion 21, parking area, 3 Aug 1995, Logan 95-129 (UCA); T13SR20W, sect. 6, sand bar- rens, 10 Aug 1995, Logan 95-135 (UCA). Ouachita Co.: T12SR18W, sect. 6, sand bar- rens, 1 Aug 1995, Lo 95-122 (UCA); Poison Springs State Forest, T125R19W, sect. 26, sand barrens, 2 Aug 1995, Logan 95-123 (UCA); T12SR18W, sect. 18, sand barrens, 3 Aug 1995, Logan 95- oe (UCA); immediately north of Arkansas Oak Natural Area, T12SR20W, sect. 28, sandy clearing in oak-pine forest, 9 Aug 1995, Logan 95-133 (UCA); T12SR18W, sect. 16, sand barren, 29 Aug 1995, Logan 95-136 (UCA). DISCUSSION AND CONCLUSIONS The typical habitat of C. grayordes is open barren areas of sandhills where enough slope is present to produce some erosion of the deep, loose sand. The only site where it was found in somewhat compacted, uneroding sand was a cemetery with only a few individuals that did not appear to be thriv- ing. C. grayoides appears to require disturbance sufficient to limit competi- tion and to provide open, loose sand for seedling establishment. Such areas are commonly shoulder and side slopes that lack the loamy A horizon re- Sipa 17(1): 285. 1996 286 Sipa 17(1) ported for the Alaga series (Catlett 1973). Instead, the appearance of the surface soil is more like that described for the white, loose C4 horizon. The most severely disturbed site with C. grayoides was on a south and southwest facing slope where the timber had been recently harvested. Here it was thriving, particularly near areas with heavy vehicle traffic. However, extreme disturbance may be harmful to the species, and, in the long run, timber harvest could result in shrub growth which would diminish the open character of the habitat. The species was never found in shaded situations, and at one site forest encroachment appeared to have limited the population size. The lack of a completed soil survey for Nevada County prevented a com- plete survey. When a soil survey for this county is available, it is possible that additional populations of C. grayoides will be discovered, ACKNOWLEDGMENTS Funding for this research was provided by the U.S. Fish and Wildlife Service under a Section 6 (Endangered Species) grant. Special thanks to William M. Shepherd for his comments and suggestions on various drafts of the manuscript. —John M. Logan, Arkansas Natural Heritage Commission, 1500 Tower Build- ing, 323 Center Street, Little Rock, AR 72201, U.S.A REFERENCES Bripces, E.L. and 8.L. Orzett. 1989. Additions and noteworthy vascular plant collections from ae xas and Louisiana, with historical ecological and geographical nores. Phytologia 66:1 Casco: a C.T. Bryson, 1991. A report of Cyperus grayoides and Cyperus oy ; ype oo new to Missouri and notes on other selected Missouri Cyperus. Sida 14: Sl. oe .R. 1973. Soil survey of Ouachita County, Arkansas. USDA-SCS, Washington, DC = D.V. 1979. Soil survey of Craighead County, Arkansas. USDA-SCS, Washing- ton, DC. FerGuson, D.V., and J.L. Gray. 1971. Soil survey of Mississippi County, Arkansas. USDA- SCS, Washington, Fie-ber, R.T., D.V. FerGuson, and J.L. HoGan. 1978. Soil survey of Clay C ounty, Arkan- sas. US 7 as SCS, Washington, DC. — 1984. Soil survey of Lafayette, Litcle River, and Miller counties, Arkansas. US Ae Washington, nes J.M. 1994. A state record fare yperus grayordes (Cyperaceae) in Arkansas. Sida 16:215— 216. Ropertson, N.W. 1969. Soil survey of Greene County, Arkansas. USDA-SCS, Washing - ton, DC Sipa 17(1): 286. 1996 Nores 287 BRACHIARIA ERUCIFORMIS AND UROCHLOA BRIZANTHA (POACEAE: PANICEAE) NEW TO TEXAS Brachiaria eruciformis (J.E. Smith) A. Grisebach and U rochloa brizantha (C. Hochstetter ex A. Richard) R. Webster, previously unreported for Texas have been found in the Edwards Plateau (McCulloch Co.) and southern Rolling Plains (Runnels Co.) respectively. Brachiaria eruciformis is native to Africa, the Mediterranean region, and east to India. Brachiaria eruciformis has been introduced into Australia (Webster 1987; Simon 1993), South America (Morrone & Zuloaga 1992), Southeast Asia (Lazarides 1980), West Indies (Gould 1979), and the United States (Hitchcock 1951; Steyermark 1963). However, Correll and Johnston (1970), Gould (1975), and Hatch et al. (1990) do not report this species as occurring in Texas. No Texas specimens were located at the following her- baria: ARIZ, BRIT, NMC, NMCR, OKL, and TEX-LL. Brachiaria eruciformis is a tufted annual with wiry to spreading culms. Plants are 10O—60 cm in height. Leaf blades are linear to narrowly lanceolate; 2-15 cm long, and 2—6 mm wide. Inflorescence is 1-8 cm long with 6—12 appressed-erect spicate primary unilateral branches, branches 1—2.5 cm long. Primary branches are minutely pubescent, bearing small closely imbricate spikelets on short pedicels. First glume minute; second glume and lower emma are of equal length, 1.8-2.0 mm long. Upper lemma is readily — deciduous, smooth, shiny, and obtuse. Voucher specimens: TEXAS. McCulloch Co.: grazed-out wheat field, 7 mi W of Brady, 15 Jun 1992, H. Hanson s.n. (TAES); in wheat stubble on H. Hanson farm, 5 mi NW of Brady on Hwy 283, 23 Jun 1993, H. Hanson s.n. (TAES). Urochloa brizantha {syn. Brachiaria brizantha (A. Richard) O. Stapf], a native of Africa. It has been widely introduced around the tropics as a for- age grass in Australia (Webster 1987; Simon 1993), Central America (Pohl 1980), South America (Morrone & Zuloaga 1992), Southeast Asia (Lazarides 1980), the Indian subcontinent (Bor 1960), and the United States (Rotar 1968: Hall 1978). However, Correll and Johnston (1970), Gould (1975), and Hatch et al. (1990) do not report this species as occurring in Texas. Urochloa brizantha is an erect perennial up to 2 m in height, culms occa- sionally branching from the middle nodes. The ligule is a short, stiff, cili- ate membrane 1.2—2.2 mm long. Leaf blades are lanceolate, 15-35 cm long and 7-18 mm wide. The inflorescence is a panicle of spicate primary unilateral branches, branches 6—12 cm long. The spikelets are 5.0—5.8 mm long and short pedicellate. The first glume is broadly deltoid, second glume and lower lemma are of equal length. The upper floret is 4.3—4.6 mm long, rigid, finely striate or minutely rugulose. Sipa 17(1): 287. 1996 288 Sipa 17(1) Voucher specimen: TEXAS. Runnels Co.: near Rowena on farm of Mr. Gerngross; plants from seed collected by Ralf Gerngross along Hwy 83 between Ballinger and Paint Rock, 18 Nov 1993, J. Landers s.n. (TAES) We would like to thank Joesph K. Wipff, HI and Hugh D. Wilson for their helpful comments in improving the manuscript—William E. Fox, IL and Stephan L. Hatch, S.M. Tracy Herbarium (TAES), Department of Range- land Ecology and Management, Texas AGM University, College Station, TX 77843-2126 U.S.A.; and Roger Q. Landers, Jv., Extension Specialist (Ret.), Department of Rangeland Ecology and Management, Texas AGM University, San Angelo, TX 76901-9782. REFERENCES Bor, N.L. 1960. The grasses of Burma, Ceylon, India and Pakistan. Pergamon Press. Ox- ford, Great ae an. Crayton, W.D. and S.A RENvoize. 1982. Gramineae (Part 3). In Polhill, R.M., ed. Flora of tropical East Africa. A.A. Balkema. Rotterdam, The Netherlands. Corrett, D.S. and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation. Renner, Texas. Gipps-Russeit, G.E., L. Warson, M. Korkemoer, L. SMooK, N.P. BARKER, H.M. ANDER- son, and M.J. Datiwirz. 1991. Grasses of southern Africa. O.A. Leister, ed. Mem. Bot. Survey South Africa 58. GouLb, EW. 1975. The grasses of Texas. Texas A&M University Press, College Station. GouLb, I. W. 1979. Poaceae. In: Howard, R.A., ed. Flora of the Lesser Antilles, Leeward and Windward Islands—Monocotyledoneae. ..1837; Amn, ey Mus. Naturgesch. 2:119. 1839. Bradburya pubescens (Benth.) O. Ktze, Revis. Gen. Pl. 1:164. 1891. Tyr: ne MicHoacan: Ad Tlalpuxahua a eae Keerle s.n. (HOLOTYPE: BR—ex hb. Martius, mounted ina plastic sleeve and p paperclippec 1). to the herbarium sheet above another specimen — cf. Fig. jon Clitoria schiedeana Schlecht., Linnaea 12(3):284. 1838. Bradburya schiedeana (Schlecht.) Rose, Contr. U.S. Natl. Herb. 8:46. 1899. Centrosema schiedeanum (Schlecht.) Will- tams & Clements. In: Schultze-Kraft and Clements, eds. se ntrosema: pug): Agronomy, and Utilization, p. 7, tab. 1 (as sned.). 1990. Ty PE: MEXICO. Vera Cruz: In dumetis pr. Jalapam Aug, Hacienda de La Laguna, Sep, oo 608 (LecroryPEr, here designated: HAL 36864; tsotecrorypes: B-destroyed, HAL 34078- sh. L!, HAL 34078-sh. 2!, HAL 37536! Clitoria grandiflora Mart. & Gal., Bull. Acad. Brux. 10:189. 1843. Centrosema grandiflorum Walp., Repert. bor. syst. eee 1846. (non Benth. 1837). Centrosema ee Fantz, Sida 8:155. 1979. Type: XICO. Vera Cruz: fl. lilac, bois, Cordill 3000 pied., Jun—Oct 1840, A. pee 3284 (LECTOTYPE: BR!-designated by Fantz, Aida 8:155. 1979; tsoLecrorypes: K!- hb. Hook., TEX!-photo of G). Taxon A now must be renamed. A name is available from synonymy that now has priority. Nomenclature for Taxon A is as follows — molle Mart. ex Benth., Comm. Leg. Gen. 55. 1837; Ann. Weiner Mus. ey 2:119. 1839. Synrypes: BRAZIL. Rio Necro: Cnc. ee amplo, albo, vexillo in medio s*** [?} violaceas, in sylvis ad Barra do Rio Negro, Martius s.n. (M 12505!). BRAZIL. Para: in pascuis et pratis, ad Para, Martius “(MD Centrosema pubescens oe .; Benth. In: MaTuADSy Et Bras. 15:131. 1859; Barbosa-Fevereiro. Roden: 42:184. 1977; non Benth. (18 Centrosema virginianum cE pro parte, Sti ee Flora Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser. 18:528. 1937; Standley & Steyermark. Flora of Guatemala, Fieldiana, ? Bot. 25:181. 1946; non L. FANTz, Centrosema pubescens complex in Central America 50) ADDITIONAL TYPE NOTES Schlechtendal (1838) cited one collection of Schiede in his protologue for Clitoria schiedeana, but did not cite an institution of deposit. Types of both Schlechtendal and Schiede material are deposited at B and HAL, with potential duplicates at several other institutions (Stafleu & Cowen 1985). Berlin material was destroyed in World War II. Four sheets were found at HAL with these duplicates incorporated into the collection on three differ- ent dates. These are treated as syntypes, as discussed by Fantz (1993). The lectotype designated herein bears the name Clitoria schiedeana sp. nov. and agrees best with the protologue. Martius type specimens are at M with types of species of other authors often at BR (Stafleu and Cowan 1981). Curator Paul Bamps (BR, pers. comm.) reported that types for C. molle were at at M, not in their collec- tion. Only one sheet, the Rio Negro syntype (M) was included in a loan request of types. R.J. Clements (pers. comm.) reported four Martius sheets at M, and recognizes the Para collection (M) to be the lectotype. ACKNOWLEDGMENTS Appreciation and thanks are extended to James W. Hardin, Richard Cowan, Rupert Barneby, Rainer Schultz-Kraft and R.J. “Bob” Clements for critical review of the manuscripts, co H. Hertel (M) and Paul Camps (BR) for responding to inquiries on types, and to the curators of the follow- ing institutions for the loan of material studied: ARIZ, BISH, BM, CR, DUKE, ENCB, F, IJ, K, LL, M, MEXU, MICH, MO, MU, NCU, NY, P, SMU, TEX, US, WIS. REFERENCES BARBOSA-FEVEREIRO, V.P. 1977. Centrosema (A.P. De Candolle) Bentham do Brazil - Leguminosae - Faboideae. Rodriguesia 42:159-219. BENTHAM, G. 1837. Centrosena DC. Commentationes de leguminosarum generibus. J.P. Sollinger, Vienna. Pp. 53-56 1859. XX XI. Centrosema. In: Martius and Eichler. Flora Brasiliensis 15(1):124-134. D'Arcy, W.G. 1980. Centrosema. In: John D. Dwyer and collaborators. Flora of Panama, part V. Family 83. Leguminosae subfamily Papilionoideae (conclusion). Ann. Missouri Bot. Gard. 67:571-578 Fantz, P.R. 1979. A new species of Centrosema (Leguminosae) from Nicaragua and a key to the species in Central America. Sida 8:152—156. Fantz, P.R. 1993. Notes on Clitoria (Leguminosae) in Southeast Asia. Sida 3:352—355. 1993. meee hye es (DC.) Benth., xom. cons. In: Flora Novo-Galiciana. 5:335— 340. of Mic n Press, Ann Arbor. Rai cones Cuart. 1985. ONC J-24, scale 1:1,000,000, 6th ed. Defense apping Agency Aerospace Center, St. Louis, Mo. Sipa 17(2) Rost, J.N. 1903. Studies of Mexican and Central American plants - ee 3. A new species of Bradburya with revision of two names. Contr. U.S. Natl. Herb. 8:45—46 SCHLECHTENDAL, D.E.L. 1838. C/itoria. Linnaea 12:284-285. SCHULTZE-Krarr, R. and R.J. Clements, ed. 1990. Centrosema: Biology, agronomy, and uti- lization. Centro Internacional de Agricultura Tropical (CIAT) publication #92, CIAT, Cali, Colombia. Stafleu, FA. and R.S. Cowan. 1981. Taxonomic literature. 3:325. Bohn, Scheltema & Holkema, Utrecht. 1985. Taxonomic literature. 5:146, 190. Bohn, Scheltema & Holkema, Utrecht. STANDLEY, PC. . Centrosema. Flora of Costa Rica. Publ. Field Mus. Nat. Hisc., Bor. Ser. 18:52 28 STANDLI ae and J.A. STEYERMARK. 1946. Centrosema De Candolle in Flora of Guatemala Fieldiana, Bor. 24(5):178-181. U.S. Orrick OF GroGRAPHY. 1956, Mexico: Official standarized name approved by the U.S Board on Geographical Names. — of the Interior, Washington, D.C. WittiAms, R.J. and R.J. Clements. 1990. Taxonomy of Centrosema. In: R. Schultze-Kraft and R.J. Clements, eds. oo naiaaee Biology, agronomy, and utilization. Centro Internacional de Agricultura Tropical. Cali, Colombia. Pp. 1-28. ¢ WOOD ANATOMY OF LIMNANTHACEAE AND TROPAEOLACEAE IN RELATION TO HABIT AND PHYLOGENY SHERWIN CARLQUIST and CHRISTOPHER JOHN DONALD Santa Barbara Botanic Garden 1212 Mission Canyon Road Santa Barbara, CA 93105, U.S.A. ABSTRACT Qualitative and eae itive wood data are provided for Limnanthes douglasii R. Brown and Tropaeolum majus Lyn scriptions of wood of Limnanchaceae or Tropaeolaceae ave been offered hitherto. ease douglasti wood is present in localized zones at the root- stem junction; imperforate tracheary elements are absent; both axial and ray parenchyma are thin-walled. Tropacolum majus has root wood in which all axial tracheary elements are wide or narrow vessels, and no libriform fibers are present; in stems, libriform fibers are — em. The wood patterns of Limnanthes and Tropaeolum are characteristic of wood of an annual and a vine, present, although narrow vessels predominate in later-formed secondary xy respectively. Alchough both are herbaceous species, wood patterns are quite different, a fact explainable by both habit and systematic position. The concept that both of the fami- lies belong to a new expanded Capparales is compatible with wood data, Key worps: Capparales, ecological wood anatomy, glucosinolate families, Limnanthaceae, secondary xylem, Tropaeolaceae, vine anatomy. RESUMEN Se ofrecen datos cualitativos y cuantitativos sobre los lenos de Limnanthes douglas BR. Brown y cna majus L.; hasta ahora no existian descripciones de los lefos de Limnanthaceae y Tropaeolaceae. El lefio de Linnanthes douglasti se presenta local lizado en union entre la rafz y el tallo. Los elementos traqueales sin perforaciones faltan completamente, y tanco el parénquima axial como el radial tienen paredes finas. oki majus tiene el lefio radical con elementos traqueales compuestos por vasos anchos : estrechos; y no hay fibras | libriformes presences. En los tallos existen fibras libriformes, aunque ee en an ema secundario formado tardiamente. Los patrones del leho de L Siena y Lropacolum son caracterfsticos de una hierba anual y una oe ee eens Aunque las dos especies son herbaceas, los modelos del leno son bastante diferentes, licable canto por el habito como por la posici6n sistematica de las dos especies. Los datos del [eto sostienen el concepto de que las dos familias se encuadran en un Capparales nuevo ampliado. INTRODUCTION No data on wood anatomy of either Limnanthaceae or Tropaeolaceae have been presented to date (Metcalfe & C halk 1950; Gregory 1994). The wood of these families is interesting with respect to their divergent kinds of habits; all members of both families are herbs. The two families are grouped here because they are the most herbaceous of the families recog- Sipa 17(2): 333-3: 1996 334 Sipa 17(2) nized among the glucosinolate families, which may be considered Capparales sensu lato, by Dahlgren (1975), Rodman (1991a, 1991b) and Rodman et al. (1993, 1994). Formerly both Limnanthaceae and Tropaeolaceae were included in Geraniales (e.g., Cronquist 1981; Thorne 1992). In some re- cent cladistic systems, Tropaeolaceae are a sister group to Akaniaceae and Bretschneideraceae; Limnanthaceae are not far away in the clade of Capparales (s.1.), and among the groups nearest to Limnanthaceae are Batidaceae, Caricaceae, and Moringaceae (Rodman et al. 1993, 1994). Tropaeolaceae and Limnanthaceae are placed close to Capparaceae in the cladogram of Conti et al. (1996), who include fewer of the glucosinolate families in their survey, and who therefore offer less precision in the place- ment of these two families. Rodman et al. (1993) concede that marked evolutionary shift in habit must be hypothesized if Tropaeolaceae are close to the tree families Akaniaceae and Bretschneideraceae. However, they do not consider this improbable and invite consideration of how these changes may have oc- curred. Similar considerations apply to Limnanthaceae. The species of Tropaeolum studied, the well known T. majus, is an annual herb with vining capabilities. The base of the stem contains a cylinder of secondary xylem I—2 mm in thickness; the upper portion of the root contains a xylem core 1—3 mm in thickness; these have been used in the present study. Limnanthes douglasii is less woody than Tropaeolum;, Limnanthes is an annual with a short basal stem to which numerous thin roots are attached; upwardly, the basal stem branches into several prostrate stems. The patches of secondary xylem in the stem of Limnanthes are evidently related to these prostrate stems. These distinctive habits, and the fact that secondary xylem in Limnanthaceae and Tropaeolaceae is minimal in comparison with that of other capparalean families provide reasons for study of wood anatomy of these two families. If marked differences in habit have occurred phyletically in the divergence of these two families from their nearest neighbors, we will have new examples of how nature of wood anatomy reflects shift in habic. To be sure, only one species of each family has been used in the present study. The reason for this selection is that for each family, the species studied is the woodiest that could be obtained readily. Further studies are needed, but they may demonstrate more herbaceous modes of structure than do the species in the present paper, judging from the habits described for Limnanthaceae by Ornduff (1971) and Tropaeolaceae by Sparre et al. (1991). MATERIALS AND METHODS Stems and roots bearing secondary xylem from plants of Tropaeolum majus and Limnanthes douglasti that were judged to be mature but not yet in de- cline were collected from plants in cultivation. In the case of the Tropaeolum, CARLQUIST AND DoNaLb, Wood anatomy of Limnanthaceae and Tropaeolaceae oe, plants were collected from a Santa Barbara garden where the species has naturalized. Plants of Limnanthes douglasii were collected from Santa Bar- bara Botanic Garden, where they have become naturalized, with the aid of Mark Stubler. The portions bearing secondary xylem were preserved in 50% aqueous ethyl alcohol. Wood of Tropaeolum was softened by immer- sion in 3% ethylene diamine for two hours. Material was infiltrated and embedded in paraffin according to the usual techniques, sectioned on a rotary microtome; sections mounted on slides were stained with a safranin- fast green combination corresponding to Northen’s modification of Foster's tannic acid-ferric chloride method (Johansen 1940). In addition, some sec- tions of unsoftened stems of JT: majus were sectioned on a sliding micro- tome, dried between glass slides, sputter coated, and examined with a scan- ning electron microscope Macerations were prepared with Jeffrey's fluid and stained with safrani Terminology follows that of the IAWA Committee on Nomenclature (1964). The term “pseudoscalariform” refers to pitting in which pits are horizontally elongate, as in scalariform pitting, but are derived from alter- nate pits that are markedly widened, and thus the ends of the pits do not correspond to angles of the vessels. Vessels were measured in term of lumen diameter; if oval rather than near-circular in outline, the mean diameter was estimated. The term “intervascular” axial parenchyma refers to a situ- ation in which occasional axial parenchyma cells are scattered among large groupings of vessels, whereas paratracheal parenchyma applies to paren- chyma surrounding a vessel or vessel group. ANATOMICAL RESULTS Limnanthes douglasii stem (Fig. 1-5). Secondary xylem very limited in extent, and also localized in several segments within the vascular cylinder (Fig. 1, top), intervening parts of the cylinder composed of primary xylem only (e.g, Fig. 1, bottom). Vessels square or polygonal in outline, in large groupings with few intervening axial parenchyma cells (Fig. 1); nieaD num- ber of vessels per group, 18.1; mean number of vessels per mm 7° (no areas adjacent to secondary xylem included): 690. Mean vessel diameter, 28.2 im. Mean vessel element length, 71.6 jim. Perforation plates all simple, ie and appreciably narrower than the vessel diameter (Fig. 4). Vessels irregular in outline, either as seen in tangential (Fig. 3) or radial (Fig. 4) sections. Lateral wall pitting of vessels consisting of pits laterally elongate, alternate to pseudoscalariform (Fig. 5). Lateral wall pits (pit cavity) about 5 jm as measured vertically (Fig. 5). Imperforate tracheary elements ab- sent. Axial parenchyma sparse, intervascular (Fig. 2), subdivided into strands or two or not subdivided, with thin primary walls only. Rays uniseriate or biseriate (Fig. 3). Mean height of biseriate rays, 238 jm; mean height of Stipa 17(2) 336 (/UIHIIL NS me UMMA ee sr a UE | Ae om ’ “ a A, or, 2. ah ie +4 /4 y B AA sy a 2 Fol Figs. l= Wood sections of Dem douglasii stems. Fig. 1. Transection; portion of ee eh secondary xylem, above, portion with only primary xylem below. ig. 2. Enlarged portion of secondary xylem to show angular nature of vessels: imperforate tri fie iry elements are absent. Fig. 3. Tangential section; rays are uniseriate or biseriate, com- posed of upright cells. Fig. 4. Radial | section; shortness of vessel elements evident. Fig. 5. Vessel element from tangential section to show ae ee alariform lateral wall pitting. Fig. 1, scale above Fig. | (divisions = 1( )yim); Figs. 2 scale above Fig. 2 (divisions = 10 jum); Fig. 5, scale above Fig. 5 (divisions = 10 a: CARLQUIST AND DoNaLb, Wood anatomy of Limnanthaceae and Tropaeolaceae Byo uniseriate rays, 91 pm. Ray cells with thin primary walls. Wood nonstoried (Fig. 3), no starch or crystals observed. Tropaeolum majus stem (Fig. 6, 8-15). Axial portions of wood consisting largely of vessels dimorphic in diameter (Fig. 6). Wide vessels solitary or in pairs (mean number of vessels per group in nonray areas, 1.8), mean diam- eter, 81 am. Narrow vessels comprising the bulk of the remainder of the secondary xylem and therefore in indefinitely large groups; mean diameter of the narrower vessels, 29 pm. Mean length vessel elements (wide and narrow vessel elements combined), 116 jm. Mean wall thickness of vessels varying from 1.8 pm for narrower vessels to 2.5 um for wider vessels. Per- foration plates mostly simple, but occasional scalariform or scalariformlike plates present (Fig. 12-15): some plates with numerous forked or intercon- nected bars (Fig. 12), some with few bars (Fig. 13), some, on narrow ves- sels, with wide bars (Fig. 14), some with a condition intermediate between scalariform and multiperforate (Fig. 15). Lateral wall pitting of vessels al- ternate, pits circular to oval (Fig. 9, left) or elliptical, about 5 pm measured vertically. Libriform fibers present, especially in earlier formed portions of secondary xylem, supplanted in later portions of axial secondary xylem by narrow vessels. Libriform fibers with simple pits, and often containing starch grains (Fig. 9, right). Mean length of libriform fibers, 462 xm; mean diam- eter, 24 pm; mean wall thickness, 2.2 pm. A few vasicentric tracheids and a few cells intermediate in pitting between libriform fibers and vasicentric tracheids present. Axial parenchyma with thin secondary walls, in strands of two cells, adjacent to the larger vessels. Rays multiseriate only, more than 1,000 pm in mean height (Fig. 8, right). Mean width of multiseriate rays, 12 cells. Ray cells with thin primary walls, bearing prominent pit fields on radial walls, separated from each other by horizontal bands of primary wall material (Fig. 10). Ray cells predominantly upright; square and procumbent cells in central portions of rays (Fig. 10). Wood nonstoried (Fig. 8). Starch abundant in ray cells (Fig. 10), starch grains spherical in shape (Fig. 11). Tropaeolum majus, root (Fig. 7). Axial portions of wood consisting mostly of vessels dimorphic in diameter. Larger vessels mostly solitary, mean num- ber of vessels per group, 1.4. Narrower vessels in groups of indefinite ex- tent. Mean number of larger vessels (ray areas excluded), 44.8. Mean diam- eter of larger vessels, 96 pm; mean diameter of narrower vessels, 27 pm. Mean wall thickness of vessels varying from 1.8 to 3.5 pm. Mean length of vessel elements (wide and narrow vessel elements together), 99 pm. Perfo- ration plates and lateral wall pitting as in stems. Libriform fibers very few; a few vasicentric tracheids present. Axial parenchyma in strands of two cells, with secondary walls, adjacent to the larger vessels. Rays multiseriate and uniseriate (Fig. 7). Mean height of multiseriate rays, 1460 jm; mean 338 SIDA 17(2) e ¢ = ——@ S eon, 6! Sa ed Ey % ~ a ) pM IK eS sia # sf, PX : m . san ie: ee 7 Fics. 6-11. ae le sections of secondary xylem from stems (Figs. 6, 8-11) and root (Fig. 7). Fi ransection; marked dimorphism in vessel diameter evident; wide ray ac right. Fig. = aie section; rays narrower, shorter than those of stems. Fig. 8. Tangential section; rays are multiseriate, wide, tall. Fig. 9. SEM photograph of vessel (left) and libriform fiber (right). Fig. 10. Portion of ray from tadial | section; primary pit fields common on radial walls. Fig. 11. SEM yh of ray cells from ia eae section to show starch grains. Figs. 6-8, scale above Fig. 1; Fig. 10, scale above Fig. 2; Figs. 9, 11, scales at lower left (bar = 5 pm CARLQUIST AND DoNaLb, Wood anatomy of Limnanthaceae and Tropaeolaceae 339 | Fics. 12-15. Perforation plates from vessels of Tropaeolum majus stems. Fig. 12. Light mi- crograph of perforation plate from transection, bars numerous and narrow. Figs. 13-15 SEM photographs of perforation plates from eae section. Fig. 13. Bars few, wide. Fig. 14. Plate from narrow vessel; bars wide. Fig. tia ‘eraedicee between scalariform and multiperforate. Fig. 12, aie above Fig. 7 - igs. 13-15, bars at lower left = 5 jm. 2A 340 Sipa 17(2 — height of uniseriate rays, 312 pm. Ray cells mostly upright, a few square and procumbent cells present in central ieee of multiseriate rays. Mean width of multiseriate rays, 5.1 cells. Ray cells thin-walled, nonlignified. Wood nonstoried, but vague storying visible in tangential sections of sec- ondary phloem. Starch grains abundant in ray cells. DISCUSSION AND CONCLUSIONS Limnanthes and Tropaeolum differ in a number of salient ways with re- spect to wood anatomy. Limnanthes douglasii has narrow vessels angular in transection, nondimorphic in diameter, with simple perforation plates ex- clusively and with mostly pseudoscalariform lateral wall pitting; imperfo- rate tracheary elements are entirely absent; axial parenchyma is sparse and with thin nonlignified walls; rays are uniseriate or biseriate, composed of upright cells. Tropaeolum majus has markedly dimorphic vessels; the larger are circular in transectional outline, the smaller circular to angular in out- line; lateral wall pitting is mostly circular to elliptical; perforation plates are simple, but with an appreciable number of perforation plates that rep- resent alterations of a scalariform pattern; libriform fibers with simple pits are present (somewhat less abundant than the narrow vessels), but a few vasicentric tracheids and cells intermediate between libriform fibers and vasicentric tracheids are present; axial parenchyma has thin lignified walls, is in strands of two cells, and is distributed adjacent to the larger vessels. In T. majus, rays are wide and exclusively multiseriate in stems in roots, rays are mostly multiseriate and less wide; ray cells are thin walled, filled with starch, and are mostly upright, but with procumbent and square cells in central ray portions. The lack of imperforate tracheary elements in Limnanthes may be corre- laced with lack of upright stems, because libriform fibers would be ex- pected where addition of mechanical strength is of selective value. The occurrence of secondary xylem in small restricted patches is related to the short duration of this annual, and the short stem. Secondary xylem appar- ently connects the branches with the basal stem, and the remainder of the basal stem contains primary xylem only. Thus, habit seems to correspond to those distinctive features. The shortness of vessel elements in Limnanthes is likely related to xeromorphy; short vessel elements are found in plants of habitats that dry as a season progresses (Carlquist 1966). Tropaeolum wood is distinctive in the dimorphism of its vessel diameter, in the sparsity of libriform fibers (supplanted to a large extent by narrow vessels), and in the very wide, tall rays. These are features common in woods of vines and lianas (Carlquist 1985). The relative shortness of vessel ele- ments in 7, mazus is likely related to the drying of soil as the plant reaches its mature size. CARLQUIST AND DonaLp, Wood anatomy of Limnanthaceae and Tropaeolaceae 341 Other differences between the wood of Limnanthes and that of Tropaeolum seem related to phylogenetic relationships. For example, the wood of Tropaeolaceae, placed close to Akaniaceae and Bretschneideraceae (Rod- man et al. 1993, 1994), shares with that pair of families a number of wood features (wood of those two families monographed by Carlquist, submit- ted): perforation plates simple but with an appreciable number of modified scalariform perforation plates; libriform fibers present; axial parenchyma sparse, distributed adjacent to the larger vessels (vasicentric in Akaniaceae and Bretschneideraceae); rays wide and tall, uniseriate rays few or absent (wide rays also a characteristic of vines and lianas, but absence of uniseriate rays is not characteristic of lianas). On the other hand, some of the features of Limnanthes wood can be found in genera of Capparales from which it is not far removed according to the schemes of Rodman et al. (1993, 1994). For example, in Capparaceae, rays are uniseriate or biseriate, rarely multiseriate (Metcalfe & Chalk 1950). The marked difference in habit between the annual Limnanthes and genera of Capparaceae, which are trees or shrubs, means that modifications related to habit are likely to be of overriding importance, and similarities related to phyletic relationship are likely to be few. Further studies on wood anatomy of the glucosinolate families, which may tentatively be considered a redefined Capparales, are planned so as to show the relative degrees of influence that habit and phylogenetic relationship have on similarities and differences among the component families of that order. REFERENCES Car ourst, S. 1966. Wood anatomy of Compositae: a summary, with comments on factors influencing wood evolution. Aliso 6(2):25—44. 1985. Observations on functional wood histology of vines and lianas: vesel dimorphism, tracheids, vasicentric tracheids, narrow vessels, and parenchyma. Aliso 11:139- as — ene snbnned: Wood anatomy of Akaniaceae and Bretschneideraceae: a case of cerca te and its systematic implications. Submitted to Syst. Bot. Cont, E., A. L ind K.J. Syrsma. 1996. Circumscription of oe and cheir os sn to a octe evidence from rbcL sequence data. Amer. J. Bot. 83:221— CRONQU ust, A. 198L. An ee rated system of classification of en plants. ae University Press, New Suse REN, ae 1975. A system of classification of the angiosperms to be used to demon- strate the distribution of characters. Bot. Not. 129:119-14 GREGORY, 1994. Bibliography of systematic wood anatomy of dicotyledons. IAWA J., Suppl. L:1- Tawa ComMMITTEE ON NoMENCLATURE. 1964. Multilingual glossary of terms used in wood anatomy. Meclapanstale Konkordia, Winterthur, Switzerland. Jouansen, D.A. 1940. Plant microtechnique. McGraw Hill, New York. Metcatre, C.R. and L. Cuatk. 1950. Anatomy of the dicotyledons. Clarendon Press, Oxford. jon ~~ 342 Sipa 17(2 ORNDULE, 1971. Systematic studies of Limanathaceae. HarON® 21:103-111. MAN, lucos producing plants, Part Dis : Phenetics. Syst. Bot. 16:598 I1b. A taxonomic canis of glucosinolate-producing plants, Pare 2: Cladistics. een , R.A. Price, K. Karor, E. Conti, K.J.K, and J.D. Parmer. 1993. Nucle- plants. Ann. Mis- rbcL - ne oe monophyly of mustard oi O oe 991a. A taxonomic enalysis of § otide sequences of the sour! Bot. Gard. 80:585— , KG. Karon, z re Prick, E. Conti, and K.J. Syrsma. 1994. Nucleotide poe KG, sequences : rbcL confirm the cap >paralean affinity of the Australian endemic ayrostemonaceae, Austral. Syst. Bot. 7:57—-69. ] Classification and geography of the flowering plants. Bot. Rev THORNE, R.F. 58:225-348. NEW GEOGRAPHICAL AND MORPHOLOGICAL DATA FOR SIDEROXYLON THORNE! (SAPOTACEAE) LORAN C. ANDERSON Department of Biological Science Florida State University Tallahassee, FL 32306-2043, U.S.A. ABSTRACT Recent discoveries of new localities for this rare species justify documentation of its presently known range. The additional specimens show that leaves and fruits can be much electron larger than previously recorded. Foliar micromorphology as viewed by scanning e microscopy demonstrates the species is distinctive and probably noc derived from S. lanuginosum and S. reclinatum as oa rly thought. RESUMEN El reciente descubrimiento de nuevas localidades de esta rara especie justifica la documenta- cién de su rango conocido actualmente. Los especimenes adicionales muestran que las hojas y los frucos pueden ser mucho mas grandes que los citados previamente. La mitromortolegia foliar al microscopio electrénico de barrido demuest ra que esta especie es peculiar y ¢ se ercnaiate no es derivada de S. /anuginosum y S. reclinatum como se pensé previamente. 2 o The buckthorns of the southeastern United States have long been known as species of Bumelia. Thorne’s buckthorn or swamp buckthorn, Bymelia thornei Cronq., was described in 1949. It was identified as a species deserv- ing Federal listing as an endangered species (Smithsonian Institution 1975) and has remained under consideration for listing ever since. Recently, Pennington (1990) made the combination Sideroxy/on thorner, he gave con- vincing arguments (1991) that Bumelia should be considered congeneric with Szderoxylon. Thorne’s buckthorn was originally known from only a few counties in southwestern Georgia and was subsequently reported (Anderson 1988) nearby in Jackson County, Florida. Recent collections extend its known range into southeastern Georgia, extreme southeastern Alabama, and along the Apalachicola and Escambia rivers in Florida (Fig. 1). The following document the known range of S. thorne: (sites identified only generally be- cause of the species’ rareness): ALABAMA: Houston Co.: Big Creek, 15 May 1994, Ss McDonald (FSU, TENN). HORIDA Escambia Co.: Escambia River, 16 Sep 1984, J.R. Burkhalter 9597 (FSU). Franklin eMedlachicola River, 17 Nov 1995, L.C. ae 16114 (FSU), Gulf Co.: ern en 17 Nov 1995, L.C. Anderson 16107 (FSU), L.C. Anderson 16108 (FSU), Sipa 17(2): 343-348. 1996 344 Sipa 17(2) a ae Known range for S. thornei Fic. 1. Known range of Srderoxylon thornei. Forbes Island, 17 Nov 1995, L.C. Anderson 16111 (FSU), Apalachicola River, 2 Nov 1995, H. Licht & M. es sn. (FSU : pens : N of Sneads, 15 Nov 1995, L.C. Anderson 16082 (FSU); 11 Oct 1994, . Burks i (FSU); 16 Aug 1986, = : Gholson 11720 (FLAS, FSU); : Jun 1988, A. : Chilen 11981 (FLAS); 1 May 1982 oo 79745 (FSU); 30 Jun 1987, R.K. Gedfrey 82429 (ELAS, FSU, MO); 30 Jun ee R.K. Godfrey 52450 (FSU-several sheets); 27 Aug 1987, R.K. Godfrey 82491 (FLAS, FSU-several sheets); 20 Oct 1987, R.K. Godfrey 82601 (GA, FLAS, FSU); 13 Jun 1988, R.K. Godfrey 82691 (GA, FLAS, FSU); 18 Oct 1990, - K. Godfrey sae (FSU, NY); N of Sne: a 19 Jun 1988, R.B. McCartney s.n. (GA); 13 Sep 1988, R.B. Mc ‘Cartney sn. (FLAS). GEORGIA: Baker Co.: Flint River, 20 Oct 1 LG. Ander son 15999 (FSU); limestone cae 3 Jun 1940, D. Eyles 7050 (GA); Flint River, 25 _ 993, L.K. Kirkman 3105 (Ichauway Herb.). Calhoun Co.: Ichawaynochaway Creek, 23 May 1990, ].R. Allison 4458 (GA). Decatur Co.: Spring Creek, 24 Oct 1990, J.R. Allison 5058 (GA). Early Co.: E of Nantz Springs, 22 May 1990, J.R. Allison 4446 (GA, MO); 3 1 Jul 1986, A.K. Gholson 11701 (FSU); 14 Aug 1986, A.K. Gholson 11713 (GA); 14 Aug 1986, R.K. Godfrey 82003 (GA, FLAS, FSU- several sheets, MO); 1 Oct 1 ae R.K. Godfrey 82155 (FLAS, FSU-several sheets); 20 Oct 1992, L.K. Kirkman 2704 (GA, Ichauway Herb.); 13 Sep 1988, R.B. et y.n. (FLAS): 22 Oct 1947, R.F. Thorne 7345 (GA-holotype, NY-isotype); Spring Creek, 24 Oct 1990, J.R. Allison 5052 (GA). Liberty Co.: Cedar Bay, 19 Aug 1993, M.O. Moore 1856 (GA). Miller Co.: Colquitt Swamp, 30 Aug 1990, J.R. Allison 4715 (GA). Wy Reports of the species from Pierce, Tattnall, and Toombs counties in Georgia were based on misidentifications; the specimens were later identi- hed as either S. reclinatum Michx. or the hybrid, 8. danuginosum Michx. x S. reclinatum. The original description (Cronquist 1949) said the species was a thorny ANDERSON, Sideroxylon thornei 345 shrub about 1.5 m tall with mostly elliptic or elliptic oblanceolate leaves, 1.5—4 cm long and 6-20 mm wide. Godfrey’s more detailed description (1988) noted the shrubs could reach 6 m in height and that leaves on a given shrub were variable in both size and shape. The variation in leaf size and shape is even greater than that recorded by Godfrey. Most leaves are oblanceolate to elliptic-oblong, but some are definitely ovate (up to 5.4 cm wide). Outlines of largest leaves from selected specimens demonstrate this variation in Figs. 2-10. The largest leaf on the holotype, Thorne 7345, was only 3.9 cm long (Fig. 2), whereas the largest leaf seen for the species (Fig. 10) was 13.9 cm long (Kirkman 3105). Part of the great size range in leaves results because some specimens were taken from the fast-growing primary canes and others from older stems that have numerous spur shoots with smaller leaves. Godfrey’s population samples from the type locality show that those plants represented about 33% of the total variation found in the entire species’ range in leaf size. Cronquist (1949) noted that Thorne’s buckthorn seemed to combine characteristics of S. /anuginosum and S. reclinatum and suggested it was prob- ably of hybrid origin; that view was repeated by Pennington (1990). The key features for S. thornei (Godfrey 1988) include relatively glabrous stems as in S. reclinatum and hairy leaves more like those of S. /anuginosum but faintly reticulate-veined with the veins somewhat impressed (not bony car- tilaginous as in S. reclinatum). The occurrence of larger leaves in many popu- — lations of S. thornez make it less similar to S. reclinatum. Scanning electron microscopy of leaf surfaces was initiated to observe the dolabrate or malpighian hairs, but the form found associated with the stomata proved more interesting. All S7deroxy/on species in the southeast- ern United States (seven to nine, depending on interpretation) have large stomatal rims that are raised funnel- or dome-like structures with long, narrow apertures that more or less hide the guard cells—somewhat like those illustrated for Gordonia (Anderson 1983). The stomatal density amongst Sideroxylon species appears to be inversely proportional to size of the stomatal apparatus; i.e., smaller stomates are more numerous per unit area than are larger ones (see Figs. 13 and 15). Leaf surface micromorphology (Figs. 11-13) distinguishes S. thorne7 from all other taxa in the genus; samples from four different localities were scanned. The stomatal rims nearly cover the stomata; the rims are 14.9— 17.2 pm long, and their apertures range from 0.9—1.3 pm wide. Numerous cuticular striae are aligned parallel to one another and generally at right angles to the long axes of the stomatal rims. Epidermal patterns for S. lanuginosum (Fig. 14), S. reclinatum (Fig. 16), and their putative hybrid (Fig. 15) have less pronounced stomatal rims with more open apertures, and the cuticular striae tend to encircle the rims. Stomatal rims in S. /anuginosum 346 Sipa 17(2) Fics. 2-10. Largest leaf per 8. thorner specimen to show variation in size and aes all che same scale. Fig. 2. Thorne 7345, holotype. Fig. 3. Thorne 7345, isotype. Fig. 4. Al/ison sueyewued 9 10 §. =< 715. Fig. 5. Allison 5058. Fig. 6. Anderson 11607. Fig. 7. Anderson 1 1614. Fig. 8. Barbbalver 9596, Fig. 9. Moore 18506. Fig. 10. Kirkman 3105, range from 14.6-18.6 pm long, whereas they are shorter in S. reclinatum (12.7-13.1 jm). The surface pattern in the putative hybrid (Fig. 15) is intermediate between those of the presumed parents and supports the no- tion of its hybridity. Clearly, on the basis of morphology of the stomatal complex, S. thornes is not derived from S. /anuginosum and S. reclinatum as some authors have sug- gested. The stomatal rims of S. thorne7 are most similar to those of S. /ycioides L., but leaf surfaces of the latter have none of the cuticular striae that are so abundant in S. thornez. Godfrey (1988) recorded fruit of Thorne’s buckthorn as dull black, 8—10 mm long, and 8-10 mm wide, whereas fruit of Light & Darst (possibly more mature) was shiny black, slightly obovate, 12 mm long, and 11 mm wide. Sideroxylon thornes is listed as endangered in Florida (Coile, 1993) and in Georgia (Patrick et al. 1995); it is a species of special concern in Alabama (Oberholster 1996). It appears to be a distinctive species in many ways. Known occurrences are still relatively few, and considerable habitat loss has been noted (Patrick et al. 1995). It deserves further field investigation of its distribution and threats and should be considered for Federal listing. ANDERSON, Sideroxylon thornei 347 Fics. 11-16. Scanning electron micrographs of abaxial leaf ae: to show stomatal rims Ay and cuticular patterns; all the same scale (scale bar in Fig. 16 = 10 pm). Figs. 11-13 er note nn stomata and similarity of cuticular relief. =a Ll. Godfrey 8 5063 (topotype). Fig. 12. Anderson 16107. Fig. 13. Moore 1856. Fig. 14.8. lanuginosum, Anderson 9057, note visible guard cells and numerous cuticular striae encircling the more open stomatal rims. Fig. 15. Hybrid of S. /anuginosum x S. reclinatum, Anderson 4858, with guard cells partially exposed and limited cuticular striae. The smaller stomatal complexes that are concomitantly more numerous per unit area may be related to hybridity. Fig. 16. reclinatum, Godfrey 63520, with partially open stomatal rims, cuticular striae, and minute flecks of epicuticular wax. Sipa 17(2) — REFERENCES ANDERSON, L.C. 1983. Trichomes and stomata of Gordonia lasianthus (Yheaceae), Sida 10:103-113 ANDERSON, L.C. 1988. Noteworthy plants from north Florida. III. Sida 13:93—100 Corte, N.C. 1993. Florida’s endangered and threatened plants. Florida Dept. Agric. and Consumer Serv., Division of Plant Industry—Bot. Sect. Contrib. 29, Gainesville. Cronguist, A. 1949. Noteworthy plants of Georgia. Castanea 14:101—108 Goprrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. Univ. Georgia Press, Athens. OBERHOLSTER, C. 1996. Species inventory list. Alabama Natural Heritage Prog., Montgomery. Parrick, T.S., J.R. ALLISON, and G.A. Krakow. 1995. Protected ‘aad of Georgia. Wild- life Hesonpees Div., Ga. Dept. Natural Resources, Social Cir PENNINGTON, T.D. 1990, Sapotaceae. Flora Neotropica Monee a 1-770. PENNINGTON, T.D. 1991. The . nera of aaa Royal Botanic Gardens, Kew. rt on endangered and threatened species of the United = © SMITHSONIAN INSTITUTION, 1975. I States, presented to — of the United States of America by the secretary, Smithsonian Institution. House document No. 94-51, 94th Congress. U.S. Govern- ment Pringting Office. MORPHOLOGICAL VARIATION AND SYNOPSIS OF THE MUHLENBERGIA REPENS COMPLEX (POACEAE) CLIFFORD W. MORDEN Department of Botany Hawaiian Evolutionary Biology Program University of Hawaii Honolulu, HI 96822, U.S.A. cmorden@hawati.edu STEPHAN L. HATCH S.M. Tracy Herbarium Department of Rangeland Ecology and Management Texas AGM University College Station, TX 77843-21206, U.S.A. s-hatth@tamu.edu ABSTRACT The Muhlenbergia repens complex consists of rhizomatous perennial species wich short contracted inflorescences and awnless spikelets. A study was undertaken to determine the amount of morphological variation that is present v an and among the species. Exami- nation of morphological variation has shown that M. ae does not have a close affinity with other species of the are Mubhlenbergia richardsonis and M. squarrosa were not found to be morphologically distinct and are considered a single highly variable species most closely related to M. repens aa M. utilis. Reinnemed variation between M. villiflora and M. villosa intergrades to a large extent, although they remain distince throughout most of their distribution, and they are thus recognized as a ak species with two variet- ies, var. villiflora and var. villosa. A key to the six species and two varieties is provided. RESUMEN El complejo Muhlenbergia repens esta formado por especies rizomatosas perennes con inflorescencias cortas contraidas y espiguillas muticas. Se realiz6 un estudio para establecer la variaci6n morfolégica intraespecifica e€ interespecifica. El examen de la variacion fe aenire? ha mostrado que M. plambea no tiene afinidades grandes con las otras especies del complejo. Se encontré que Muahlenbergia richardsonis y M. squarrosa no son distintas erie y son consideradas como una sola especie de variabilidad alta, mas afin a M. repens y M. utilis. La variacion morfoldgica entre M. villiflora y M. villosa se intergrada en una gran extensid6n saad permanecen diferentes en la mayor parte de su distribucisn, y por ello se reconocen como una Unica especie con dos variedades, var. villiflora y var. villosa. Se ofrece una clave de las seis especies y las dos variedades. Sipa 17(2): 349-365. 1996 350 Sipa 17(2) INTRODUCTION The genus Muhlenbergia Schreb., with about 160 species, has a center of distribution in the southwestern United States and northern Mexico. In general, these species are found in hot, arid habitats alchough some species occur in diverse situations such as moist shaded woodlands (M. schrebers Gmel. and its allies) and alpine meadows {M. richardsonis (Trin.) Rydb. and M. filiformis (Thurb.) Rydb.}. Members of the Muhlenbergia repens comple (Morden & Hatch 1987) are distributed throughout North America (ex- cluding the southeastern United States) and in the Andean Highlands of South America. This complex is characterized by a rhizomatous perennial habit, short culms seldom exceeding 40 cm, short involute leaf blades, and a short contracted inflorescence with awnless or mucronate spikelets. As presently understood, this complex consists of eight species which may be placed into two groups based on floret vestiture. Six species are typically considered to have glabrous spikelets, several of which are minutely pubes- cent or scabrous on the lemma when observed closely (above 20x magnification). These include M. repens (Presl) Hitche., M, wtilis (Torr.) Hitche., M. richardsonis (Trin.) Rydb., M. squarrosa (Trin.) Rydb., M. fastigiata (Presl) Henrard, and M. ies (Trin.) Hitche. Two species, M. villiflora Hitche. and M. villosa Swallen, have florets that are densely villous on both the lemma and palea. Muhlenbergia plumbea, which typically grows larger chan is usual for . complex and often has an open inflorescence, is included in this complex upon the suggestion of Charlotte Reeder (pers. comm.). Most of the species occur in the southwestern United States or adjacent Mexico. The species of the M. repens complex have been traditionally separated into two groups on the basis of glabrous or villous florets (Hitchcock 1950; Gould 1975). Analysis of these species using scanning electron microscopy (SEM) found that only three of the species (M. fastigiata, M. plumbea, and M. utilis are completely devoid of vestiture on the floret, whereas all others have macrohairs present on the lemma and/or palea (Morden 1985). Both M. villiflora and M. villosa have copious macrohairs evident without the aid of magnification. However, M. repens, M. richardsonis, and M. squarrosa all have macrohairs, albeit smaller and in much lower frequency than M. villiflora and M. villosa, distributed on the lemma and palea in a pattern similar to that of the densely villous species (near the base of the lemma and between the two nerves of the palea). Of these three species, macrohairs of M. repens are typically longer and more densely distributed over the cal- lus of the floret and the lemma and palea. In some populations the hairs are visible at 20 magnification with a dissecting microscope. Macrohairs of M. richardsonts and M. squarrosa are shorter and less frequent, and were seen only with SEM. Morben AND Hatcu, The Muhlenbergia repens (Poaceae) complex 35] Little is known of the variation among the taxa of this complex. Morden and Hatch (1987) studied the variation present within leaf anatomical traits and found that all species of this complex were discernible with the excep- tion of M. richardsonis and M. squarrosa. There is an apparent overlap in morphological characters and sympatry in distribution (occasionally grow- ing side by side). In one instance, three different species of this complex were found growing within a five square meter area. Morphological charac- ters which have often been used to separate species within this complex often show considerable variation. For example, of the species with gla- brous florets, only M. richardsonis and M. squarrosa were reported to have nodulose roughening on the culms. However, upon close inspection this is found to a limited degree in all species of the complex. Further, it is not uncommon within some populations for the lower branches of the inflorescence to become relaxed, and in some cases reflexed, similar to what is found in M. plumbea, although most taxa typically have contracted panicles. There are two problematic species groups within this complex. In one, M. richardsonis and M. squarrosa have been found throughout North America north of Mexico (including Alaska and the provinces of Canada) except in the southeastern United States. They have previously been reported as sepa- rate species based on culm morphology or as a single species (under M. richardsonis; Cronquist et al. 1977); M. squarrosa has stout, decum bent, and spreading culms and occupies drier sites with little competition from other species, whereas M. richardsonis has more slender, erect culms and 1s usually growing in moister soils in a meadow-like association. The South Ameri- can species, M. fastigiata, appears similar to M. squarrosa and M. richardsonts in all characters except that the vegetative morphology is depauperate. This species occurs in the Andean Highlands of Argentina, Bolivia, and Peru, and is not found below 3000 meters elevation. Given the broad distribu- tion of M. richardsonis and M. squarrosa, and that it often occurs at high elevations in the Rocky Mountains, one previously unexplored possibility is that M. fastigiata is a smaller South American form of this species. Al- though M. richardsonis and M. squarrosa are apparently morphologically distinct from others of the complex, they are most similar to and appar- ently intergrading in some regions of their range with AM. repens (in the southwestern United States) and M. wti/is (in Texas and California). As a result of the similar morphology herbarium specimens are frequently misidentified cond problematic species group includes M. v7/liflora and M. villosa. ie villiflora forms extensive stands across gypsiferous soils in north- ern Mexico and is a dominant component of that ecosystem. In contrast, M. villosa is found in very localized populations on alkaline and calcareous soils of western Texas and adjacent New Mexico. The nature of the rela- 352 Stpa 17(2) tionship between these two species has not been previously explored, and the paucity of populations of M. villosa available for comparison (four prior to this study) has made population studies difficult. A morphological analysis of the M. repens complex was undertaken to elucidate biological relationships among the species. A key and descrip- tions to the taxa are presented based upon population studies and results of analysis of anatomical variation (Morden & Hatch 1987). MATERIALS AND METHODS Field collections were made during the flowering periods of the species (August through October) in the northern states of Mexico, the souchwest- ern United States, and the Sierra Nevada and Rocky Mountains. To insure isolation between populations, a minimum distance of 10 miles was trav- eled between successive collection sites. In most cases, ten independent specimens from each site were semi-randomly collected and pressed; selec- tion was biased in favor of mature plants that did not show any signs of damage from insects, herbivores, trampling, etc. As always with rhizoma- tous species, the possibility exists that a population is made up of a single clone with specimens displaying mosaic or somatic variation rather than individual genetic variation. Therefore, specimens were collected far enough apart (e.g. 2 meters) within any single population to avoid this possibility. In addition, herbarium specimens on deposit in several herbaria were used to represent species we were not able to collect. In some cases, sufficient material of a single collection was available to serve as a “real” population; in other cases, “artificial” populations were made by selecting specimens from the same geographic region and grouping chem together. A total of 117 populations (real and artificial) were available for analysis. Vouchers for the population scady are deposited at the S. racy Herbarium (TAES). able 1 lists the 34 characters identified that show variation among the populations analyzed. Several characters show variations unique to this com- plex and warrant further description. Plants in this complex are often de- cumbent at the base, and as such culm height will represent culm length. Culm nodules were counted within a 5 mm distance in one surface view of the culm, generally below the inflorescence. Leaf blade curvature was re- corded as a two state character and observed as straight or arcuate; degree of leaf longitudinal posture was recorded as a multistate character and re- corded as flat, involute, or folded. Spikelet length was measured from the base of the spikelet to the apex of the lemma (or the glume if they exceeded the lemma) excluding the lemma awn. Occasionally, a second floret was associated with the floret, and was measured but excluded from the analy- sis. Vestiture of the lemma and palea was recorded as a four state character: MorbeN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 353 TasLe 1. Thirty-four ehatactets used to access Rieck ae vapiaein 3 in 1 the M. npers comple All characters were ‘ } tho State (BS) or multistate (MS). Vegetative: rhizome diameter, rhizome scale length, culm height, culm diameter, culm internode length, culm nodules (MS), leaf sheath length, culm/sheath ratio, ligule length, leaf blade length, leaf blade width, leaf blade curvature (BS), degree of leaf longitudinal posture (MS). Inflorescence: inflorescence length, inflorescence width, nodes per inflorescence (MS), branches per node (MS), inflorescence branch length, inflorescence branch angle (MS), spikelets per inflorescence branch (MS), pedicel length Spikelet: ae length, first glume length, first glume nerves (MS), second — length, second glume nerves (MS), lemma length, lemma awn length, lemma vestiture (MS), palea length, palea vesticure (MS), anther length, caryopsis length, caryopsis width glabrous, scabrous or minutely pubescent, with distinct trichomes (although short and/or sparse), and densely villous. Univariate statistics (mean, standard deviation and range) were performed using SAS computer packages (Goodnight 1979). A nested analysis of vari- ance (ANOVA) was done to compare intrapopulation, interpopulation, and interspecies variation. Multivariate statistics of principal components analy- sis (PCA) and cluster analysis were performed using the NT-SYS package of Rohlf et al. (1980). Principal components were derived using the corre- lation matrices; cluster analysis used the unweighted pair group method using arithmetic averages (UPGMA) on both correlation and distance ma- trices. To assess the degree of divergence among populations, the multi- variate analysis of variance (MANOVA) and canonical analysis programs in SAS were used. RESULTS AND DISCUSSION Initially, all populations were analyzed with PCA and MANOVA to determine various groupings within the complex. Results from PCA indi- cated that all populations, except the nine populations of M. plambea, clus- ter together. By analysis of eigenvector coefficients, the characters identified as contributing to the separation of these two groups were culm nodules (absent in M. plumbea, but present in all other species of the complex at least to a limited degree), leaf blade longitudinal posture (always folded in M. plumbea and flat or involute in other species), and inflorescence width. Two additional groupings were evident when all populations, excluding those of M. plambea, were analyzed using PCA: populations with promi- nently villous florets in the spikelet were clearly separated from those with glabrous or scabrous florets. Results of an analysis of all populations using ANOVA in which the three groups just described are presented in 354 SIDA 17(2) Figure 1. Because M. p/umbea is so clearly distinguishable from the other species of the complex it was not considered in further population analyses. Soderstrom ( 1967) suggested the leaf anatomy of M. plambea might be more closely related to M. wniflora or M. arizonica, a placement in which we are in agreement. Based on the results found here for all species of the complex, the remaining species will be discussed in their traditional group- ings of “glabrous” taxa (macrohairs on the floret absent or not readily evi- dent without magnification) and “villous” taxa (copious macro dent without magnification). Glabrous taxa.—Five taxa with _— airs evi- “glabrous” florets (M. fastigiata, M. repens, M. rvichardsonis, M. squarrosa, and M. utilis, were identified in the initial analysis. An ANOVA and PCA were used to determine which characters accounted for most of the variation in distinguishing among populations. The characters identified were ligule length, spikelet length, first and sec- ond glume length, lemma length, lemma vestiture, palea length and cary- opsis length. Populations of M. richardsonis and M. squarrosa are not consis- tently distinguishable from one another (Fig. 2) on the basis of morphological characters analyzed. This was also corroborated using clus- ter analysis and MANOVA (Morden 1985) and henceforth will be referred to as a single species, M. richardsonis. Populations of M. wti/is and M. repens appear to be clearly distinguishable from those of M. fastigiata and M. richardsonis whereas populations of M. fastigiata and M. richardsonis appear to intergrade in morphological form. To investigate this last point in more detail, analyses were performed on populations of M. fastigiata and M. richardsonts to determine the extent to which these species are similar. The results of PCA (Fig. 3) and cluster analysis (Morden 1985) clearly show these populations to be distinct. Char- acters which consistently distinguish these populations are plant size (culm height, internode length, and inflorescence length) and floret vestiture (M. fastigiata being completely glabrous whereas M. richardsonis is usually sca- brous or minutely pubescent). Inflorescence length and floret vestiture are of particular importance because they may be indicative of a more conser- vative distinction between the two species, whereas differences based only on plant size may be environmentally induced. A cluster analysis using UPGMA was performed on all four of these taxa. Characters which did not account for variation among the taxa as determined from analysis of variance and PCA were eliminated from the analysis. These results indicate that the “glabrous” taxa are represented by four distinct groups, each of which is separated from the others based on morphological characters (Fig. 4). The cluster analysis also depicts M. richardsonis being more closely aligned with M. repens rather than M. fastigiata as was previously hypothesized. The alignment of M. fastigiata and M. MorbDeEN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 355 cy %e ‘ ee e ®e = &° 1 Yj e ‘ e ¥Le-. — e e ap e ee oe? hag Pe ge N ° 98 re} e .3) o ee > e @ ee ses e e © e 38 ” . e e ee ° Vector 1 Fic. 1. Two dimensional representation of all populations of the M. repens complex ana- lyzed with MANOVA. Clusters 1, 2, and 3 represent glabrous taxa, villous taxa, and M. plumbea, respectively. — utilis here may not be entirely unexpected; both species have completely glabrous florets and similar spikelet size, although vegetatively they are very distinct. Villous taxa. —An ANOVA and PCA of the 15 population samples of species with distinctly villous florets, M. vi/liflora and M. villosa, identified ten characters for which there is relatively high level of variation at the species level. These characters were ligule length, leaf blade length, inflorescence branch length and spikelets per inflorescence branch, spikelet length, firse and second glume length, lemma and palea length, and cary- opsis length. The first two components in the PCA (Fig. 5) accounted for 63.4% of the variation present. Most of the populations segregated by species classification within the plot. However, three populations of special interest include Morden 514 (“a”; Fig. 5) Valdes s.n. (‘b”), and Spellenberg 4565('c"). Samples of Morden 514 and Valdes s.n., were collected at the same locality (Rancho Experi- mental “Los Angeles”) in southern Coahuila during different seasons. Analy- sis of the specimens in Morden 514 showed that these plants were very 356 Stipa 17(2) oO . re) | | » ese e o oO ” " ee e = Z e3% a oe a) Oo ‘| e ° e a = e c | a fab) On oO 2 c e a Oo | a a o O oO £ oo 6 al — oO Po Oo ' a a 9°} fa) ra] ims o.0 0 (a) a = oO = re) Qo al QO. fa] = | ra oO Principal Component 1 Fic. 2. Principal components analysis of populations of the “glabrous” taxa using all char- ae acters measured. The symbols present represent the species M. repens (m), M. utilis (@ M. richardsonis (0), and M. fastigiata (Q.). The first and second axes accounted for 53.2% ; of the variation. typical of M. vi/liflora whereas those of population Valdes s.n. were more similar to those of M. vi//osa and clustered with them in the analysis. The distinguishing characteristic between these two species has been plant size; M. villosa is larger than M. villiflora in most morphological characters. It is evident that there exists enough plasticity in these populations that under different growing conditions the plants can exhibit the characters of the other group. The Spellenberg 4565 collection segregated from the rest of M. villosa ina way that indicated it was very distinct from other populations of this spe- cies. In analyzing individual specimens of this population, it was apparent that they represent a larger form with longer leaf blades and sheaths, longer internodes, and more nodes in the inflorescence than is typical of the spe- cies. Results of cluster analysis (Morden 1985) also show this population as segregating distinctly from the remainder of the populations. These results support the conclusion that M. v///iflora and M. villosa can not be consistently separated on the basis of morphological characters and MorbEN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 357 fe) ° f°] o °o N — e} c © Fo o fob fe] ° 5 ie) oO fe) o oO a. fe] oO 0 09° 9 0 £ 90 9 ° 2 6 oO Ps %o 4 oO 6 — oO wo o. oO c= Q. e e ue Pe Principal Component 1 Fic. 3. Principal components analysis of Ara of M. richardsonis(O)and M. fastigtata (@). The first and second axes accounted for 54. | % of the variation as such should be regarded as a single species. In general, M. vi//osa tends to be larger than M. vi/liflora and occupies a different habitat (alkaline and calcareous versus gypsiferous soils, respectively). As such, they have been treated as distinct varieties under M. vi//iflora (Morden 1996). However, these differences may be a result of phenotypic variability induced by the different habitats. Spe/lenberg 4565 may also represent a distinct variety that is geographically isolated from other populations. However, we re- frain from taxonomic recognition of this form until more information about — the population is obtained. Subsequent trips to the collection region by the first author, R. Spellenberg (New Mexico State Univ., pers. comm.), and others have failed to rediscover it, thus its true status may never be re- vealed. KEY TO THE SPECIES . Inflores , lower panicle branches usually greater chan 5.5 cm oe leat ns fol ided or flat, prominent midvein present. M. plumbea . Infl ‘ted, lower panicle branches usually less than 5 cm ae eel open or reflexed; leaf blades flat or involute, a prominent midvein absent 2. Lemma and palea densely villous. 358 SIpA 17(2) / repens richardsonis utilis fastigiata Fic. 4. Cluster analysis of the * eas taxa from the distance matrix with all extraneous ysis eee rs removed from the ana 3. Spikelets usually less than 2.0 mm long; plants of gypsiferous soils of northern Mexico 5a. M. villiflora var. villiflora Spikelets usually greater than 2.0 mm long; plants of alkaline or cal- careous soils, west Texas and New Mexico ............... yb. M. villiflora var. villosa 2. Lemma and palea glabrous or scabrous, not densely villous. {. Mature spikelets 2.7 mm long or greater; glumes greater than 1/2 the length of floret; aan tapering to a mucro or short awn ............0.. . M. repens . Mature spikelets less than 2.7 mm long, seldom greater; glumes 1/2 or - than length of floret; lemma awnless or mucronate, not tapering. 5. Ligule less than 0.6 mm long, seldom to 0.8 mm; inflorescence, at least basally, usually included in uppermost culm sheaths 5. M. utilis 5. Ligule greater than 0.7 mm long, seldom less; inflorescence usually on an exserted peduncle G. Plants usually less than 10 cm tall; leaves less chan 9 mm long: lemma completely glabrous: Andean highlands ........0.c00. 1. M. fastigiata 6. Planes seldom less than 15 cm tall. leaves greater than 10 mm long; lemma scabrous or minutely pubescent, at least caret North America . M. richardsonis he garam fastigiata (J. Presl) Henrard, Meded. Rik Herb. 40:59. 1921. Basionyn: Sporobolus fastigiatus J Presl, Rel. Haenk. 1:241. 1830. Vilfa fastigiatus Meyen, Reis. Erd. 1:484. 1834 (nom. nud.). Typr: Sener Haenke s.n. (HOLOTYPE: PR!; Type fragment: US!). Perennial with scaly rhizomes; rhizome scales (4.5—)5.5—8.1(—10) mm long, acute, often deteriorating with ace Ajlims 246( 1.1) em tall, 23 5) mm diam., mostly erect, branching, glabrous; internodes 1—3(—5) mm long, compressed, nodulose-roughened slightly below inflorescence. Leaves Pp) distichous; sheaths 3—6(—8) mm long, overlapping, margins hyaline. Ligules (0.6—-)0.7—1.1(-1.4) mm long, membranous, acute or slightly rounded, decurrent. Blades (2—)4—8(—9) mm long, 0.6-1.1 mm wide, involute, ar- cuate, glabrous, margins scabrous. Inflorescence a panicle, 1-2 cm long, 1— MorbDeEN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 359 @ aoe @ e S < ® ee c e a4 = C = re) 8| a Oo oO c O = QO. oO oO O Principal Component 1 lous” taxa, M. villiflora( ©) and M. vi//osa Fic. 5. Principal components analysis of the “vi (@). Populations “a” (Morden 514) and “b” (Valdes y Vasquez 5.n.) were collected at the same locality (Rancho Experimental “Los Angeles”, Coahuila, Mexico) in different years and growing seasons. Population “c” (Spe/lenberg 4565) was collected in Otero Co., New Mexico. 3(-4) mm wide, narrow, contracted usually exserted above foliage; inflorescence branches solitary at each node, with 3—7 nodes per inflorescence; branches ascending. Pedicels 0.3—0.8(-1.3) mm long, minutely setose. Spikelets (1.8—)2.0—2.4(—2.5) mm long, usually crowded on branches, |— 5(—8) spikelets on lowermost branches. Glumes (0.6) 1.0=1 41.6) mim long, equal, about one-half the length of spikelet, acute, straw colored to dark green, one-nerved. Lemmas (1.7—)1.9—2.3(—2.4) mm long, 3-nerved, glabrous, narrowing to a mucronate apex, usually dark green. Paleas (1.6) 1.8—2.2(—2.4) mm long, glabrous, dark green. Anthers 0.9—1.6 mm long, yellow. Caryopses ca 1.2 mm long, 0.4 mm wide, narrowly elliptic, dark brown. Chromosome number not reported. Distribution. —Andean highlands of Argentina, Bolivia, and Peru, usu- ally above 3000 m elevation. Muhlenberga fastigiata resembles M. richardsonis in many of its vegetative and reproductive characteristics. However, this species differs from M. 360 SIDA 17(2) richardsonis by its short stature, overlapping and distichous leaves, and florets lacking any vestiture. Za. eeu ae plumbea (Trin.) Hitche., Contr. U.S. Natl. Herb. 17:296,. 1913. Bastony: Vilfa plumbea Trin., Mem. Acad. Imp. Sci. Saint- Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. A(1): ae 1840. Sporobolus plumbeus (Trin.) Hemsl., Biol. Centr. Amer. Bor. 3:546. 1885. rE: “Mineral del Monte, Schlechtendal” Schiede s.n. (HOLOTYPE: LE!; Type oe U S!). Herbarium labels on the ty fragment bear Schlechtendal’s name, however it was probably collected by Schiede be and — and sent to Schlechtendal, and later to Trinius. Perennial with scaly rhizomes; rhizome scales (7.5—)8.5—13(-16) mm long, acute, often deteriorating with age. Culms 13—34(—51) cm tall, (0.4-) 0.8—1.2(-1.4) mm diam., glabrous, freely branching at lower culm nodes; internodes 4—40(—60) mm long, variable, smooth, not nodulose-roughened, nodes green or purple, constricted. Sheaths (10—)15—32(-51) mm long, shorter or longer than internode, margins hyaline. Ligules 0.3—0.5(—0.7) mm long, membranous, truncate, decurrent. Blades (1.7—)3—7(-10) cm long, (1.1—)1.6—2.4(—3.0) mm wide, glabrous, flat or folded along a promi- nent midvein. Inflorescence a panicle, (4—)5—9(—14) cm long, 0.5—4(-8) cm wide, open, upper panicle branches ascending or reflexed, usually on a well exserted peduncle; inflorescence branches solitary at each node, with (6—)8—12(—13) nodes per inflorescence; branches usually lax, occasionally ascending, seldom appearing contracted. Pedicels (0.5—)0.7—1.3(-1.9) mm long, minutely setose. Spikelets (2.4—)2.6—3.2(—3.5) mm long, usually not appearing densely eee on branches, 3-30 spikelets per branch. Glumes (1.0—)1.2—-1.6(-1.9) mm long, equal or second glume slightly longer than first, acute, 1/3—1/2 the lenge h of the floret, usually dark green, one-nerved. Lemmas (2.2—)2.4—3.0(—3.3) mm long, rounded and tapering to an acute apex, glabrous or eee scabrous, dark green, 3-nerves obscure. Paleas (2.2—)2.4—2.8(—3.1) mm long, about equal to the a glabrous, dark green. Anthers (1.4—)1.6—2.0(—2.1) mm long, dark green at maturity. Cary- opses 1.4—1.6 mm long, 0.7-0.9 mm wide, elliptical to slightly ovate, greenish-brown. Chromosome number 27 = 40 (Reeder 1967). Distribution.—Valley of Mexico, and Guatemala. Moist soils in mead- ows, hillsides, or ditches. This species has often been misidentified as M. richardsonis. However, it differs by its larger spikelets, open inflorescence, and flat or folded leaf blades. 3. See wana ee Pres) Hitchc., In: a ie Calif. 1(3):111. 1912. Bastonym: Sporobolus repens J. Presl, Rel. Haenk,. 1830. Vilfa repens (J. Presl) Trin., Mem. Acad. Imp. Sci. Saint-Pétersbourg, a @ Sci. Math., Seconde Sct. Nat. 4(1):102. 1849. Typr: MEXICO, Haenke s.n. (HOLOTYPE: PR!; Type fragment: US! isorypr: MO!), MorbeEN AND Harcu, The Muhlenbergia repens (Poaceae) complex 361 Mubhlenbergia — Nees, Linnaea 19:689. 1847. Type: MEXICO, Aschenbom 206 (Ho- LOTYPE: B-los t; Type fragment: US!). Muahlenbergia abata 1.M. Johnston, J. Arnold Arbor. 24:387. 1943. Type: NEW MEXICO. Valley of the Rio Grande, Wright 1982 (HOLOTYPE: GH!). Hitchcock’s name of M. repens was incorrectly circumscribed when he transferred the name from Sporobolus. However, he did not explicitly exclude the type and in later publications corrected his error. Johnston provided a new name for this species assuming that Hitchcock's name was a later homonym of Sporobolus repens J. Presl and thus illegitimate. Perennial with creeping, scaly rhizomes; rhizome scales (3—)5—9(—20) mm long, acute, often deteriorating with age. Culms (5—)8—20(—42) cm long, 0.3—0.6(-1.1) mm diam., decumbent and usually forming dense mats, freely branching above, glabrous; internodes 4—16(-50) mm long, vari- able, lightly nodulose roughened below inflorescence, usually smooth on lower culm intemodes. Sheaths (2—)6—14(—34) mm long (occasionally longer), shorter to longer than internodes, margins hyaline, “dvenlaspine at base, open near collar and diverging from culm near collar. Ligules (0.1—) 0.4—1.0(-1.8) mm long, membranous, truncate, decurrent, occasionally becoming split. Blades (0.4—)1.1—3.0(—6.0) cm long, (0.5—)0.7-1.5(-4) mm wide, glabrous, involute, straight or usually arcuate to spreading, midvein absent, margins scabrous. Inflorescence a panicle, 1-4(—9) cm long, 1—6(— 32) mm wide, contracted, usually included at base in uppermost leaf sheath; inflorescence branches solitary at each node with (3—)4—8(-10) nodes per inflorescence; branches usually ascending, rarely lax. Pedicels (0.2—)0.5— 1.5 mm long (rarely longer), minutely setose. Spikelets 2.7—3.3(—3.6) mm long (sometimes longer), usually with one floret, occasionally two, not densely distributed, 1—7(—19) spikelets usually on lowermost panicle branch. Glumes (1.1—)1.6—2.4(—3.6) mm long, about equal or first a little longer, acute, 1/2 to equaling length of floret, usually light green, l-, 2-, or 3- nerved. Lemmas 2.6—3.2(—4.2) mm long, acute to attenuate, often narrow- ing into a mucro or short awn, 3-nerved, appearing glabrous or scabrous along midnerve apically, occasionally short hairs are visible near base or along margins, opaque to dark green or mottled; awn 0.1—0.3(-1.4) mm long. Paleas 2.1—-2.7(—3.4) mm long, appearing glabrous or scabrous be- tween the two nerves, color similar to lemma. Anthers (0.7—)1.0—1.4 mm long, yellow, becoming purple at maturity. Caryopses 1.1—-1.5 mm long 0.4-0.7 mm wide, elliptic to ovate, brown, usually with a straw colored coating on pericarp. Chromosome number 2” = 60, 70—72 (Brown 1951; Morden 1985; Reeder 1967, 1968). Distribution.—Southwestern United States and northern Mexico south through the Valley of Mexico to Chiapas. On open sandy grounds of dry mea- dows, canyon bottoms, and along roadsides, often forming dense stands. Soils are variable, ranging from alkaline to calcareous and heavily gypsiferous. 462 Sipa 17(2) This species is similar to M. #t//7s in many respects, such as vegetative characteristics and the nature of the caryopsis. However, M. wi//is can be distinguished by its smaller spikelets, more slender culms and spreading blades. Some populations tend to show intergradation with M. richardsonis. Vegetative apomixis in this species has been previously reported (Morden & Hatch 1986). 4, Muh a richardsonis (Trin.) Rydb., he Torrey Bot. Club 32:600. 1905. Basionym: Vilfa richardionis Trin., Mem. Acad. Imp. Sci. Saint- Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. 4(1).103. 1840. Sporobolus richardsonis (Trin.) Merr., Rhodora 4:46. 1902. Muahlenbergia ie geen aes var. richardsonis (Trin.) M.E. Jones, Contr. W. Bot. 14:12. 1912. Type: NORT AMERICA, Richardson s.n. (HOLOTYPE: LE!; Type G voinent US). Muhlenbergia aspericaulis Nees ex Trin., Mem. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4(1):103. 1840 pro syn. Sporobolus pias ee ex Trin.) Scribn., Bot. Gaz. 21:15. 1896. Type: Richardson s.n. (HOLOTYPE: BM; Type fragment: US!). The types of the basionyms Vi/fa richardsonis Trin. and Wee aspertcaulis N ees ex Trin. may be the same. Vi/fa elie Trin., Mem. Acad. Imp. Saint- ees Sér. 6, Sci. Math., Seconde Prt. Sci. ae. A(1):100. 1840. salina Squarrosa (Trin.) Rydb., Bull Torrey es Club 36:531. 1909. Type: Menzies Island, Columbia River, Washington, Northwest America Dowe/as s.n. (HO- LOTYPE: LE! Type fragmenc: US!). Vilfa depauperata Torr. in Hook., Fl. . Amer, 2: an — 236. 1840. Sporobolus depauperatus (Torr.) Scribn., Bull. ee Bot. C ae 9:1 1882. Type: sandy parts of Columbia River, from Menzies Island upward, he 2. (HOLOTYPE: K; Type frag- ent: US!; tsorype: GH!, NY!). Not Muhlenbergia ae Scribn. (Bot. Gaz. 9:187. 1884). Perennial with scaly rhizomes, often a knotty base, mat-forming; rhi- zome scales (4—)6—9(-13) mm long, acute, often deteriorating with age. Culms typically (S—)L0—30(—64) cm tall, occasionally taller, 0.3—0.6(-1.1) mm diam., wiry, glabrous, decumbent or erect, freely branching above or not; internodes (4—)11—40(—94) mm long, variable, variously nodulose- roughened, occasionally appearing smooth. Sheaths usually (1—)9—22(—50) mm long, shorter or longer than internode, margins hyaline, overlapping at base, open near collar and diverging from culm. Ligules (0.5—)1.0—1.9(— 3.0) mm long, membranous, acute to truncate, erose, or shallowly toothed. Blades (0.4—)1.6—4.0(—6.5) cm long, (0.5—)1.0—1.6(—4.2) mm wide, flat or becoming involute when desiccated, straight or usually arcuate-spreading, midvein absent, margins scabrous. Inflorescence a panicle, (1—)2—5 cm long (rarely up to 15 cm), 1-4(-17) mm wide, contracted, usually on a well- exserted peduncle, inflorescence branches solitary at each node, (4—)7—12(— 18) nodes per inflorescence; branches usually ascending, seldom lax. Pedicels (O.2—)0.5—1.1(—2.0) mm long, minutely setose. Spikelets (1.7—)2.1—2.7(— 3.2) mm long, with one floret, rarely two, crowded on branches, 1—1 3(—40) MorbDEN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 363 spikelets on lowermost panicle branch. Glumes (0.6—)0.9-1.4(—2.0) mm long, about equal or second slightly longer, acute, 1/3—1/2 the length of floret, green, 1- (rarely 2-) nerved. Lemmas (1.7—)2.0—2.6(—3.1) mm long, acute or acuminate, often mucronate, glabrous or scabrous near apex, usu- ally dark green or mottled, 3-nerves obscure, mucro 0.1—0.2 mm long. Paleas (1.0—)1.8—2.4(—2.9) mm oe glabrous or scabrous between nerves, color similar to lemma. Anthers (0.9—)1.2—1.4(-1.6) mm long, yellow, becoming purple at maturity. Caryopses (0.9—)1.1—1.4(-1.6) mm long, 0.3— 0.5(—0.6) mm wide, narrowly elliptic, brown. Chromosome number 27 = 40 (Delay 1950; Morden 1985; Stebbins & Love 1941). Distribution.—This is the most widely distributed species of the com- plex occurring from New Brunswick and Maine, west to Alaska, south through the Sierra Nevada Mountains into Baja California, and the Rocky Mountains to New Mexico and Arizona. On open soils of moist meadows or drier sites away from saturated soils, sandy arroyo bottoms, occasionally alkaline soils Muhlenbergia richardsonis is similar to M. repens with which there appears to be some intergradation in the southwestern United States. There are two intergrading forms of M. richardsonis. One form, which has been re- ferred to as M. sguarrosa, has stout, decumbent, and spreading culms and occupies drier sites with little competition from other species. The other form, referred to as M. richardsonis, has more slender, erect culms and is usually growing in moister soils ina meadow-like association. This species is often confused with M. caspidata (Torr.) Rydb. from the Midwestern U.S., which differs by lacking rhizomes and which has puberulent culm inter- nodes, a shorter ligule, and long glumes which are acuminate or cuspidate. nie ek filiformis (Thurb.) Rydb. is an annual or weak perennial re- ing M. richardsonis, but is distinguishable by its non-nodulose rough- ened as and ovate to acute glumes (these often slightly erose or ciliate along the margins). 5. Muhlenbergia utilis (Torr.) Hitchc., J. Wash. Acad. Sci. 23:453. 193. BasionyM: Vilfa utilis Torr., U.S. Rep. Expl. Miss. Pacif. 5:365. 1858. tees Hie ea U.S.D.A. Div. oe Bull. 17:17 1. f. 467. 1899. Tyee: Lost t. Spring, trip from Tejon to the Lost Hills, cae in oe places, Blake 5.n. lees PE: NY!; Type fragment: US!; tsorypes: GH!, — ci - anes Mex. PI. 2:101. 1886. Sporobolus sacatilla Griseb. ex E. Fourn., PL 1886 pro syn. Types: Cha i as c pr. Mexico, Schaffner 165 pl. ed ae : es a Luis Potosi, Virfet 1455 (P); Texas, Wright (in herb. Durand, P). (TYPE ee US!). Fournier (1886) also listed Agrostis brevifolins Nutt. as a syn- onym. However, this is a synonym of Muahlenbergia cuspidata (Torr.) Ryc o Perennial with scaly rhizomes; rhizome scales (3—)5—9(—13) mm long, 364 Stipa 17(2) acute; scales often deteriorating with age. Culms 7—20 cm high, 0.2—0.6(— 0.9) mm diam., much branched above, erect on smaller plants, decumbent, long and trailing on larger ens up to one meter long, usually ascending, glabrous; internodes (3—)8—22(—37) mm long, variable, lightly nodulose roughened or with siliceous ee ae below inflorescence, smooth or not as pronounced on lower internodes. Sheaths (3—)7—13(—24) mm long, shorter or longer than internodes, margins hyaline, overlapping at base, open near collar and diverging from culm. Ligules (0.2—)0.3—0.5(—0.8) mm long, membranous, truncate, decurrent. Blades (S—)11—24(-47) mm long, (0.2) 0.8-1.3(-1.8) mm wide, glabrous, flat or mostly involute, straight or be- coming arcuate with age, blades of major culms often at right angles to culm, midvein absent, margins scabrous. Inflorescence a panicle, 1—3 cm long (rarely longer), 1-4 mm wide, contracted, usually included at base in uppermost leaf sheath; inflorescence branches solitary at each node, with 3-13 nodes per inflorescence; branches ascending, occasionally becoming lax. Pedicels (O.1—)0.2—0.6(-1.1) mm long, glabrous. Spikelets (1.4—)1.6— 2.0(—2.3) mm long, with one floret, these crowded on branches, (1—)2—7(- 4) spikelets on lowermost panicle branch. Glumes (0.5—)0.6—1.0(-1.4) mm long, about equal, acute, 1/3—1/2 length of floret, straw colored to light green, 1- or 3-nerved. Lemmas (1.3—)1.5—2.0(—2.3) mm long, 3- nerved, acute, not mucronate, green or purple near base at maturity, gla- brous, rarely with short hairs present. Paleas (1.0—)1.4—1.8(-2.0) mm long, glabrous, color similar to lemma or with more purple at maturity. Anthers (0.7—)1.0-1.2(-1.4) mm long, yellow, becoming purple at maturity. Cary- opses (0.7—)0.8-1.0(-1.2) mm long, 0.4—0.6 mm wide, elliptic to ovate, brown, usually with straw-colored coating on pericarp. Chromosome num- ber 2” = 20 (Gould 1966; Morden 1985). Distribution. —United States: Arizona, southern California, Nevada, and central Texas; Mexico: Chihuahua, Coahuila, Durango, Guanajuato, Jalisco, Mexico, Nuevo Leon, Puebla, Sonora, and Veracruz; Guatamala. In wet soils, usually with slow flowing fresh water or along margins of faster flow- ing streams or rivers, depressions in grasslands, and alkaline or gypsiferous plains. This species is similar to ML. repens from which it differs by its more wiry culms, slender leaves, shorter ligules, and smaller spikelets. This species was used by people of Mexico for stuffing packsaddles, hence the common name “aparejo grass.” 6a. Muhlenbergia villiflora Hitchc. var. villiflora, see Morden 1996 (p. 29) 6b. Muhlenbergia eee Hitche. var. villosa (Swallen) Morden, see Morden 1996 (pp. 29, 30). MorDEN AND Hatcu, The Muhlenbergia repens (Poaceae) complex 365 ACKNOWLEDGMENTS This is technical article T. A. No. 25712 of the Texas Agricultural Ex- periment Station. This paper represents a portion of a dissertation submit- ted by the first author to Texas A&M University. We thank Kelly Allred, Paul Peterson, and J. K. Wipff for reviewing earlier versions of this manu- script. Appreciation is expressed to the curators of the following herbaria for specimen loans for this and related studies: A, ARIZ, CAS, DS, F, GH, LL, MICH, MO, MSC, NMC, NY, RM, SD, TEX, US, and UTC, This study was funded in part by the Department of Rangeland Ecology and Management, Texas A&M University, a grant from the Texas A&M Asso- ciation of Former Students, and a grant-in-aid from Sigma Xi. REFERENCES 2 V. 1951. Chromosome numbers of some Texas grasses. Bull. Torrey Bot. Club. =299. a A.H. Hotmacren, N.H. Hoimcren, J.L. REVEAL, and P.K. HOLMGREN. 1977. Intermountain flora: Vascular plants of the intermountain west, vol. 6. New York: Co- lumbia University Pres Deay, C. 1950. Nombres chromosomiques chez les phanerogames. Rev. Cytol. Biol. Veg. 12:1-368. GoopniGutT, J.H. 1979. SAS users guide 1979 edition. SAS Institute Inc., Cary, North ‘arolina. GouLtp, EW. 1966. Chromosome numbers in some Mexican grasses. Canad. J. Bot. 44: 1683-1696. __«d'975. The grasses of Texas. Texas A&M Press, College Station. Hrrcucock, A.S. 1950. Manual of the grasses of the United States. U.S.D.A. Misc. Publi- cation No. 200. Morpen, C.W. 1985. A ps = of the iene repens complex (Poaceae). Ph.D. dissertation, Texas A&M. University, Colleg eee A new Ae nae in pi arenas os Phytologia 79:28—30. and S.L. Hatch. 1986. Vegetative apomixis in Muhlenbergia repens (Poaceae). Sida 11:28 9-285, nd . 1987. Anatomical study of the Sa repens com- plex Poaceae Chloridoideae: Snowe eo. Sida 12:347-35 RrEDER, J.R. 1967. Notes on — grasses VI. MMiscelbencots: chromosome numbers. Bull. es Bot. Club 94:1 . 1968 tes on es oo VUI. Miscellaneous chromosome num- bers-2. Bull. Torrey Bot. Club 95:69-8 Router, EJ., J. KisHpauGu, and D. Kirk. 1980. NT-SYS: eons taxonomy system of multivariate statistical pro ams. New York: University of New York at Stoney Brook. SoperstROM, T.R. 1967. Taxonomic study of subgenus Podosemum a section Epicampes of Mublenbergia Gramineae). Contr. U.S. Natl. Herb. 34:75-189. STEBBINS, G.L. and R.M. ie 1941. A cytological study of California forage grasses. mer. J. Bok 28:371- 366 Sipa 17(2) BOOK NOTICE Raven, Perer H., Ray F Evert, and Susan E. EicHHorn. 1992. Biology of Plants, Fifth Edition. (ISBN 0-7901-532-2, hbk.) Worth Publish- ing Inc., 33 Irving Place, New York, NY 10003, (212) 475-6000, $59.95. 791 pp. Though not a degreed biologist by any stretch, I chose to review this book because I was initially struck by its cover. At first glance, I took it fora book on art history ... well, it is in fact an art book, but rather one on the art of Biology—a textbook on Biology to be exact. Science and art have been successfully blended here. My compliments to the design- ers and artists involved and to the publisher and authors who had the vision and under- standing of how important exciting visuals are to the success of any publication and espe- cially to textbooks. Knowledge is acquired so much more readily with the use of good visual material and it is not lacking here. This is the fifth printing of this well known textbook with more full color throughout the book. Clear and more informative with new findings, chapters have been re-organized for more logical sequence. For instance, the Evolution chapter was moved to directly fol- ee 25 low Genetics which is now in a chapter of its own. Extensive change is seen in Ch due to the many discoveries being made in plant hormones, particularly to molecular basis of hormone action, plant biotechnology and genetic engineering (recombinant DNA) tech- nology. Boxed essays have been included in the Ecology section on the great Yellowstone Fire, on logging of ancient forests in the Pacific Northwest and competition for light in plant communities The fifth edition book maintains its interlocking themes of: 1) the plant body as a dynamic result of processes of growth and development carried out by chemical interac- tions, 2) evolucionary relationships as the guide to understanding form and function in organisms, 3) ecology as a way of cage our dependence on plants to sustain our lives and those of all other living organ To quote the authors ... “The fifth ie of Biology of Plants remains true to the book’s original goals: to present basic information about plants in an accurate, accessible, well- illustrated, and interesting way.”’—Linny Heagy. Sipa 17(2): 366. 1996 ANNOTATED CHECKLIST OF TREES AND SHRUBS OF ALABAMA STEVEN L. YOUNGHANCE 2870 Hwy 171 South Fayette, AL 35555, U.S.A. JOHN D. FREEMAN Department of Botany and Microbiology Auburn University, AL 36849-5407, U.S.A. ABSTRACT A total of 72 families, 163 genera, and 443 species (including 14 represented by at least two subspecies or varieties each) are listed for Alabama. This includes 346 native and 60 naturalized species as well as 37 others believed likely to occur in the state. RESUMEN Se listan un total de 72 familias, 163 géneros y 443 especies (incluyendo 14 que estan eS por dos SUDCSSClES o variedades cada una) para Alabama. Se incluyen 346 especies 2 sy 60 asf como otras 37 que se cree que pueden encontrarse en el ee INTRODUCTION The vascular flora of Alabama is perhaps as diverse as that of any area of comparable size in the eastern United States. Although many new records for species have been obtained for Alabama in recent years, the state re- mains less well documented than any other based upon relative size, geo- logical complexity, and the number of herbarium collections available for study (Holmgren et al. 1990). Much of the published material that deals directly with the Alabama flora is out-dated, out-of-print, nomenclaturally obsolete, or taxonomically inaccurate. Some of it was based (in part) upon observations that apparently remain undocumented by herbarium speci- mens, and so these references are equivocal if not actually misleading. Thus, even for some trees and shrubs chat are aspect dominants, the identity of species in the flora of this state remains somewhat vague and uncertain. CONTENT AND FORMAT OF THE LIST The main reason for developing this checklist was to bring together information from various scattered sources to provide a basis for further floristic work by others as well as ourselves. Our list includes the names of all native and naturalized tree and shrub species that are known to occur Stipa 17(2): 367-384. 1996 368 Sipa 17(2) oe within Alabama to which were added others that seem reasonably likely to be discovered. Documentation for taxa that are listed herein as new for Alabama and for ones previously listed by Clark (1971) without county records are mainly those deposited in the Auburn University Herbarium (AUA) or the University of Alabama Herbarium (ALU/UNA). Because geographical distribution patterns often are helpful in locating additional populations, counties of occurrence are provided for newly listed and pre- viously undocumented taxa. Some of these records, particularly for a num- ber of the uncommon taxa, were obtained either from the computerized database at the Alabama Natural Heritage Section, Department of Conser- vation and Natural Resources (Anonymous 1996), or from cited references. n all-inclusive list of every tree and shrub species of Alabama is not presently attainable simply because the flora of this state, as mentioned above, has not been as thoroughly collected as is the case for its neighbors. Some native species that are more common elsewhere and some that are naturalized or weedy may show up in Alabama and indeed be new taxa for the state, but further additions are more likely to be uncommon species that until now have not been detected. For this reason, names of several woody species from the Vascular Plant Tracking List of the Alabama Natu- ral Heritage Section (Anonymous 1996) were included to encourage fur- ther field studies. Three publications concerning plants of Alabama (Dean 1961; Clark 1971; Diamond & Freeman 1993) provided names for the first version of our list. On account of their content regarding the flora of Alabama, other references that deal with a broader geographical range were reviewed for additional names. These were mainly studies concerning the flora of other states (Radford et al. 1968; Coile & Jones 1985; Jones & Coile 1988; Godfrey 1988; Brown & Kirkman 1990, Wofford & Kral 1993), woody plants of the Southeast (Godfrey & Wooten 1979; 1981; Duncan & Duncan 1988; Foote & Jones 1989; Meyer et al. 1993; Luteyn et al. 1996), and compre- hensive treatments for the U.S. (Little 1979; Elias 1987; Kartesz 1994). With very few exceptions, up-to-date and currently accepted nomencla- ture following that of Kartesz (1994) was used throughout this list, and relevant synonymy was included. The names of plant families used herein agree with the treatment by Zomlefer (1994), which was closely adopted from Thorne (1992). Since earlier lists and much of the available taxo- nomic literature have employed more traditional family names, users of this list should find our inclusion and citation of alternative family names useful. Various notes were used to guide persons trying to deal with Alabama plants to taxonomic treatments in which keys helpful for deter- mining their identity were published. We also used notes to lead readers to YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama 369 references concerning other aspects of many listed species. References that were closely followed in the preparation of this list, whether published prior to or more recently than Kartesz and Zomlefer (both 1994), were also cited in the list itself to make the content more helpful to students. Within the major classification categories used herein, the names of fami- lies, genera, species, taxa of lower rank, and their synonyms are respec- tively arranged in alphabetical order. Author citations for plant names fol- low the standardized abbreviations of Brummitt and Powell (1992). Common names were also included, either with the preferred name first or arranged in alphabetical order when no particular name was evidently “pre- ferred,” for all species and varieties for which such names were available in the literature reviewed. Bailey (1976) served as a major source for common names of naturalized non-native species and also some native taxa that have attracted the attention of horticulturists. Both naturalized non-native species and taxa likely to occur within the state but lacking definite or recent confirmation are denoted by symbols to distinguish them from documented native species. Names of non-native species that apparently survive and reproduce outside of cultivation (by either seed or asexual means) are preceeded by an asterisk (*), whereas other species that seem reasonably likely to occur in Alabama, including some undocumented or not recently collected taxa, are identified by a question mark (?) before the name. A few taxa that may long persist following cul- tivation (such as apple, pear, and peach) are herein listed as “naturalized” as suggested by one reviewer or more, but other woody ornamental and agri- cultural species that seem to be dispersed and established on/y as a result of human activities were mostly excluded in compiling this list. Maintenance of biodiversity in commercial forests as well as in protected areas and preserves is increasingly being recognized as an important long- term objective of forest management, one that also has practical economic benefits in addition to obvious conservation goals (Hansen et al. 1991; Probst & Crow 1991; Burton et al. 1992). Since Alabama has a greater percentage (ca. 68%) of its area in forest than any other southern state (Powell et al. 1993), correct data about the woody species present in this state’s ecosystems are no less than essential if sound management decisions are to be made. For this list to remain useful, it will need to be updated and upgraded from time to time so that gains in taxonomic knowledge and changes in nomenclature for these plants will be readily at hand. Sipa 17(2) LIST OF TREES AND SHRUBS PHYLUM PINOPHYTA (OR CONIFEROPHY TA) GYMNOSPERMS: CONIFERS NOTE: This phylum has been treated as the Class Gymnospermae within the Division Spermatophyta (seed plants) in traditional classi- fication systems. Names of classes of the Pinophyta are not often used in manuals, Horas, guides, etc. used for identification, a all of the mem- bers of this phylum in Alabama are now placed in the Class Pinopsida. Cupressaceae (incl. Taxodiaceae) NOTE: Some authors, including Kartesz (1994), treat the Taxodiaceae as distinct, but most do not. Chamaecyparis thyordes (L.) Britton, Sterns & Poggenb. — Atlantic white-cedar, white- edar, false cypress Juniperus communis (. ground juniper, oldfield common juniper: Calhoun Co. J. virginiana 1. — eastern NOTE: Plants ce ee previously treated as Juniperus silicicola (Small) LH. Bailey are regarded as J. virginiana (Flake et al. 1969; Adams 1986 Taxodinm ditcban (L.) Rich: var. dist — baldcypress var. eed THM {Taxodinm ascendens Brongn. } {Taxodium distichum var. nutans (Aiton) Sweet | var. depressa Pursh — redce Nutt. — pondcypress Pinaceae Pinus clausa 7 hapm. ex Engelm.) Vasey ex Sarg. : ehinetta Mall = -shortleat pine a Walter — spruce pine P cain Mill. — longleaf pine [Pinus australts Michx. | P. serotina Michx. — pond pine *P. strobus L. — eastern white pine: Lawrence, Lee, y, Tuscaloosa, Winston cos. P. taeda L. - "loblolly pine Po virginiana Mill. — Virwinia pine Tivga canadensis (1...) Carriére — eastern hemloc PHYLUM MAGNOLIOPHYTA (OR ANTHOPHYTA) ANGIOSPERMS, HOWERING PLANTS NOTE: This phylum has been treated as the Class Angiospermae within the Division Spermatophyta in traditional classification systems. CLAss MAGNOLIOPSIDA DICOTYLEDONS OR Dicots NOTE: These plants were formerly placed in the Subclass Dicoty- ledoneae of the Class Angiospermae. Aceraceae (see Sapindaceae) Adoxaceae yambucus and Viburnum have usu- NOTE: § ally the srifoliaceae 1 treated as members of Sambucus canadensis L. — elderberry, Ameri- can elde Viburnum scerifolinm L. — mapleleaf vibur- num ’ bracteatum Rehder — limerock arrowwood: Jackson, Madison cos. var. dentatun — southern ar- bracted arrowwood, ot ’ dentatum L. _ rowwood [Viburnum semitomentosum (Michx.) Rehd Vi nudum L. — possumhaw viburnum var. cassinordes (L.) Torr. & A. Gray Viburnum cassinoides L.| var. udu V. obovatum Walter — small-leat viburnum: ouston Co, \V. ae ies — blackhaw Vv. pane ci Schult. — os arrow- wood, Rafinesque viburnum: Etowah Co. V. rufidulum Raf. — rusty blackhaw YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama 371 Anacardiaceae Cotinus obovatus Raf. — American smoketree, wttam-wooc [Cotrnus americanus Nutt.} Rhus avomatica — fragrant sumac R. copallina L. — ‘Gaenelent sumac, shining mac ; saa L. — smooth sumac . hirta (L.) Sudw. — staghorn sumac: Lawrence Co. [Rhus typhina 1.4 R. michauxii Sarg. — false poison sumac, Mi G) x's sumac Toxicodendron pubescens Mill. — poison-oak toxicodendron L. [Toxicodendron quercifolinm (Michx.) S aoe = Greene { Toxicodendron toxicarinm Siete 1. radicans (L.) K [Rhus radia NO hrubby forms of poison-ivy are _ — poison- pene mistaken for poison-oak, s¢ the common names are often ee T. vernix (L.) Kuntze — poison-sumac {Rhus vernix L.] Annonaceae Kral var. spathulata Kral af pawpaw Asimina longifolia & — longlez A. ale (Michx. : Dunal — small Hower awpaw, dwarf pawpaw A, le (L.) Dunal — pawpaw Aquifoliaceae Ilex ambigua (Michx.) Torr. — Carolina holly x cavoliniana (Walter ) Trel. urtis in Chapm. — sarvis r: Mobile ne — = I. amelanchier N (service) Ralls , Washington cos. I. cassine L. — [. coriacea (Pursh) Chapm. — large gallberry 1, decidua Walter — possumhaw L. glabra (L.) A. Gray — inkberry 1. longipes Chapm. ex Trel. — Georgia holly Ilex decidua var. longipes (Chapm. ex Trel.) hles | 21, montana Torr. & A. Gray — mountain holly, winterberr {Ilex ambieua var. montana (Torr. & A. Gray) H.E. Ahles lee monticola A. Gray] I. myrtifolia Walter — aia holly: Covington, Mobile Ilex cassine var. mmyrtifolia | I. opaca Aiton — oo holly L. verticillata (L.) A T. vomttoria Aiton — Walter) Sarg.] stay — winterberry yaupon 1acea (Umbelliferae; incl. Araliaceae) NOTE: Araliaceae has ey use| been regarded as distinct from Apiace Aralia spinosa L. — teenie Her- -club Araliaceae (see Apiaceae) Asteraceae (Compositae) Baccharis angustifolia Michx. — groundsel tree B. baliifelia L. ~— eastern baccharis Borrichia fritescens (L.) DC. — sea ox-eye ee pauciflosculosa aici) Greene — each goldenroc (Soldeep eee elses Michx. } va frutescens L. — marsh elder 1. imbricata Walter — seacoast marsh elder Bataceae ?Batis maritima L. — turtleweed Berberidaceae Berberts canadensis Mill. — American bar- berry: Lee Co *B. sargentiana C.K. Schneid. — Sargent’s barberry: Conecuh Co. Betulaceae Alnus serrilata (Aiton) Willd. — hazel alder ? Betula ero Bricton — yellow birch etitla Intea F. Mic B. lenta L. — sweet bh cherry birch B. nigra L. — river bir Carpinus caroliniana a — American hornbeam, ironwood Corylus americana Waleer — American hazel C. cornuta Marshall — beaked hazel Ostrya virginiana (Mill.) Koch — eastern hophornbeam Bignoniaceae Catalpa bignonioides Walter — southern Catalpa *C. spectosa Warder ex Engelm. — northern catalpa Cactaceae Opuntia humifusa (Raf.) Raf. — prickly-pear cactus {Opuntia compressa J.P. Macbr.] O. pusilla (Haw.) Nutt. — little prickly- pear {Opuntia drummondii Graham] *0. vulgaris Mill. — common prickly-pear Calycanthaceae Calycanthus floridus L. — sweetshrub var. floridus var. glaucus (Willd.) Torr. & A. Gray {Calycanthus ci ne var. ee (WwW I lld. ) Tor Gr Caprifoliaceae (see Adoxaceae) Dyervilla loniera Mill. — bush-honeysuckle: shall Co. D. rivilaris Gatt. — streamside bush-honey- -kle (Dierila He var. rivilaris (Gatt.) H.E. D. es Buckl ley — southern bush-hon- ysuckle Symphoricarpos orbiculatus Moench — coral- erry Celastraceae Enonymus americanus L. — strawberry-bush E. atropurpureus Jacq. — eastern burningbush, wahoo Chrysobalanaceae NOTE: Some authors include this family within the Rosaceae. Lucania michauxit Prance — gopher-apple [Chrysobalanus oblongifolius Michx.} Clethraceae lethra acuminata Michx. — mountain pepperbush: Etowah Co. Sipa 17(2) C. alnifolia L. — pepperbush Clethra alnifolia var. tomentosa (Lam.) Michx. } {Clethra tomentosa Lam.]} Clusiaceae (Guttiferae; incl. Hypericaceae) og NOTE: Hypericaceae has traditionally been sated as a distince fam 7 cum fe ee Spach ) Steud. — astal p t. Joh [Hperi cum A ne se Wi Ild., H. cistifolinm Lam. — roundpod St. Johnswort H. crux-andreae (L.) Crantz — St. erswort [Hypericum stans (Michx. ex Willd) WP. dams & N. Robson} in part] [Ascyrum stans Michx. ex Willd.] H. densiflorum Pursh — bushy St. Johnswort 1. fasciceulatum Lam. — peelbark St. Johnswort H. frondosum Michx. — cedarglade St. Johnswort {Hypericum aureum Bartram} H. galivides Lam. — traw St. Johnswort Hl. hypericoides (L.) Crantz — St. Andrew’s Cross ssp. soars rum bypericoides L.} ee, (Michx. ex Willd.) N. Robso [Hypericum stragulum “XP. Adams & SSP. N. Robson} H. Hoydii (Svenson) W.P. Adams — Lloyd St. in ae St. Johnswort: Randolp| A, nyrufiam Lam. sandhill — myrtleleaf St. Johnswort H. cane Lam. — Carolina St. Johnswort: covington Co. H. nudiflorum Michx. ex Willd. — early St. Johnswort, pretty Sc. Johnswort: Bibb, Choctaw, enry, ee, Madison, H. prolificum L. Rae! St. Johnswort H. reductum (Svenson) W.P. ams — Atlan- tic ie Baldwin, Covington, nee Geneva, Mobile, Washington i Fapeelnn aspalathoides Willd., in part] YOUNGHANCE AND FREEMAN, Trees and shru Et. ee W.P. Adams & N. Robson ineland St. Joh ne pumilum Michx.} nswort Cornaceae (incl. Nyssaceae) NOTE: Nyssaceae has Sas eee been treated as a Cornus alternifolia L.f. — ee dogwood . amomum Mill. — silky dogwood aspertfolia Michx. — roughleaf dogwood . drummondii C.A. Mey. — Drumn wood, roughleaf oo C. florida L. — flowering dozwood C. foemina Mill. — swamp dogwood {Cornus stricta Lam.] s distinct from Corna ae nond dog- ?Cornus racemosa Lam. — gray dogwood {Cornus foemina ssp. racemosa (Lam.) J.S. Wilson} {Cornus paniculata LHér.} Nyssa aquatica L. — wate N. biflora Walter —swamp tupelo, blackgum ssa sylvatica var. biflora (Walter) Sarg.] *N. ogeche Bartram ex — Ogeechee- lime, Ogeechee-plum, Ogeechee tupelo: Barbour Co. r tupelo Marshall : This species was planted along ite Pakeale in Eufaula National Wild- life Refuge to help stabilize eroding shoreline and to provide food for water- fowl. It is not yet proven beyond doubt o be naturalized. N. sylvatica Marshall — blackgum, black tu- pelo, sourgum Cyrillaceae Cliftonia shee (Lam.) Britton ex Sarg. uck Cyiill ee L. — black titi, summer titi, swamp cyrilla at-tree, spring titi Ebenaceae Diospyros virginiana L. —common persimmon Elaeagnaceae *Elaeagnus pungens Thunb. — Russian olive, silverberry *E, ywmbellata Thunb. — autumn olive, au- tumn elaeagnus bs of Alabama 37 Wo Empetraceae NOTE: This family is sometimes (but usu- ally nor) included in the Ericaceae (Luteyn et al. 1996). Ceratiola ericoides Michx. — beach rosemary, sand heath Ericaceae (see Pyrolaceae) OTE: We 2 Famosos follow Luteyn et al. (1996) in this treatment. ?Bejaria racemosa Vent. — tar-flow OTE: This species was listed for’ “Ala.” by Luteyn et al. (1996), but documenta- tion is lacking at both AUA and UNA Epigaea repens L. — trailing arbutus Gaylussacia baccata (Wangenh.) Koch — a huckleberry: wer Co. ews) Tor A. Gray — gre huckleberry G. frondosa (L.) Torr. & A. Gray ex Torr. — blue huckleberry, dangleberry G. mosieri (Small) Camp — hirsute huckleberry G. nana (A. Gray) Small — dwarf dangleberry [Gaylussacia Mies var. nana A. Gray} G. tomentosa (A. Gray) Small — dangleberry {Gaylussacia frondosa vat. tomentosa A. ao . dumosa (Andre Gray} Kalmia hirsuta Walter — wicky, hairy laurel K. latifolia L. — mountain-laurel Lencothoe axillaris (Lam.) D. Don — swamp fecterbush (St Sleumer — dog-hobble a ialee gore var. editorum (Fernald Schub.) H.E. Ahles} { Leacothoe editorum Fernald & B.G. Schub.]} L. racemosa Gray — fetterbush ?L. recurva (Buckley) A. Gray — mountain fetcerbush onta ligustrina (L.) DC. — male- cl L. lucida Koch — — Oxydendrum arboreum (L.) DC. — meee Pieris phillyreifolia ee DC. — vine- wicky, climbing fetterbush NOTE: Growth habit of this shrub is commonly eee heen’ on the bark of baldcypre Rhododendron al, (Lam.) shamense Rehder — Alabama rhododendron R. arborescens (Pursh) Torr. — smooth azalea, sweet azalea 374 ?R. ans wn (Ashe) Rehder — dwarf azalea TE: A few collections from south-cen- a Alabama were widely distributed and labeled as R. atlanticum. We regard those populations to be hybrid swarms of R. alabamense X BR. canescens. R. austrinum (Small) Rehder — Florida aza- ea, orange azalea, southern azalea 2B. iene oe ans ) Torr. — flame azalea "E: Credited to Alabama by Luteyn et nee (1996) and others. The Alabama re- cords seem to represent R. camberlandense. R. canescens (Michx.) Sweet — Piedmont aza- lea, mountain azalea, southern pinxter- o pe} n azalea, woolly aza R. catawhiense Mi bloor ichx. — Cati ne i rhododen- a) ron, rosebay R. cumberlandense E.L. Braun — Cumberland azalea, diploid flame azalea [Rhododendron bakert of Alabama authors, not (Lemmon & McKay) Hume] ?R. maximum nL. — roseb say, great laure NOT E: Credited to Alabama by Luteyn etal. (1996), but documentation is lack- ing at AUA and UNA R. minus Michx. — dwarf rhododendron, rose- . i 8. periclymenoides (Michx.) Shinners — ees oom azalea, pinxter-flower, elec- tion-pink: Clay, Crenshaw, Elmore, Lawrence, Randolph cos. sea on nudiflorum (L.) Tor !: Considered rare in eee by . (1996), The county records above probably represent R. canescens. R. prunifolium (Small) Millais — plumleaf ie et al lea R. viscosum (L.) Torr. ~_ OTE: The varieties of this species are distniewisned by habit and habitat dif- ferences that are not preserved well in her- barium specimens. Many authorities fail to recognize them because these distinc- tions are not evident in the material on which past taxonomic judgements were usually made. var. serrufatum (Small) H.E. Ahles — ham- mock sweet azalea: aria Conecuh, Covington, Mobi S. [Rhododendron nil apy (S Millais} mall) IDA 17(2) var. viscosum — ge azalea: Chilton, clay, Henry, Lee, Tuscaloosa cos. JaCCIRLUMM Marshall — sparkleberry, farkleberry a arborenum tree V. corymbosum L. — highbush blueberry {Vacemninm asher Reade {Vaccinium atrococcum (A. Gray) A. Heller} {Vaccinium fuscatum Aiton {Vaccinium virgatum Aiton] NOTE: Vander Kloet (1980) and Luteyn . (1996) treated Vaccinium ashe, V. elliotit, V. fuscatum, and V. vir ingatum (V. amoenium ee as : art of the — et al atrococcum, V. Al aad kl comp dle follow Godfrey (19 these blueber- ries and maintain V. e//rotris as distinct because it can easily be identified by morphological charac ters in the field. — ?V. crassifolium Andrews — creeping blue- berry V. darrowii Camp — Darrow’s blueberry, dull licclele - oo evergreen blueberry V. elfiottir Chapm. — Elliortt’s blueberry Vier apn ie bearberry, moun- tain cranbe ITE: Credite d to Alabama based upon its listing by Dean (1961), but documen- tation is lacking atc AUA and UNA V. myrsinites Lam. — ground blueberry, shiny littleleaf blueberry V. pallidum Aiton — hillside blueberry, low- bush blueberry, Blue Ridge blueberry { Veecernium vacillans V. staminenm LL. erry, squaw eenty [ Vaccinium be ee C.M [Vaccinium staminenm var, melanocarpum (C Mohr) C. Mohr} V. tenellum Aiton — slender blueberry, south- Kalm ex Torr. } ~ ern blueberry Euphorbiaceae Andrachne phyllanthoides (Nutt.) Coult. — maidenbush Croton alabamensis E.A. Sm. ex Chapm. var. dlabamensis — Alabama croton *Sapium sebiferum (L.) Roxb tallowtree, — Chinese popcorn-tree Sebastianta friticosa (Bartram) Fernald bastian-bush [Sebastianta ligustrina (Michx.) Mill. Arg. } YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama 375 Sen aquatica Chapm. — corkwood *Vernicia fordii (Hemsl.) Airy-Shaw — tungoil-tree [Aleurites fordii Hemsl.] o + Fabaceae (Leguminosae) NOTE: We follow Isely (1990) in this treat- ment. * Albizia julibrissin Durazz. — silktree : morpha fruticosa L. — indigobush ! f oir. — mountain indigo 2A. nitens Boynton — shiny indigobush schwerinit C.K. Schneid. indi h — Schwerin’'s Cercis canadensis L. — eastern redbud Cladrastis kentukea (Dum.Cours.) Rudd — yellowwood {Cladrastis lutea (F. Michx.) Koch *Cytisus scoparius (L.) Link — Scotch broom Erythrina herbacea L. — eastern coral-bean ?Gleditsia aquatica Marshall — waterlocust triacanthos L. — honeylocust Cea dioicus (L.) Koch — Kentucky coffeetree: Madison Co. NOTE his species is questionably na ee: bicolor Turcz. — bicolor lespedeza *L. thunbergii (DC.) Nakai — Thunberg espedeza Parkinsonia aculeata L. — cee eas Robinia a en istly locus NOT Rosinia aren and ae spp. of ae are very complex with many cul- tivated and naturalized forms that for- merly were treated as species but now are probably best regarded as varieties; sev- eral of these are established and possibly naturalized in Alabama. *var. fertilis (Ashe) Clausen [Robinia - tilis Ashe] *var, hispida var. Eliot — ex Hutch.) Isely — Kelsey loc [Robinia ‘ae Cowell ex Hutch.} var. nana fee DC. — downy locust, Elliott locust se elliottii Oe ta ) Ashe} t} Robinia nana Elli Sesbania puntcea (Cav.) B *var. rosea Pursh [Robinia boyntonii Ashe — Boynton ocust R. ecm ta L. — black locust *R. viscosa Vent. ex Vauquelin var. ate egii (Koehne) Ashe — Hartweg scosa — clammy locust enth. — purple ses ae > an Daubentonia punicea (Cav.) D Wisteria floribunda (Willd.) DC. wisteria — Japanese he common ie habit of , but shrubby — Wi Sterld Spp. 18 as a rms also occur in nt ‘of the species. W. eee (L.) Poir. — American wisteria *W. srnensis (Sims) Sweet — Chinese wisteria Fagaceae Castanea dentata (Marshall) Borkh. — Ameri- ca tnu *, mollissima Blume — Chinese chestnut C. pumila (L.) Mill. rar, ozarkensis (Ashe) Tucker — Ozark chinquapin {Castanea ozarkensis Ashe} {Castanea alabamensis Ashe] var. pumila — Allegheny chinquapin [Castanea abnifolia Nute. — trailing chinquapin] Fagus grandifolia Ehrh. — American beecl *Onercus acutissima Carruth. — sawtooth oak NOTE: Quercus is the largest tree or shrub genus in Alabama. Clark (1971) recog- nized 28 species of Quercus, including four with two varieties each. A number of taxa that were then considered to be conspecific are currently accepted as dis- tinct species by many authors, and their names together with additions such as the one above account for the total of 38 Quercus species listed herein. . alba L. — white oak arkansana Sarg. — re oak O10 {Onercus caput- a he} Q. austrina Small — bluff bastard white oak {Ouercus durandii var. austrina (Small) E.J. Palmer} 376 Q. bicolor Willd. — swamp white oak boyntonii Beadle — running post oe Boynton oak: Bibb, Etowah, $ Q. chapmanii Sarg. — Chapman oak Q. coccinea Miinch k O. falcata . geminata Small — sand live oak (© Shelby cc « h. — scarlet oa ichx. — southern red oak ri tO Quercus virginiana var. geminata (Small) Sarg. ] ee virginiana var. maritima of au- thors, not (Michx.) C.H. Mill.] georgiana M.A. Curtis — Georgia oak: Chambers, St. Clair . bemisphaertca ae ex x Willd. var. hemisphaerica — Darlington oak var. maritima (Michx.) C.H. Mull. — coast ive oak LOwercus maritima (Michx.) Willd.] [Owercus a ante var. maritima (Michx.) C.H. Miill.] _ mbricavia aris - eee oak to a toto incana Bartram — bluejack oak [Quercus cinerea Michx laevis Walter — turkey oak {Quercus sai Michx. } . laurifolia N — laurel oak {Quercus aa elle 1.) Ashe} alter — overcup oak, swamp oe o~ (O = Kk = Q ~ a Q a: . macrocarpa Michx. — bur oak, mossy-cup oak: Montgomery, Pickens cos t© CN a Q. margaretta Ashe ex Small — sand post oak ITE: Kartesz (1994) used Qvercus margarettiae, but Ashe’s original spelling is accepted here [Quercus stellata var. margaretta (Ashe ex Small) Sarg. } Q. marilandica Miinchh. — blackjack oak Q. michauxti r utt. — swamp chestnut oak, ~ — t oa . minima (Sarg.) Small — dwarf live oak: win Co. . muehlenbergii Engelm. — chinquapin oak uercus prinoides var. acuminata (Michx.) Gleason] ene Willd. — myrtle oak neva L. yak pagoda Raf - “herb ark oak {Quercus falcata var. pagodifolia Elliott} [Quercus eee (Elliott) Ashe] tO to S to Sipa 17(2) 2Q. palustris Miinchh. — pin oak phellos L. — willow oak oo Willd. — ee oak . print chestnut o me montana Willd] lias Walter — running oak ubra L. — ae red oak ray) Fernald FE. Michx. } sass Dil! var. ambigna HerCMS ee var. rubra [Quercus maxima (Marshall) Ashe] Q. shumardii Buckley Q. sinuata Walter — shallow-lobed oak, bas- tard oak var. brev — (Torr.) C.H. Mill. — white shin ee breviloba (Torr.) Sarg.] [Quercus durandit var. breviloba E.J. Palmer] var. s/nvata — Durand white oak: Dallas Co. {Quercus durandti Buckley var. durandit Q. stellata Ae eau post oak Q. texana Buck exas ae Nuttall oak { Overcus meer E.J. Palmer] Q. velutina Lam. — black oak Q. virginiana Mill. — live oak — Shumard oak (Torr.) Grossulariaceae NOTE: Itea and Ribes have traditionally been placed in the Saxifragaceae. Itea virginic — Virginia willow Ribes curvatum Seal — granite gooseberry R. cynosbati L. — dogberry, prickly gooseberry Hamamelidaceae Fothergilla gardenit Murray — dwarf witch- alder F. major (Sims) Lodd. — witch-alder Hamamelis virginiana L. — witch-hazel Liquidambar styraciflua L. — sweetgum Hippocastanaceae (see Sapindaceae) Hydrangeaceae NOTE: Hydrangea and Philadelphus have traditionally been placec Saxifragaceae : YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama 377 Hydrangea arborescens L. — wild hydrangea, smooth hydrangea H. cinerea Small {Hydrangea arborescens ssp. discolor (Ser. ex DC.) E.M. McClint quercifolia Bartram — oakleaf hydrangea Phan as Nutt. — hairy mock- orange P. ae L. — mock-orange ?P. pubescens Loisel. — hoary mock-orange Hypericaceae (see Clusiaceae) Illiciaceae NOTE: Some authors place [//iciwm within the Magnoliac Ilicinm flori bans, ‘Ellis — Florida anise-tree Juglandaceae Carya alba (L.) Nutt. ex Elliott — mockernut hickory {Carya tomentosa (Lam. ex Poir.) Nutt.] C. aquatica (F. Michx.) Nutt. — water hickory [Juglans aquatica F. Michx.} C. carolinae- i eshaneee (Ashe) Engl. & Graebn. hickory, southern shagbark hickory [Carya ovata var. australis (Ashe) Little] C. cordiformis (Wangenh.) Koch — bitternut ickory C. glabra (Mill.) Sweet var. glabra — pignut hickory var. hirsuta (Ashe) Ashe — swamp hickory {Carya leiodermis Sarg.] *C, illinoensis (Wangenh.) Koch — pecan {Carya pecan (Marshall) Engl. & Graebn. } C. laciniosa (F. Michx.) Loudon — shellbark hickory C. myristiciformis (PF. Michx.) Nutt. — nut- meg hickory C. ovalis (Wangenh.) Sarg. — red hickory, sweet pignut hickory [Carya glabra var. odorata (Marshall) Little} — U C. ovata (Mill.) Koch — shagbark hickory C. pallida (Ashe) Engl. & Graebn. — sand jos cinerea LL. — sere white walnut J. nigra L. — black walnu Lamiaceae (Labiatae) Calamintha coccinea (Nutt. ex Hook.) Benth. — red basi {Clinopodium coccineum (Nutt. ex Hook.) Kuntze {Satureja coccinea (Nutt. ex Hook.) Bertol.] C. georgiana (R.M. Harper) Shinners — Geor- gia basi [Clinopodium a. R.M. Har [Satureja georgiana (R.M. Harper) HL. E. Ahles} le lear ei (Torr. & A. Gray) A. Gray se rosemary, wild rosemary Lauraceae Haale benzoin (L.) Blume — spicebush melissifolia (Walter) Blume — hairy pets ush NOTE: Probably extirpated from Ala- bama. The only record is one from Wilcox County in 1840 (Freeman et al. 1979). Le Paes B.E. Wofford — pondberry: Mobile ?Litsea pee (L.) Fernald — pond-spice Persea ee (L.) Spreng. — redba P. palu ae Sarg. — swamp redbay, : Baldwin, Crenshaw, Dale, eee Conewar Mobile, Pike cos {Persea borbonia var. pubescens (Pursh) Little] Sassafras albidum (Nutt.) Nees — sassafras Leitneriaceae ?Leitneria floridana Chapm. — corkwood Loganiaceae *Buddleja lindleyana Fortune ex Lindl. — butterfly bush Loranthaceae Phoradendron Hiei ie i Reveal & M.C. Johnst. — mist [Phoradendyon serotinum ce ) M.C. Johnst.} Lythraceae Decodon verticillatus (L.) Elliott — swamp oosestrife *T agerstroemia indica L. — crapemyrtle 378 Magnoliaceae (see Iliciaceae) Liriodendron tulipifera L. — yellow-poplar Magnolia acuminata L. — cucumbertree [Magnolia cordata Michx.} M. grandiflora — southern magnolia M. macrophylla Michx. — bigleaf magnolia M. pyramidata Bartram — pyramid magnolia [Magnolia fraseri var. (Bartram) Pamp M, try, M. virginiana i pyramidata vetala L.. — umbrella magnolia — sweetbay Malvaceae = * Hibiscus syriacus L. — rose-of-Sharon Meliaceae Melia azedarach L. — Chinaberry Moraceae (see Urticaceae) Myricaceae NOTE: The nomenclature and taxonomic treatment for Myricaceae follows Wilbur (1994). ?Comptonia peregrina (L.) Coult. — sweet fern Morella caroliniensts (Mill.) Small — evergreen bayb [Myri vita ideale Rat.} M. cerifera (L.) Small — southern bayberry, waxmyrtle {[Myrica certfera L.} [Myrica cerifera var. pumila (Michx.) Small ] M. tnodora (Bartram) Small — odorless bay- Derry [Myrica tnodora Bartram] Nyssaceae (see Cornaceae) Oleaceae Chionanthus virginicus L. — fringetree, grandsir graybeard Forestiera acuminata (Michx.) Poir. — swamp- privet E a (Michx.) Poir. — upland swamp- American ash, white ash [Fraxinus oe eana Beadle} Sipa 17(2) EB caroliniana Mill. — Carolina ash, pop ash XINUS ae _ Nutt. } F. per nNS) oink rshall — green ash, red ash [Praxinus see ata Borkh E. profunda (Bush) Bush — pumpkin ash: Cullman, Madison cos. [Praxinus tomentosa F. Michx.] E. quadrangulata Michx. — blue ash *Lieustrum japonicum Thunb. — Japanese privet *L. sonense Lour. — Chinese privet Osmanthus americanus (L.) Benth. & Hook.f. xray — devilwood Platanaceae Platanus occidentalis L. — American sycamore Polygonaceae nas Polygonella americana Small — sou Pisch. & C.A. Mey.) ichern jointweed P. eli Small — largeleaf jointweed: Baldwin Co. P. polygama (Vent.) Engelm. & A. Gray — jointweec {Polygonella croomit Chapm. } Pyrolaceae NOTE: Members of the genus Chimaphila are subshrubs that have traditionally been Gaultheria, Epigaca (both Ericaceae) and Chimaphila all share similar growth form and ever- green foliage, but both Gau/theria and Epigaea are m Ericaceae. placed in the ore commonly treated as shrubs than Chimaphila Chimaphila maculata (L.) Purs wintergreen, pipsissewa 1 — spotted Ranunculaceae Nanthorbiza stmplicissima Marshall — yellowroot Rhamnaceae Ceanothus americanus L. — New Jersey tea C. microphyllus Michx. — littleleaf redroor, littleleaf ve horn > f, acl alter — Carolina buck- thorn R. lanceolata Pursh — lanceleaf buckthorn YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama ?Sageretia minutifiora (Michx.) Trel. — calciphile buckthorn NOTE: The only record for Alabama ts a ) collection from Mobile *Ziziphus jujuba Mill. — common jujube Rosaceae (see Chrysobalanaceae) Amelanchier arborea (F. Michx.) Fernald var. alabamensis (Britton) G.N. Jones — Ala- bama serviceberry, Alabama shadbush A. canadensis (L.) Medik. — thicket service- 2A, ee (Michx.) Ashe — coastal plain serviceberry, shadbush NOTE: Credited to Alabama by Dean ( ene l and others, but collections of this e lacking at AUA and UNA. Aronia arbutifolia ( (L. pee —red chokeberry [Pyrus arbutifolta (L.) [Sorbus pa ae (L.) aa 1] Crataegus aestivalis (Walter) Torr. & A. Gray —apple haw, May hawthorn C. brachyacantha Sarg. & Engelm. — blue- berry hawthorn C. chrysocarpa Ashe — fireberry hawthorn, scarlet hawthorn a coccinea L., in part} C. crus-galli cockspur hawthorn C, fala ae sach) Kirchn. — fanleaf haw- C. es ava Aiton — yellow hawthorn C. har ue Beadle — Harbison hawthorn: illman Co C. marshallit Ege. — parsley hawthorn C. ee ea (L.f.) Medik. — red hawthorn, ngton-thorn ?C. pile beans Ashe — beautiful hawthorn . punctata Jacq. — dotted hawthorn Ms a i ai Michx. — littlehip hawthorn threeflower hawthorn CG uniflora NMeachh.: — oneflower hawthorn, dwarf hawthorn ), triflova Chapm C. viridis L. — green hawthorn Matus i eel (Aiton) Michx. — south- ern crabapple [Pyris ne ole Aiton} *M. pumila Mill. - common apple M. coronaria (L.) — sweet ope [Malus bracteata nen Bailey) Rehder] [Malus elaucescens Rehder} [Pyrus coronaria eee. Neviusia alabamensis A. Gray — Alabama snow-wreath Physocarpus opulifolius (L.) Maxim. — ninebark Prunus ina C. Mohr — Alabama black see {Prunus serotina var. alabamensis (C. Mohr) Little} P. americana Marshall — American plum P. angustifolia Marshall — seca plum fill.) A laurelcherry, ap P. mexicana 8. Watson — Mexican p P. nigra Aiton — Canada plum {Prunus americana var. lanata Sudw.) {Prunus americana vat. caroliniana (M on — Carolina um nigra (Aiton) Waugh} {Prunus lanata (Sudw. . Mack. & Bush] *P_ persica (L.) Batsch. — peach P. serotina Ehrh. — black es wild cherry P. umbellata Elliott — flatwoods plum, hog m, sloe plum *Pyracantha coccinea M. Roem. — =P, ae (Maxim.) H.L. L {Pyracantha crenato-serrata (Hance) ae} firechorn Rehder} *P. koidzumii (Hayata) Rehder rus calleryana Decne. — Bradford pear *P. communis L. — common pe *Rosa bracteata J.C. Wendl. - eeu se, he a rose *R, canina L. — dog rose: Mobile Co. R. carolina L.- Carolina rose, wild rose [Rosa sede Marsha R. eglanter — sweetbrier rose Rosa aa ie L.} *R, se Michx. — Cherokee rose ?R. moschata Herrm. — musk rose KR, mai hunb. — baby rose, multiflora R. pabusri Marshall — swamp rose R. set chx. — prairie rose, climbing rose R. virginiana Mill. — Virginia rose ER, eased Cré NOTE: ‘Dor teera Mi — ép. — memorial rose othy Perkins’ is the cultivar ea syle from old homesites and along highway Rubus allegheniensis eae — Allegheny blackberry: Madison Co. R. argutus Link — common blackberry, sawtooth blackberry 380 [Rubus betulifolins Small} [Rubus floridus Tract. *R. bifrons Vest ex Tratt. — European black- berry, Himalayan berry: Bibb, Tuscaloosa cos. R. canadensis L. — smooth blackberry, thorn- less blackberry: Clay, Lee cos R. cunetfolius Pursh — sand blackberry R. flagellaris Willd. — northern dewberry R. hispidus L. — bristly dewberry, swamp dewberry *R. occidentalis L. — ?R. odoratus LL. raspberr NOTE: wore from Jackson Co. by Dean (1961 R. trivialis Michx. — southern dewberry black raspberr — thimbleberry, flowering * Spiraea cantoniensis Lour. — Reeves spiraea *§. thunbergit Siebold ex reath S. tomentosa L. 28 Blume — bridal — hardhack _ virgimana Britton — Virginia spiraea Rubiaceae Eagan occidentalis L.. — buttonbush oneya bracteata (Bartram) Raf. — pinckneya, fever-tree {Pinckneya pubens Michx.} Rutaceae *Poncirus trifoltata (L.) Raf. — trifoliat — trifoliata L. — common hoptree, a t- ee americanum Mill. — common prickly-ash L. — Hercules-club [Zanthoxylum macrophyllum Nutt.} Z. clava-herculis Salicaceae Nae alba L. — white poplar ? deltordes Bartram ex Marshall — eastern cottonwooc . heterophylla L. — swamp cottonwood P. nigra L. — Lombardy popla *Salix babylonica L.. — weeping lion S, caroliniana Michx. — Coastal Plain willow ae — Shuttlew. ex E.S. Anderson} v4 ssy willow, large gray wil- — Missouri willow: cto the large number of Alabama records Sipa 17(2) cited by Argus (1985, 1986) for this spe- cies should be added Calhoun, Coffee, Covington, Dale, Escambia, Geneva, and Monroe cos. 2S. floridana Chapm. — Florida willow S. humilis Marshall — prairie willow, upland willow S. nigra Marshall — black willow 2S. rigida Muhl. — heartleaf willow S. sericea Marshall — silky willow Santalaceae Nestronta umbellula Rat. — nestronia Pyrularia pubera Michx. — buffalo nut Sapindaceae (incl. Aceraceae and Hippocastanaceae) NOTE: Aceraceae and Hippocastanaceae have traditionally been regarded as dis- from Sapindaceae. Acer barbatum Michx. — Florida maple [Acer floridanum (Chapm.) Pax] [Acer sai var. floridanum (Chapm.) Small & A [Acer saccharum ssp. floridanum (Chapm.) Desmarais A. leucoderme Small — chalk maple Heller r saccharum var. leucoderme (Small) Rehder] {Acer saccharum ssp. leucoderme (Small) i a smarais } A. negundo L. — boxelder A. nigrum F. Michx. — black maple [Acer saccharum var. nigrum (F. Michx.) Small [Acer il: ssp. Desmara 2A. eae L. — striped maple A. rubrum L. — red maple var. rubrum var. drwmmondit (Hook. & Arn. ex Nutt.) Sar nigrum (FE. Michx.) arg. [Acer drummondii Hook. & Arn. ex Nutt.] A. saccharinum UL. A. saccharum Mars NOTE: — silver maple all — sugar maple me eee follow Desmarais (1952) “ne treat A. barbatum, A. nigrum, and A. leucoderme as subspecific taxa within an A, saccharum complex. YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama Aesculus flava Soland. ex Hope — yellow buckeye [Aesculus octandra anarer A. glabra Willd. — Ohio A. parviflora Walter — Borla buckeye A. pavia L. — red buckeye A. sylvatica Bartram — es buckeye [Aesculus georgiana Sarg.} * Koelrenteria paniculata Laxm. — golden rain tree ge ae! Hein L. — wingleaf soapberry, oapberry iSapindies marginatus Willd.} Sapotaceae NOTE: Arca (1991) treats Bumelia as synonymous with oo Except for ee i994 ¢ very few American authors have done likewise for more than acentur Sideroxylon lanuginosum Michx. — gum buck- thorn, gum bully [Bumelia lanuginosa (Michx.) P S. /ycioides L. — buckthorn, buckthorn bully (Bumelia /ycioides (L.) Pers S, reclinatum Michx. — Saad buckthorn: Mobile Co. [Bumelta reclinata (Michx.) Vent. } S. thornei (Cronquist) T.D. Penn. — swamp kchorn: Houst on Co {Bumelia thornei Cronquist} Saxifragaceae (see Grossulariaceae and Hydrangeaceae) Scrophulariaceae *Panlownia tomentosa (Thunb.) Siebold & Zucc. ex Steud. — princess-tree Simaroubaceae *Adlanthus altissima (Mill.) Swingle — ail- nthus, tree-of-heaven Solanaceae Lycium carolinianum Walter — Christmas- berry, Carolina wolf-berry Staphyleaceae Staphylea trifolia L. — American bladdernut Sterculiaceae *Firmiana simplex (L.) W. Wight — Chinese asol- ree, varnish-tree tFirmiana platanifolia (L.f.) Schott & Endl.] Styracaceae NOTE: The nomenclature used for Halesia follows Reveal and Seldin (1976) Halesia carolina L. — little silverbell [Halesia parviflora Michx.} H. diptera J. Ellis — two-wing silverbell H. tetraptera J. Ellis — Carolina silverbell ae hieadg of authors, not L.]} St am. — American snowbell [Sry vax x pier wlentus Michx. NO r of the cas shea has cae a source ane . It is neuter according to Wood and ee _ ee ) and Howard (1974); feminine according to Gonsoulin (1974); and masculine ac- cording to Nicholson and Steyskal (1976). We believe Nicholson and Steyskal to be correct. S. grandifolius Aiton — bigleaf snowbell Symplocaceae Symplocos tinctoria (L.) L Hér. — horse sugar, ‘Theaceae Gordonia lasianthus (L.) J. Ellis — loblolly-bay Stewartia malacodendron L. — Virginia stewartia [Stuartia malacodendron (L.) LHér} S. ovata (Cav.) Weath. — mountain camellia [Stuartia pentagyna L’Her. Thymelaeaceae Dirca palustris L. — leatherwood Tiliaceae NOTE: Some authors consider Tilia caroliniana, T. heterophylla, and T. mithauxti to be synonymous with T: americana (Hickok & Anway 1972). We follow Hardin (1990) in this treatment. 382 Tilia americana UL. var. americana — American basswood, American linden var. caroliniana (Mill.) Castigl. — Carolina basswood Tilia cavoliniana Mill.| var. heterophylla (Vent.) Loudon — white dasswood [Tilia heterophylla Vent.) {Tilia michauxii Nutt. — Michaux basswood | Ulmaceae Celtis aes (Jacq.) Sarg be Cc. —— Willd. — sugarberry C. occidentalis L. — common hackberry C. tenuifolia Nutt. — dwarf hackberry Celtis georgiana Smal [Celtis mississippiensis Bosc] [Celtis smalli (Beadle) Sarg.] [Celtis tenuifolia var. ‘ernald & B.G. Planera aquatica J.F. Gmel. — water-elm — iguana hack- Lamm] at georgiana (Small) Schub. | _ Ulmus alata Michx. — winged elm U. americana L. — American elm, white elm U. rubra Muhl. — slippery e [Ulmus fulva Michx.} U. seroting Sarg. — September elm m 2U., thomasit Sarg. — rock eln Ulmus racemosa D. Thomas] Urticaceae (incl. Moraceae) NOTE: Moraceae has traditionally been Urticacea *Broussonetia papyrifera (L.) Vent. — paper mulberry * Maclura pomifera (R treated as distinct from af.) C.K. Schneid. — Osage-orange, bois d’are NC TPE: Historical evidence suggests that this midwestern species did not occur east of the Mississippi River prior to Euro- pean settlement (Elias 1987) Americans may earlier have extended the , but native original geographic range through trade because the tree was so much valued for bow-wood. SipA 17(2) *Morus alba L. — white mulberry M. rubra L. — red mulberry Verbenaceae Callicarpa americana L.— American beautyberry f —common lantana Bonpl. & Kunth — spiny *Lantana camara L. *L. horrida Humb., lantana * Verbena brasiliensis Vellozo — tall verbena * Vitex agnus-castus L. — chaste-tree CLaAss LILIOPSIDA MONOCOTYLEDONS OR MONOCOTS NOTE: These plants WCrC formerly classified as the Subclass Monocoryledoneae within the Class Angiospermae. Agavaceae NOTE: ° been included within the Liliaceae The genus Yucca has traditionally Yucca aloifolia L. —Spanish dagg sialon Y. filamentosa L. — Adam's needle [Yucca flaccida Haw.] [Yucca filamentosa var. smalliana (Fernald) H.E. Ahles ucca ; bear-grass Let LYneca smattiana Fernalc Y. gloriosa L. — Spanish bayonet, moundlily yucca ?Y. recurvifolia Salisb. — curveleaf yucca Arecaceae (Palmae) pak a ie 2 stvix (Pursh) H. Wendl. & Drude ¢ ude — needle palm : an minor (J Pers. — blue palm, dwarf palmetto any ie: Lodd. ex Schult. — tac pal- Mobile ee ane (Bartram) Sm: all —saw palmetto G. Nicholson} metto: Houston, Serenod serrulata (Michx.) Poaceae (Gramineae) Arundinaria gigantea (Walter) Muhl. ssp. gigantee ssp. fecta (Walter) McClure — switch cane [Arundinaria tecta (Walter) Muhl.} — Slant cane YOUNGHANCE AND FREEMAN, Trees and shrubs of Alabama SS) io) Oo REFERENCES AbDAMS, R.P. 1986. 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Native shrubs and ae vines of the Southeast, landscape uses and identification. Timber ie ee and, € Freeman, J.D., A.S. Causey, J.W. Suortr, and R.R. Haynes. oe Endangered, threat- ened, and special concern plants of Alabama. oe and Microbiology Dept. Ser. No. 3. Auburn Univ. Agr. Exp. Stat., Auburn, Goprrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. Univ. of Georgia Press, Athens, eee * Wooten. 1979. Aquatic and ee sae of the southeastern United ae 7 , Monocotyledons. Univ. of Georgia Press, Athens, id: ve uatic and wetland plants of the southeastern U ied States. Vol. 2, ee an of Georgia Press, Athens, GA. GONSOULIN, G.J. 1974. A revision of — (Styracaceae) in North America, Central America, and the Caribbea an, "Sida 5 5:L91— HANSEN, A.J., TA. Spies, FJ. Sw. ENON ane J.L. OuMANN. 1991. Conserving biodiversity in managed forests. Bioscience 4 Harbin, J.W. 1990. Variation patterns and recognition of varieties of Tilia americana 8.1. ©.92 sf,0Q Syst. Bot. 15:33— 384 Sipa 17(2) Hickok, L.G. and J.C. ANway. 1972. A morphological and chemical analysis of geographical variation in Ti/7a L. of eastern North America. Britt 24:2- HoitmGREN, P.K., N.H. Ho_mGren, and L.C. BARNETT. on hex Herbariorum, Ed. 8. Regnum Veg. 120:1-639. Howarb, R.A. 1974. Further comments on Styrax L. Sida 5:33: — 37. Isety, D. 1990. Naccdlee flora of the southeastern United States. Vol. 3, Part 2, Leguminosae iabaceae): Univ. of North Carolina Press, Chapel Hill, NC. Jones, S.B., Jk. and N.C. Coe. 1988. The discribution of the vascular flora of Georgia. Dept. of ae Univ. of Georgia, Athens, see bres . A synonymized checklist of ce vascular es of the United States, Canada, me ial 2nd ed. Timber Press, Portland, OR. Lirrie, E.L., Jr. 1979. Checklist of United States trees (native naturalized). U.S. Dept. Agr. Handb. No. 541. USDA Forest Service, Washington, DC. Luteyn, J.L., W.S. Jupp, S.P. VANDER Kioet, L.J. Dorr, G.D. Wattace, K.A. Kron, PE. STEVENS, and S.E. CLEMANTS, 1996. Ericaceae of the southeastern United States. Castanea 61:101—-144 Meyer, EG., P.M. Mazzeo, and D.H. Voss. 1993. A catalog of cultivated woody plants of the si hencn United States. U.S. Nat. Arbor. Contrib. No. 7. Nicoison, D.H. and G.C. StrrysKat. 1976. The masculine gender of the generic name Styrax [acne (Styracaceae). Taxon 25:581—587. PENNINGTON, T.D. 1991. The genera of Sapotaceae. Royal Botanic Gardens, Kew, England. PoweLL, D.S., J.L. FAULKNER, D.R. Darr, Z. ZHu, and D.W. MacCieery. 1993. Forest resources of the United States, 1992. General Technical Report RM-234 (Revised 1994). USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, CO. Prosst, J.R. and T.R. Crow. 1991. Integrating biological diversity and resource manage- enc. J. For. 89:12 fae. A.E., H.E. Anes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. Univ. North Carolina Press, Chapel Hill, NC. Reveal, J.L. and M.J. Serpin. 1976. On the identity of Halesia carolina L. (Styracaceae). Lxon 35:123-140 THORNE, R.F. 1992. Classification and geography of the flowering plants. Bot. Rev. (Lancaster) 58:225—348. si Kort, S.P. 1980. T Canad. J. Bor. 58:1187-12 Wirsur, R.L. 1994. The Mri aceae of the United States and Canada: genera, su and series. Sida 16:93-107. he ae of the highbush blueberry, Vaccinium corymbosum. ~— S — »genera, Worrorb, B.E. and R. aa 1993. Checklist of the vascular plants of Tennessee. Sida, Bot. Misc. No. 10 Woop, C.E., Jr. and R. B. CHANNELL. 1960. The genera of Ebenales in the southeastern Uniced States. J. Arnold Arbor. 41:1—35 ZOMLEFER, W.B. 1994, Guide to flowering plant families. Univ. of North Carolina Press Chapel Hill, NC. SUPPLEMENTAL NOTES ON RHYNCHOSPORA CRINIPES AND RELATED SPECIES IN SECTION FUSCAE (CYPERACEAE) ROBERT KRAL Herbarium, Department of Biology Vanderbilt University Nashville, TN 37235, U.S.A, ABSTRACT Rhynchospora crinipes Gale and related species are considered as to sectional disposition, taxonomic relationships, geography and ecology. New distributional information is added. RESUMEN Se consideran Rhynchospora crinipes Gale y las especies pr6ximas en cuanto a su disposicion seccional, relaciones taxonémicas, geografia y ecologia. Se afiade informacién nueva de su distribuci6n. INTRODUCTION Until recently, RAynchospora crinipes Gale has been one of the least known of native beakrushes, all knowledge of it being confined to Gale’s descrip- tion and illustrations (1944: 173, & pl. 823, figs. 2A, 2B) of two speci- mens on deposit at the U.S. National Herbarium. Somewhat later G. Kiikenthal (1950: 106) in his revision of the the “fuscae,” still with only these two specimens to go on, recognized the taxon but at a reduced rank, treating it as a variety of R. fi/ifolia. Both specimens were collected in June 1868 by Dr. Charles Mohr from the vicinity of Mobile, Alabama. Over the ensuing years many botanists, myself included, searched in vain to relocate Mohr’s distinctive plant, particularly since no one knowledgeable working in the genus doubted the morphologic distinctness of the taxon, whatever their judgement as to its appropriate nomenclatural rank. Reasons for difficulty in finding R. crinipes are plain enough. Mohr’s la- bel information is scanty and at the same time somewhat misleading. The holotype label reads “ditches, borders of ponds, June 1868,” that on the paratype “low but dry places, roadsides (exsiccated), June 1868.” This in- formation, while adequate to describe habitat for most RAynchospora known “ac ” from the Mobile area, provides no lead to the very specialized habitat of R. crinipes. It remained for Dr. Loran C. Anderson (1988) to do this in his excellent article, which is the first authentic report of the continued exist- ence of R. crinipes. In it he notes finding, during botanical expeditions by Sipa 17(2): 385-411. 1996 386 Sipa 17(2) canoe in the summers of 1987 and 1988, several viable populations from within the Big Coldwater drainage in Santa Rosa County, northwestern Florida. Big Coldwater Creek is one of the finer examples of a pristine blackwater stream in the Gulf South and he describes it very well. Since such streams are not rare in both the Atlantic and Gulf Coastal Plain, other botanists, armed for the first time on where precisely to look, have suc- ceeded in finding this plant over a five-state area. More information on such discoveries will be given under R. crinipes in the treatment below. This particular work is intended to bring recent fieldwork and studies on Rhynchospora crinipes into a taxonomic and ecologic context, in hope that it would serve as an aid to conservation of a rare species and its unique habitat. Thus, below, is a synoptical treatment of RAynchospora crinipes and related North American species in the Section Fuscae. TAXONOMY The two outstanding students of Rbynchospora in recent years remain Dr. Shirley Gale for her 1944 revision of “Eurhynchospora” (now treated as subgenus Rhynchospora) and Dr. Georg Kiikenthal for his revisional studies of the entire genus (1949, 1950). Of particular interest, I think, is the fact that neither had much if any field experience with RAyachospora, their re- markable effort coming instead from a careful study of herbarium material. Their taxonomy can scarcely be improved on by those who are accustomed to study these plants in the field. Their work is made even more useful simply because both were so consistent in their concepts of species and so willing to express doubt when doubt was justified. Of the two, the more conservative in assigning specific rank appears to have been Dr. Kiikenthal, perhaps because of his having to do the genus worldwide. This difference also shows up as to how complexes of species should be ranked. Gale (L.c.) broke what was then treated as Section Exrhyncho- spora of Subgenus Dysty/is into 12 “series” basing these largely on what previ- ous workers such as C.B. Clarke, J.K. Small, etc. had designated as sections, in some cases simply reducing them in rank, in others breaking a section into series, as was true in regard to Sect. Fuscae C.B. Clarke. Here, her de- cision seems to have merit since her Series Fuscae and Fasciculares do consti- tute two natural and distinct sections within the section Fuscae as con- ceived by Clarke, later by Kiikenthal. A key to the proposed sections is below: 1. Tubercle edges with at least some setae; fruits narrowed to a stipe-like vari- ously setose or setulose receptacle, the lenticular body under 1 mm wide and mostly broadest above the middle, the surfaces lustrous, ire with a pale, a ral disc Sect. Fuscae L. Tuber ae neu fruits not narrowed toa stipe-like base, the lenticu- lar a over | mm wide (except for R. ferna/dii) and broadest at the middle, the surfaces smoochish but dull ect. Fasciculares Kral, Rhynchospora crinipes (Cyperaceae) and related species 387 Since this particular report bears only on R. crinipes and its relatives in the proposed Section Fuscae | am deferring a consideration of the Fasciculares until a later date. The species accounts below are based upon the examina- tion of many hundreds of specimens (and in field work) but for economy only recent collections records will be cited, along with maps to show gen- eral distribution. Rhynchospora Vahl, Enum. Pl. 2:229. 1806, subgenus Rhynchospora Sect. Fuscae (C.B. Clarke) Kral, stat. nov. Rhynchospora Series B. Diplostyleae Sect. 4, Fuscae C.B. Clarke in oe Symb. Ant. 2:105. 1900 (in part); RAyachospora V. Claes Small, Man. SE Fl. 175. 1933 (in part); sy) ail Sect. Eurhynchospora Ser. 4. Fuscae (C.B. Clarke) ae Rhodora 46:169. 1 Solitary to cespitose, sometimes scaly-rhizomatous, perennials. Culms mostly erect to ascending-arching, subtrigonous, leafy, the leaves filiform to linear, blades flat to conduplicate or canaliculate, apically narrowed to triquetrous tips. Inflorescence of (1—)2—4(—7) mostly turbinate to hemispheric compounds of fascicles. Spikelets lanceoloid to narrowly ellip- soid, producing 2—4 or more fruits, the scales loosely spirally imbricate; receptacle often stipe-like, frequently setose; perianth of 6 ieee barbellate bristles, these always overtopping fruit body, often setose at or toward base; stamens 2 to 3; fruit body mostly obovoid to ellipsoid, bicon- vex, under 1 mm wide, broadest above middle, the faces smooth, usually glossy; tubercle triangular to triangular-subulate, flattened, its edges vari- ably setulose-ciliate. Type species: —Rhynchospora fusca (L.) W.T. Aiton. Six well-marked species, with R. fusca in eastern Canada and New England south in the Coastal Plain to Maryland, the other five primarily southward, mostly in the Atlantic and Gulf Coastal Plains with some ex- tending yet southward into the Antilles and/or Mexico and Mesoamerica. Below is a key to species, this followed by some notes on the individual species as I see them. KEY TO SPECIES OF SECT. FUSCAE 1. Spikelets (4.5-)5—7 mm lon 2. Fruit body broadly e llipsoid to obovoid; see blades filiform to linear. 3. Plants sparsely or densely cespit but also strongly clonal by slender rhizomes; stipe joint stout or ene leaf bl ades filiform. 4. cL few per clump; longer involucral bracts of terminal fascicle much exceeding it; fruit from stipe base to tubercle tip ca. 2.5 mm, the stipe smooth; fruit body a biconvex; tubercle ae wa setulose only at base; north .R. fusca y .Culms densely disposed; ee involucral bracts of terminal a. cicle barely if at all exceeding it; fruit from stipe base to tubercle tip m, the short stipe and tubercle edges setose and setulose respectively;fruit body more shallowly biconvex;southern........ 2. R. pleiantha 388 Stipa 17(2) 3. Plants cespitose or, in no. 4 stoloniferous, rooting from elongated bur- ied lower nodes when top} sled noe flood, then strongly clonal; stipe joint short to elongate, it and receptacle, smooth to variously setose or setulose; leaf blades filiform to ee Principal leaf blades no wider than 2 mm; fruit f am; fruit stipe joint smooth to sparsely setose, . | mm wide; spikelet clusters s1tOse 3. R. harperi rom stipe base to tubercle tip to 2.7 to 0.4 mm; fruit body obovoid, ca few, mostly Eo tubercle triangular su ibulace; plants strictly Ges even when toppled 5. Pri eas al leaf blades 2-5 mm wide; fruit from stipe base to tubercle 2.8—3.3 mm; fruit stipe joint slightly to densely setose, 0.5—1 mm; fruit body narrowly conta ca. 0.8 mm wide; spikelet clusters several, usually 4 or more; tubercle triangular, or sides slightly con- ; plants cespitose but clonal ae toppled by Hood, then root- Cave . R. crinipes ing at buried lower nodes and stoloniferous . Fruit body narrowly oblong-ellipsoid; leaves filiform ......000...00cccee. oO. R. filifolia by ee 3-4 mm long 1. Rhynchospora fusca (L.) W.T. ee Hort. Kew. ed. 2,1:127. 1810. 763 Schoenus fuscus L., Sp. Pl. ed. 2, 2:1664 Phaeocephalum fuscum House, Amer. Midl. Naturalist. 6:202. 1920. Rarely solitary, mostly sparsely cespitose perennial to 4(—5) dm high, clonal by slender (1—2 mm) imbricate-scaly rhizomes. Culms wiry, ascend- ing to excurved, subterete, multicostate, at midculm 0.5—1 mm thick, leafiest at and toward base. Leaves erect or bowed outward, the principal ones mostly 0.5—2 dm long, the sheath with broad, scarious red-mottled border, at apex with a convex “V” shaped low scarious ligule; blades nar- rowly linear to filiform, at base to 1.5(-2) mm wide, narrowly involute, upblade gradually narrowing to broadly, then narrowly canaliculate, taper- ing toa sulcate triangular apex, the tip typically blunt, the margins smooth. Fascicles of spikelets mostly compound, appearing to be 1—3(—4), all ex- ceeded by one or more subtending, setaceous-tipped bracts, cylindric or (mostly) narrowly to broadly turbinate, rarely hemispheric, 1-1.5 x 0.5— 1.5 cm, sparse to dense, the terminal complex of up to 3 or even more approximate fascicles, the laterals progressively more distant longer-pe- duncled, longer-bracted, downculm; spikelets broadly to narrowly ellip- soid-lanceoloid, 5-7 mm, narrowly acute, pale to deep red-brown, scales several, the lowest barren, often cuspidate-aristate; fertile scales, mostl 3(-4), narrowly ovate to oblong-lanceolate, 4—5(—G6) mm, acute, convex to navicular, the strong midcosta excurrent as mucro or cusp; stipe and recep- acle stubby, 0.2—0.3 mm, usually smooth; periancth mostly of 6, antrorsely barbellate bristles to 2.5 mm, the longer extending past base of tubercle, mostly past its tip; stamens 2—3, anthers 2—2.5 mm long; fruit body tumidly lenticular, obovoid,1—1.3 mm, narrowly and inconspicuously margined, at apex with a shallow trough below the sharply raised tubercle base, faces KRaL, Rhynchospora crinipes (Cyperaceae) and related species 389 pale to deep brown, lustrous, longitudinally finely lined, transversely very finely rugulose with wavy rows of very narrow, vertical cancellae, some- times also with lines of shallow pits; tubercle strongly subulate, (0.7—)1— 1.2(-1.5) mm, the edges scabrid-setulose only toward base. Figure 1. Wet to moist sandy (often sphagnous) peats of bogs, pondshores, seeps, acidic wetlands, northeastern North America, in Canada from the maritimes westward to Lake Superior region, in the U.S. from New England west- ward from New York to northern Illinois, Wisconsin and Minnesota, south- ward in the Coasal Plain to New Jersey and Maryland. Europe. Relationships of this widespread sedge are plainly closest to R. pleiantha and will be taken up there. No specimen citations are given here. 2. Rhynchospora pleiantha (Kiik.) Gale, Rhodora 46:171. 1944. R. ce var. pleiantha Kiik. Feddes Repert. se Nov. Regni Veg. 23:208. 1926. Type: C PINAR DEL Rio: Laguna Sta. Maria, 22 Aug 1923, Ekman 17242a penis < ISOTYPE: L Densely cespitose perennial (1.7—)2—4.5(—5.3) dm high, strongly clonal by slender (to 2 mm thick) scaly rhizomes. Culms wiry, ascending to excurved, obtusely angled or subterete, multicostate, at midculm mostly 0.4—0.7 mm thick, leafiest at and toward base. Leaves erect to bowed out- ward, principal ones 1—2 dm long, the blades filiform, proximally shal- lowly to deeply canaliculate, upblade becoming convolute or conduplicate, then triquetrous to a short, blunt, scabrid-edged tip. Fascicles of spikelets 1—2(—3), mostly narrowly to broadly turbinate, rarely hemispheric, each with few to many pale to deep red-brown, lanceoloid spikelets, the termi- nal fascicle largest, sometimes lobed, equalled or slightly exceeded by a stiff, erect, linear-setaceous bract. Spikelets 5-7 mm long, acute, scales several, the 2—5 fertile ones narrowly ovate to broadly oblong-lanceolate, 3.5-)4-5 mm, acute, convex to navicular, only the midcosta evident, this excurrent as an aristula; stipe and its receptacle short, rarely to 0.3 mm, from SerunOse to nearly smooth; perianth of 6 antrorsely barbellate bristles ca. 2-3 mm, extending to tubercle tip or beyond; stamens 2(—3), anthers iene: ca. 2mm; fruit body biconvex, narrowly to broadly obovoid, 0.8-1 mm, margined and apically broadly but shallowly sulcate below raised edge of style base, usually lustrous brown, often with a paler oval “disc” at cen- ter, the surface very finely longitudinally lined, sometimes finely reticu- late-cancellate; mature tubercle triangular-subulate, 0.7—0.9 mm setulose only toward slightly decurrent base. Figure 2. Wet fine sands or sandy peats, mostly around karst ponds or in their environs, Atlantic and Gulf Coastal Plain, North Carolina south to penin- sular Florida, western Florida and contiguous southern Alabama. Cuba. Figure 3. 390 Sipa 17(2) ee FIG _ 1. Rhynchospora fusca (Kral 78328). A. Habit sketch. B. Leaf sheath-blade junction, adaxial view (left), abaxial view (right). C. Sectors of leaf blade from discal (bottom) to apical (top). D. Apical compound fascicle. E. Spikelet. F. Fertile scales, lower (left), upper (right). G. Anther and upper filament. H. Mature fruit & perianth, edge view (perianth removed) at left, face view (right). Krat, Rhynchospora crinipes (Cyperaceae) and related species 391 Fic. 2. Rhynchospora pletantha (Kral 81119), A. Habit sketch. B. Leaf sheath-blade junc- tion, abaxial view (left), adaxial view (right). C. Series of sections of leaf blade from near base (bottom) to tip (top). D. Apical fascicle. E. Spikelet. F Spikelet scales, lowest sterile (left), lower fertile (middle), upper fertile (right). G. Receptacle and stipe with 3 bristles. H. Top of one stamen. I. Mature fruit with stipe and perianth (left), side view of fruit body (right). 392 SipA 17(2) { § Rhynchospora “<2 pleiantha iam, Rhynchospora barperi Rhynchospora crinipes Rhynchospora curtissit Fic. 3. Distribution of RAynchospora pleiantha, harperi, crinipes, and curtissii in the southeast- ern United States. Gale (L.c., p. 172), as mentioned earlier here, appears to have it right in determining that R. pleiantha is a close relative of the more northern R. fusca. As she mentioned in the synonymy, Dr. R. Harper, a very astute observer and collector of sedges, considered the two conspecific. Both are low, cespitose and with wiry culms and leaves of similar dimensions; both have few fascicles per culm, these generally rather closely spaced at and near culm apex, these similarly narrowly to broadly turbinate and of simi- lar color; the fruits, including receptacle-stipe, are similar in shape, color Krat, Rhynchospora crinipes (Cyperaceae) and related species 595 and most surface features. Interestingly, Gales description of R. plezantha excludes the slender scaly rhizomes abundantly produced by this species and which are likewise produced by R. fusca, a feature which ties the two even more closely together. Other differences mentioned by Gale are how- ever consistent and enough to “make” species, as in the short stipe-recep- tacle and the perianth bases of R. fusca being smooth or nearly so, the more tumid fruits consistently, if slightly, longer and broader. Also, the seta- ceous bract subtending the terminal fascicle much exceeds it in R. fusca, whilst in R. plezantha is ranges from slightly shorter to slightly longer than that fascicle. But a consistent taxonomy would, if the two were sympatric at all, be to make them geographic variants. In habitat R. pletantha is remarkably uniform over its range. It can form bands along the shores of karst ponds and lakes, these fluctuating borders providing a substrate of fine sand and peat. While many more specimens from herbaria were studied from loans, I am citing only those here at VDB, several of these representing numerous duplicates of my numbers, now discsibuced or soon to be (here and subse- quently indicated by “t.b.d.”): Specimens examined: U.S.A. ALABAMA. Covington Co.: sandy upper shores of Blue Pond, Blue Springs Wildlife Mgme. Area, SW Andalusia, 1 Jun 1970, Kra/ 39499 (VDB & are distributed). Houston Co.: sandy peaty shores of limesink pond by co. 4, ca. 4 W Chattahoochee State Park entrance, Kral 47261 (VDB & widely distributed); sandy i at edge of Indigo Pond, SE of Cottonwood, 20 Aug. 1977, R. Kral 60845 (VDB & widely distributed); sandy peaty edge of doline by co. 8, W of entrance to Chattahoochee State Park, 20 Aug. 1993, R. Kral 83230 (VDB, set to be distributed). FLORIDA. Bay just to § side of FLA 20, 4.4 mi E its jet. with FLA 77, generally N of Vicksburg, 14 Aoet O05 Pei: ee with A. Gholson 8: faa (FSU, GH, VDB); sandy peat of shores of small doline S side FL 20, N of Bennett, cz i W of Iconfina River, 18 Jul 1992, R. Kral 81199 (BM, BRCH, CM, CTB, FSU, GH, K. KANU, M, MICH, MO, MSC, NCU, NY, PH, SMU, VDB). Calhoun Co.: sandy peaty exposed shores and bottom of cypress-myrtle- leaved holly pond, U.S. Rt. 90, 3 mi E of Bay Co. line, W of Clarksville, 10 Aug. 1976, R.K. Godfrey 75328 (FSU, VDB, etc.). Hernando Co.: 3 mi N of Weeki Wachee Springs; abundant on moist to wet sands at margin of hypericum pond, 9 Jun 1958, R. & M. Kral 6786 (FLAS, FSU, GH, USE, VDB, etc.). Jackson Co.: sandy peaty shores of Race Pond, P , R8W, W of Apalachee Wildlife Mgmt. Area on Lake Seminole, R. Kral with A. Gholson 60789 (VDB). Leon Co.: sandy soil, upper exposed margins of pond, sinkhole pond amidst sand ridges near Springhill Rd., ca. 1/4 mi S of Capital Circle, S$ of Tallahas- see, 12 Jun 1976, R.K. Godfrey 75062 (FSU, VDB); ca. 4 mi S of Tallahassee on sandy peat of upper margin of limestone pond in longleaf pine-turkey oak forest; plants scoloniferous 15 Aug 1962, R. Kral & R.K. Godfrey 15581 (FSU, VDB), ae a shores of Dog Lake, SW of Tall ge in Apalachicola National Forest, 3 Jul 1 R. Kral 82783 (BRCH, MO, VDB). Wakulla Co.: damp sandy ae of a lake oes ee and St. Marks, anno ca 1843, 7 ae 601 (BM, FLAS, B). Walton Co.: ca 2 mi E of county line by US 98; sandy border of small pond within nih of Gulf, 12 Jul 1963, R. Kral 17750 (VDB), seed peat at edge of Lake Chipley SW of S arm Juniper Lake and ca 1 mi E of US 331, side De Funiak Springs, 23 Jul 1993, R. Kral 82930 with G. Moore (BRCH, CM, 394 SIDA 17(2) CTB, GA, GH, IBE, MICH, MO, NCU, NY, OSC, PH, VDB). Washington Co.: sandy shores of Chain Lakes, W of eee. 6 Jul 1963, R.K. Godfrey & N.C. Henderson 62955 (FSU, VDB); meee eat pains pond, aces Pine Log State ae ca 0.5 mi W of FL 79, 15 Aug . Kral aon (BR — , CTB< GH, MO, VDB); sandy peat of shore of S-acre one ine ca oF coi N of FL 20, ca < es mi E of jet. FL 77, : Aug 1992, R. Kral 81541 (FSU, oad B). NORTH CAROLINA. Brunswick Co.: sandy peat at edge of Blue Pond, ca 4.2 mi o Jackies Creek and NC 133, ca 500 yds NW from McKenzie Pond, | Jun 1980, R. Kral 65265 (VDB). 3. Rhynchospora harperi Small, Fl. SE U.S. 182. 1903. R. fascicularis (Michx.) Vahl var. harper? (Small) Kiik., Bot. Jahrb. Syst. 75:111. 1950. Type: U.S.A. GEORGIA. Putaski Co.: wet pine barrens ca 3 mi E of Hawkinsville, R.M. Harper 1337 (HoLoryPE: NY; tsorype: US). R. leptorhyncha sensu Small, Fl. SE U.S. 195. 1903, non C. Wright, 1871. Cespitose perennial (3—)5—8(—9) dm high. Culms erect to arching, ob- tusely angled to subterete, rarely filiform, more often distinctly broader than leaf blades,at ca midculm (0.5—)1—2 mm thick, strongly costate, leafiest at and toward base. Leaves erect to ascending, mostly straight, principal ones 1—2 dm long, blades linear-filiform, 0.5—1.0 mm wide, proximally slightly inrolled (broadly canaliculate), upblade increasingly narrowly condupli- cate with midcosta strongly raised abaxially, thence gradually narrowing, adaxially sulcate, to a triangular or subtriquetrous apex, the very tip slightly flattened, blunt, scabrid-margined. Fascicles (1-)2—3, turbinate to hemi- spheric, 0.7—1.5 x 1-2 cm, each with few-to-many neler pale to deep red-brown spikelets, the terminal fascicle largest, densest, often lobed or with another immediately subtending, usually atop a distinctly bent or excurved internode, equalled or exceeded by one or more subtending lin- ear-setaceous bracts; spikelets S—7 mm, narrowly acute; scales several, the —4 fertile ones narrowly ovate to broadly oblong-lanceolate, (2.5—)4—5 mm, acute, convex to navicular, only the midcosta evident, this distally scabrid, excurrent as a cusp or aristula; stipe and its receptacle short, 0.2— 0.3 mm, usually sparsely setose and setulose; perianth of 6 antrorsely barbellate bristles 1.5-2.5 mm, extending from mid-tubercle to beyond tubercle tip; stamens 2(—3), anthers linear, ca. 2 mm; fruit body biconvex, — broadly obovoid, 1—1.4 mm, margined and apically broadly grooved below the narrow, sharply raised tubercle collar, faces glossy, pale to dark brown, often with a paler oval disc at center, finely longitudinally lined, variably low papillate-cancellate, also often transversely with wavy lines of dark punctae; mature tubercle triangular-subulate, (0.8—)0.9—1(—1.1) mm, edges densely pale setulose-scabrid. Figure 4 Moist to wet sands and peats of pond shallows and shores, Atlantic and Gulf Coastal Plain, from Maryland and Delaware, then North Carolina south to southern Florida, westward to panhandle Florida and southern Alabama. Figure 3. Krat, Rhynchospora crinipes (Cyperaceae) and related species 395 —— E a Fic. 4. Rhynchospora harperi (Kral 85337). A. Habit sketch. B. Leaf sheath-blade junction, abaxial side (below), adaxial side (above). C. Series of sectors of leaf blade from distal (below) to distal (top). D. Leaf apex. E. Terminal fascicle. F. Spikelet. G. Fertile brace. I. Face view of mature fruit. 396 Sipa 17(2) Richard LeBlond has reported records for this species, until then thought confinec eon to Florida, Georgia and Alabama so that we now know it from several additional states. My own experience with it thus far is within its area of greatest abundance, namely panhandle Florida, where optimum habitat is pond shallows in Taxodinm ascendens-Nyssa (biflova, ursina)-Llex myrtifolia-Cyrilla in the pine-saw palmetto flats, where it is mostly ad- mixed with various species of shrubby myriandrous Hypericum, or along ditches or shallow moats bordering pine savanna bogs. It should be noted that the Delaware and Maryland R. harper is a far cry from that of North Carolina southward, being lower, narrower-leaved, with smaller fascicles, the terminal internode mostly lacking the “bend,” the fruits on the short side of the range. However, this material, while strongly resembling R. filifolia, would be difficult to place there or elsewhere except in R. harperi. Specimens examined: As is the case with R. plesantha (above), | am citing only those specimens here at VDB: U.S.A. DELAWARE. Sussex Co.: Assawoman ee 17 Sep 1991, RJ. LeBlond 2552 (VDB). FLORIDA. Bay /Washington Co.: sandy peat of sloughs in pine-cypress savanna, FL 20 by cty marker, E of Ebro, 9 M: ay 1995, oy, 84643 (FSU, GH, MO, VDB). Franklin Co.: deep-banked drainage ditch i with ipsa fascicnlatum 2 mi-. E of rte 65 S side Buck Siding Rd, SEI/4 of NW1/4 Sec.9, T7S,R7W, L- Jul 1988, L.¢ poe 1611 (FSU, VDB); se: Dy water of cypress stringer, S side forest e 148, etween open savannas, 0.4 mi W of rte 65, 6.5 mi S of Sumatra, 23 May oak ». Anderson 12004 (FSU, VDB); wet peaty sand, a cypress in Tate’ s Hell Swe amp, mi - of High Bluff Creek, ca 7.5 mi NE Easepoint, 28 May 1991, L.C. Anderson 13429 me VDB); abundant over large area of ee f pond-cypress forest, Tate’s Hell Swamp, E of FL 6, R. K. Godfrey 8 3880 on W. Baker (FSU, GH, VDB); sandy peats of cucover cypress-gum-pine flats by FL 65, 2.2 mi N of jet. US 98, 3 Jul 1993, R. Kral 82800 (BM, BRCH, CLEMS, CM, CTB, FSU, GA, GH, K, KANU, MBM, MICH, MO, MSC, NCU, NY, TENN, TEX, US, USCH, VDB, VSC, WAT, WILLI, WIS); sandy peat of edge of small cypress dome in scrub, end of ay lone East Point, ENE end of FL 65, 20 Jul 1993, R. Kral 828406 with G. Moore (FSU, GH, K, KANU, MICH, MO, NCU, NY, TEX, USCH, VDB, WIS). Gulf Co.: bog eee - a ae chapmanti in clearing of thick Cyri/la scrub beew. Daniels Rd. and Wetappo Creek, 4.3 mi S of rte 22, ca 5.5 mi SW Wewahitchka, 20 Jul 1989, L.C. Anderson 12170 (FSU, ae. sandy peaty swale in Hypericum-Cyrilla- Nyssa-pine, flats by Fl 71, 6-7 mi S of Wewahitchka, R. Kral 82854 (GH, MICH, MO, NY, USCH, VDB); sandy an of ditch through slash pine flats, S side US 98, 5 mi W of Franklin Co. line, 21 Jul 1993, R. Kral 82868 (BRCH, CM, GA, GH, K, MICH, MO, CU, TEX, <6 VDB): and same locale but with dwarf cypress, R. Kra 82869 (VDB). Then Co.: common in shallow water of a Hens along railroad border- ing slashpine flats : of rte 65 a few mi NE Wilma, 10 Jul 1 , L.C. Anderson 13706 (FSU, VDB); by FI. 65 plus or minus 5 mi N of Sumatra, — pit, aspect dominant Hypericum wines P. Adams, 26 Jul 1993, R.K. Godfrey 84647 (FSU, GH, VDB + 43 duplicates elsewhere); sandy ae oF shallow ee a ey w side FL 65, 22 mi S of Telogia, oe ant, 3 Jul 199 . Kral 82809 (CLEMS, KANU, MO, NCU, NY, TEX, US VDB, VSC, ed "Martin Co.: : ash pine . at $ side Stuart off US 1 pond shi ee 18 Sep 1973 3, R. Kral 51780 (VDB & distributed); ae peat of shi ire sa Taxodium pond by Willoughby Ave., ca. 1/4 mi N of j 22, scarce, 8 Jul 1994, R. Kral 83706 (BRCH, FSU, MO, NCU, VDB, NY). Okaloosa Co.: small gum a + Krat, Rhynchospora crinipes (Cyperaceae) and related species 397 pond in longleaf pine savanna near gravel rd intersect | mi N of Beaver Creek, ca. 3.5 mi of Fl 4 ca 7 mi E of Munson; Danley Ponds, Blackwater State Forest, 10 Jul 1990, S.L. Orzell & E. L. Bridges 14275 (TEX, VDB). Volusia Co.: upper transitional zone from open eae flatwoods pond to clearcut slash pine flatwoods, ca 4.7 mi W jet. Fl 40 and US 7 in Barberville, 12 Jun 1990, S.L. Orzed/ & E. - cee 13 3921 (TEX, VDB). Walton c o.: sandy peat of aa cypress- car savanna, FL 2 E of Bruce, frequent, 18 Jul 1995, R. Kral 85337 (BM, BRCH pro Coa. aa - KANU, MBM, MICH, MO, MSC, NCU, NY, US, VDB). Whingion Co.: sandy peaty Nyssa-Taxodium ascendens- Hypericum flats ca. 1/2-1 mi W of Bay Co. line marker by FL 20 just E of Ebro, 19 Jul 1993, . ee 820 (CLEMS, CTB GH , MICH, MO, MSC, NCU, NY, TEX, US, USCH, WAT); Fla 20, 1/8-1/4 mi W of Bay or es in cypress-pine savanna, sandy peat, R. Po 85322 (NY, US, VDB); & t.b.d.). RTH CAROLINA. Onslow Co.: Camp Lejeune Marine Corps Base, wet peat of gr ee forb es meadow, 2 Aug 1990, RJ. LeBlond 1549 (VDB); Camp Lejeune Marine Corps , ina 3-acre shallow pond § of the jct. of Sneads Ferry and Marines Rds.,6 Aug. mn RJ. LeBlond 1552. (VDB) 4. Rhynchospora crinipes Gale, Rhodora oe 173. 1944. R. flifelia A. Gray var. crinipes (Gale) Kiik., Bot. Jahrb. 75(1):106. 1950. Type: U.S.A LABAMA. Mosite Co.: ditches, borders of ponds, Mobile 18 Jun 1868, Charles aa (HO- LoryPe: US!); dry places, roadsides (exsiccated), Mobile, Jun 1868, Charles Mobr s.n. ries nen US!). — Solitary or cespitose perennial 6—10(—15) cm long, often strongly clonal through toppling by flood, lower internodes thus elongating, rooting at nodes stoloniferously. Culms lax, erect to ascending, excurved distally, at midculm mostly 2—3 mm thick, obscurely trigonous to terete, multicostate, leafy. Leaves lax, the lower ones often trailing after flood, ae ones toward and at culm base, 20-50 cm long; blades narrowly linear, 2 i>) mm wide, proximally flattened, at base slightly concave, midcosta slightly raised abaxially, thence upblade more level with midcosta strongly raised abaxially, chence to “v” shaped, with midcosta a sulcus adaxially, very raised abaxially, thence tapering and triquetrous to a subulate tip. Fascicles of spikelets mostly compound, (3-)4—7(-10), mostly pee to pia! tur- binate, less often hemispheric, ellipsoid or ovoid, 1.2—2 * 1—2.5 cm, all short-pedunculate, the lower ones distant, and ee exceeded subtend- ing foliaceous bract, upculm progressively more approximate and shorter- bracted, the apical ones exceeding or but slightly overtopped by linear- subulate bract(s); spikelets lanceoloid, 5—5.5 mm, acute to acuminate, light red-brown (on flooding darkening to very dark brown, appearing nearly black); fertile scales narrowly ovate to lanceolate, 4—4.5 mm, convex with the poe midnerve excurrent as aristula or mucro; stipe and receptacle (.5—0.8(—1.0) mm sparsely to densely setose; perianth of 6 ascending, se- tose-based, antrorsely scabrid bristles 2-2.5 mm reaching above tubercle base, more often to or beyond its tip; stamens typically 2, anthers linear, ca 2 mm; fruits 2—4 per spikelet, the body lenticular, pa le-margined, nar- 398 Sipa 17(2) rowly to broadly obovoid, 1.2—1.5 mm, faces apically concave below sharply raised tubercle base, brown to even castaneous, lustrous, often with a pale glassy midzone, the surface minutely striolate, sometimes transversely mi- nutely rugulose and with wavy rows of dark puncticulae; tubercle triangu- lar, sometimes slightly subulate (concave-edged), 0.7—1.1 mm, setulose- ciliate from base to tip, the slightly lunate base flowing to the strong fruit margin. Figure 5. Banks, bars, shoals of blackwater streams, less often ditchbanks and slug- gish drainage from flatwoods bogs and ponds, Atlantic and Gulf Coastal Plain, North Carolina; Georgia south to northwestern Florida, southern Alabama, southeastern Mississippi. Figure 3. Rhynchospora crinipes is unquestionably the most distinct species of its subcomplex in Sect. Fuseae. In habit it is callest, has the thickest culms, the broadest leaf blades. While the plants are typically cespitose, in common with the other species, they are unique in their reaction to toppling, in that culm bases when buried by wash produce elongated lower internodes anc increase stoloniferously, thus rooting at one or several lower nodes. The plant bases therefore are often more deep-set, stubbornly hanging on through flood cycles, forming extensive clones in the “best” habitat when other species of KAynchospora may wash away. In fact, dislodged plants may for the same reason reroot more readily in similar habitat downstream. So, while rootstocks of the species have been described as rhizomatous, they are truly stoloniferous, very unlike the true rhizomes produced in R. fusca, R. pletantha of the subcomplex which are narrower with spirally imbricate scales. Ic differs from the others in numbers of fascicle compounds, rarely as few as 3 per culm, (generally the maximum number for the others) more often 5 or more per culm. There is overlap in fascicle and bract shape and dimension, as well as in shape and size of spikelet and number of fruit per spikelet, but none in length and indument of stipe and receptacle, this being the longest and shaggiest of the complex. Actually, in the field, a common companion RAyxchospora from another complex, namely R. glomerata (L.) Vahl bears the strongest resemblance in general dimensions of culm, leaf, and inflorescence (but not clonal by stolon), differing mainly in char- acter of flower and fruit. Within the complex only R. filifolia may be abun- dant in such habitat but is easily distinguished on a basis of habit, being lower, with narrower leaves and culms, but non-clonal by stolon, with fewer and smaller spikelet clusters, smaller spikelets. Differences between R. crinipes and R. filifolia are even stronger when one consults the flower and fruit, the former with a longer, hairier stipe- receptacle, a similar shaped fruit body, but longer and not as smooth, a tubercle that is longer, often less triangular (straight-sided). Thanks to support from Botanists Cary Norquist (U.S. Dept. of Interior, en epee, Kra., Rhynchospora crinipes (Cyperaceae) and related species \Nt i Wy , ay WW ft GULL 0 Xd Sy wy ‘ \ hy NEE f Vie, if % i \ | Fic. 5. ine fades ee 85409). A. Habit sketch. B. Leaf apex. C. Sector of leaf tor of leaf midblade, abaxial view. E. Leaf sheath-blade midblade, adaxia junction, adaxial view. F. poe fascicle. G. Spikelet. H. Face view of mature fruit. J. Fruit with sprouting seed. 400 Sipa 17(2) Fish & Game, Jackson Office) and Jeryl L. Hilton (Alabama Natural Heri- tage Program, the Nature Conservancy office, Montgomery) I have been able to do a proper field survey of this RAynchospora over what is presently known to be its geographic range. My notes below on habitat are based on personal encounters from 33 sites, as well as from consultation of vouchers through loans or sendings of duplicates from knowledgeable colleagues such as Bruce Sorrie (North Carolina) and Loran Anderson (Florida). The survey has resulted in re-establishment of it in Alabama, establishment of Georgia records, and extension of its known range to Mississippi. While efforts to locate it in South Carolina and eastern Louisiana have thus far not been successful, such should be continued since there is plenty of suitable habitat there. Thus, “suitable” habitat is discussed below. The geographic distribution of R. cr7zipes as presently understood is rather remarkable. A general realization of it can come from a map of the south- ern Atlantic and Gulf Coastal Plains, on which Chamaecyparis thyoides (L.) B.S.P. (including C. benryae) is shown (Little, E.L., Atlas of United States Trees, Vol. 1, Map 11-E. 1971) since most of cng optimal sites are in, or are upstream or downstream from Chamaecyparis habitat. In my experience, R crimipes 1s never far from clean, clear, active streams or in seeps directly by them. Such streams, generally a pale wine in color eas tinting by acidic- humic seepage are mostly swift and well aerated. They run clear soon and recede quickly after rain when the watershed is healthy. They have distinc- tive and stable woody and herbaceous bank floras. They have clean, sandy or gravelly sandy bottoms and bars, these often underlain by lenses of gray clay with iron mottling which often are exposed in the swifter runs. They meander, thus cut deeply into steep slopes and deposit large bars of sands and gravels opposite cuts, these bars often of white sands freckled with rounded stones and pebbles of quartz, hardened clay, chert. On the steep sides large trees are often undercut by floodwater, and topple into the stream often to create considerable “jams” and “dams” with their attendant accu- mulations of floodwash. In more level stretches, particularly as the valley broadens, sandy-gravelly-silty floodplain develop, these optimum sites for some “high” forest of Chamaecyparis, Magnolia virginian, Liguidambar, Fraxinus, Acer rubrum (sometimes Acer saccharum), various “willow” oak, Lirtodendron, etc. with Cyrilla, Cliftonia (southern part of range), various Myrica, Ilex, Vaccinium, thickets of Arandinaria, etc. in the understory. As one moves downstream and gradient is less, as floodplains broaden and channels deepen, the amount of good habitat for the rhynchospora becomes less. Within the range of a stream of this sort, the bese clones of R. crinipes are in full sun or light shade along the narrow banks below cuts, on tails of bars, in seepage areas in cuts and broader banks, or on low, frequently tem- porarily washed islands. Most of my finds have been around bridgesites, — Krat, Rhynchospora crinipes (Cyperaceae) and related species 401 these of course cleared back to limit of road right-of-way to what is usually a dense vegetation under which the rhynchospora becomes quickly more scarce. Wading up and downstream from a bridge gives one a perspective on how the species maintains under natural conditions. In broader streams this sun plant has no problem, but along narrower stretches where a channel] is N-S-oriented there is too much shade. In such situations the rhynchospora is usually found only where overstory trees have toppled to create open- ings. In these, over time, clumps of culms are toppled by flood, their bases buried by floodwash, their lower internodes lengthening, their lower nodes rooting and extensive clones thus form. Heavy flood washing may dislodge parts of clones, some re-lodging downstream. The phenology is broad since this species flowers and fruits as early as May and will continue to do so until frost; therefore over a long season an abundant supply of fruits is swept downstream. Most fragments and fruits are doubtless lost, others may survive for short periods in shaded stretches, but in such active streams many openings are formed or reformed and are ideal. I encountered a per- haps anomalous but yet very interesting situation when sampling a large clone on Escambia Creek, Escambia County, Alabama, northeast of Atmore (Kral 85234), where many of the toppled culms had many spikelets in which the fruits had germinated, some actually beginning to take root where fascicles were flat against a patina of slick sandy peat. This again shows a possibility for a remarkable adaptation. Several other Rhynchospora do occur in and immediately around popula- tions of R. crinipes, some aleeadly noted, such as R. filifolia, R. glomerata, but also R. cephalantha, R. chalarocephala, R. corniculata, R. gracilenta, R. macrostachya (in the northern range), R. microcephala, R. nitens, R. rariflora. It may, in shaded bottoms, sometimes also be associated with such species of shaded bottoms as R. caduca, R. decurrens, R. miliacea, or R. mixta of distinctively different section. But none of these associates have the combi- nation of features that allow as much adaptation to “temperamental” and dynamic blackwater stream habitat that this tenacious and deeper-rooted species has. There are some other sedges and monocots that can deal well with such sites, most notably Eleocharis olivacea, Scirpus etuberculatus, S. subterminalis, Orontium, Sparganium, various Juncus, particularly J. vobicepbalas: the sedges and Sparganium often streaming in massive clones along bars and islands and in shoals. But none of these is better adapted. I suggest that R. crinipes once had a wider distribution along some of the larger streams, since a wider channel would also admit more light, particu- larly if this was a braided system with many shoals and bars. But, since larger stream systems also have (or had) broader displays of quality trees along them, and since logging of such systems has been generally disas- 402 Sipa 17(2) trous, consequent extremes of flooding and receding, accompanied by in- creased siltation and accelerated erosion have washed out or buried these sorts of plants. In short, the continuance of R. cr7nipes very much depends on careful management of adjacent forest and watershed. An overview of watersheds themselves would show the best ones to be in longleaf pine-deciduous scrub oak, “yellow” sandhills or where, toward the coast (as in Florida, Alabama) streams issue from or flow through Sand pine-evergreen scrub. These systems are fed by seepage or springs, some- times karst type. The pine-oak-hickory uplands grade downslope to pro- gressively more mesic systems of pines, oaks, hickories and other hard- on woods such as Liriodendron, Liquidambar, maple, downward to a Fagus-Magnolia (grandifolia)-Acer saccharum (southern subspecies) type, to the actual streambottoms where are higher silt fractions and Chamaecyparis- Taxodinm-Virginia bay-Red maple-willow oaks-Persea-Hex-Liquidambar etc. hold forth. It is significant that there are often cutoff meanders, but while these may have abundant sedges along them, R. crinipes does not persist long in such places; it has to be part of an actively combed and well aerated stream. The only areas where there seems to be some transition of type is in the Florida Panhandle where some small and comparatively slack streams are flowing into or out of cypress ponds or bogs in a pine flatwoods system (localities in Bay, Gulf, Liberty counties). In these places, particularly along “ditched” flow, R crinipes can be very close to habitat of R. harpert, R. curtissit. Vouchers from the 33 sites I have inspected as of 1995, cogether some sent by Dr. L. Anderson, Dr. M. LeLong, and Mr. B. Sorrie are cited below. Since most of the sites I] have sampled had few plants, most of my collec- tions are represented by but one or two specimens and few large sets were therefore made: Specimens examined: U.S.A. ALABAMA. Baldwin Co.: N side Bay Minette Creek directly across from Buzbee Landing, at waters edge, 23 Jul 1993, Howard Horne, s.n. (USAM, VDB); peat-muck slicks and in shallows of branch of Perdido River by county 61 bridge, ca 10 mi E of Bay Se 26 Jul 1990, R. Kral 78170, first record re- pee species for Alabama! (BM, BRCH, CM, CTB, FSU, GH, K, MICH, MO, MSC, NCU, TEX, USCH, VDB); same site, 21 Aug 1993, R. Kral 83269 (VDB , VSC); grave Hom along Major's Creek off cty 96, W of Rabun, 18 Aug. 1994, R. Kral 84088A (MO, MSC, VDB). Conecuh Co.: gravelly-sandy-silty bank ping creek by gravel road 5.4 mi W of Range, 18 Sep 1993, R. Kral 83325 (MO, NCU, VDB, WIS); recollected from seep bank, same site, 6 Jun, 1995, Kral 84890 (NY, US, VDB, ae Escambia Co.: sandy-gravelly seeps and banks along Escambia Creek near I-65 crossing SW of Barnett Crossroads, 21 Aug 1993, R. Kral 83267 (GA, MO, NCU, NY, US, VDB); Co. 27 crossing of Escambia Creek ca 10 mi NE of Atmore; sandy gravelly | banks along stream, 18 Sep 1993, R. Kral 83329 (VDB, ae and downstream 13 Jul ie (fruit germinating in spikelets on toppled culms!), Kra/ 85234 (CLEMS, CTB, GH, K, MICH, MO, MSC, NY, TENN, TEX, US, USCH, VDB, ca VSC, WAT, WILLI); gravelly bar in Juniper Creek by Al 49, ca 4.9 mi N of Brewton, scarce (the “Juniper” is Chamaecyparis), 18 Aug 1994, R. —— Krat, Rhynchospora crinipes (Cyperaceae) and related species 403 Kral 84070 (VDB); and abundant along tributary just N, 6 Jun 1995, Kra/ 84888 (BM, BRCH, FSU, GA, GH, KANU, MICH, MO, NCU, NY, TEX, VDB). Mobile Co.: bars and banks ee sat also es ‘islands” above sande Beall shallows, Puppy Creek, N end of AL 2 2 1 mi S its jct. with Prime Rd., WSW Citronelle above a ai bridge. pen or present, many toppled, 20 Sep. i R. Kral 83374 (BRCH, CM, PSU, GH, MO, VDB); revisited 10 Jun 1995, Kra/ 84994 (ALU, K, MBM, ae NY, USCH, VDB). Washington Co.: shallows and sandy silty banks of Bassett Creek, cypress- gum-maple, just N of Chatom by US 43, R. Kral 78266 (BRCH, CM, CTB, FSU, GA, GH, MICH, MO, NCU, NY, TEX, US, VDB, VSC, WAT), revisited 10 Jun 1995, Kral 84998 (GH, KANU, MICH, MO, MSC, NCU, NY, TEX, US, USCH, VDB, WIS); seeps and shallows of coldwater stream 0.5 mi N of Cty | on US 43, 2.7 mi N of Sunflower, 10 Aug 1992, R. Kral 81313 (CLEMS, GA, GH, MBM, MICH, MO, NCU, NY, TENN, USCH, VDB, WIS); shoals and sandy silty baal of oe s Creek, above and below US 43 bridge, 3.4 mi S of Milry, forming large colonies, 10 Jun 1995, R. Kral 85000 (APSU, BM, BRCH, CLEMS, CM, CTB, FSU, GA, GH, K, KANU, MBM, MICH, MO, NCU, NY, TENN, TEX, UNA, US, USCH, VDB, VPI, VSC, WAT, WILLI, WIS). FLORIDA. Bay Co.: small boggy stream in slash pine-titi flats by FL 22, 7.8 mi W of Sandy Creek bridge, 20 Jul 1993, R. Kral 82861 (ASPU, BM, CM, CTB, FSU, GA, GH, K, KANU, MBM, ; , US, USCH, eae Gulf Co.: sandy silty banks and shallows of Wettapo Creek at FL 2? crossing ca 3. mi W of We Merch 20 Jul 1993, R. Kral 82857 (AKG, BRCH, GA, GH, KANU, MIC H, MSC, NCU, VDB). Liberty Co.: sandy silty gravelly banks and shallows of Moccasin Creek, ie -Nyssa-Taxodium, Apalachicola Nat. Forest, SR 379, ultimately draining into Kennedy Creek. Abundant, 24 Aug 1995, R. Kral with A. Gholson 85600 (AKG, FSU, GH, MO, MSC, USCH, VDB), small stream near Larkin’s Farm Rd. by Co. 379, just S of Alligator nt 24 Aug 1995, R. Kral with A. Gholson ee (AKG, FSU, GH, MO, MSC, USCH, VDB). Santa Rosa Co.: moist sandy loam of low left bank of Big Coldwater Creek Shes canoe) ca 8 air mi NNE of Milton. New to Florida! 26 Jul 1986, L.C. Anderson 9827 (FSU, VDB); turf banks along East Fork of Big Coldwater Creek, 6.4 km above Tomahawk Landing, 22 Jul 1990, JA. Churchill 90164 (VDB, sent from set by Dr. Churchill); sandy-peaty banks of spring fed stream, along Big Juniper Creek at FL 191 biidee 2 mi SW of Munson, ca. 18 mi NE Milton, Blackwater River SF, 20 Sep 1989, S.L. Orzell & E.L. Bridges 12443 (TEX, VDB, etc.); FL 191, SW of Munson 2.8 mi; seeps along Juniper Creek just above bridge, i Sep 1993, R. ial 83353 (GH, NY, US, VDB); ca. 4.5 mi W of eee at FL 4 crossing Coldwater Creek; sandy-gravelly seep banks along stream, 19 Sep 1993, R. Kral $3339 (GH, VDB). GEORGIA. Marion/Talbot Cos.: sandy silty gravelly banks and low cuts of Upatoi Creek, both sides (these from N side), GA 41 bridge, ca 4 mi S of Geneva and E of Juniper, suffering from roe by DOT mowers, 13 Aug 1995, R. Kral 85554 (GA, GH, VDB). Montgomery Co.: sprigs atop Nyssa biflora stumps in small stream just below beaverdam, bz pees by US 221, 7 mi N of Uvalda, full sun, scarce, stream slack and turbid, 12 Aug 1995, R. Kral 85506 (MICH, MO, VDB). Toombs Co.: sandy silty gravelly banks of Pendleton Creek, scarce, by GA 152 bridge, NE of Lyons, 9 Aug 1995, R. Kral 85428 (VDB); sandy silty gravelly banks of Pendleton Creek, scarce, at Wixtrum’s Bridge, by GA 297, N of Vidalia below Interstate, 9 Aug 1995, R. Kral 85420 (TEX, USCH, VDB). Treutlen Co.: sandy silty gravelly banks and shallows, Red Bluff Creek by GA 29, NW of Soperton, scarce, first formal record for Georgia, 14 Jul 1994, R. Kral 83808 (CTB, FSU, GA, GH, NCU, VDB); sandy seep bank of Pendleton Creek, 0.5 mi $ of I-16 by US 221; made scarce by DOT cutting, 12 Aug 1995, R. Kral 85505 (MO,VDB). Wheeler Co.: wet sands, silts and gravels of swift creek (Ochwatkee trib.) above and below GA 46 bridge in full sun and light shade, 9 Aug 1995, R. Kral 85409 (GA, GH d 404 Sipa 17(2) NY, VDB, VSC). MISSISSIPPI. Greene Co.: peat-muck and sand of clear, flowing small stream through bay bottom in pine flats by MS 56, 17.7 mi S of State Line, 27 Aug 1991, first record for eas R. Kral 79807 (CTB, MICH, VDB); gravelly shallows of pine barren stream by MS 57, 11.3 miS of State Line, 11 Aug 1992, R. Kra/ 81349 (CTB, MO, VDB), gravelly sandy i - pine barren coldwater stream by MS 57, 15.7 mi S of State Line and 6.1 mi N jet. MS 63, 11 Aug 1992, R. Kral 81360 (CTB, GH, VDB), sandy gravelly seeps and shallows sf coldwarer pine barren stream by MS 57, 5.1 mi N of jet. MS 63 (Leaksville), | Aug 1992, R. Kral 81369(VDB). Jackson Co.: bars and shallows, grav- elly-silty-sand over clay, narrow of Bluff Creek, E side of MS 52 at S side of Van C ieee large colonies, culms mostly toppled by flood, inundated ee looking black; Sl a oresent, 7 Jun 1995, R. Kral 84936 (BM, BRCH, CM, CTB, FSU, GA, GH, , KA MBM, MICH, MO, MSC, NCU, NY, TENN, TEX, US, H, VDB, VPI, a cue WIS); sandy silty banks and shallows of small stream by MS 52, ca 5 mi S of George Co. line, occasional, some in full sun, some shaded by Cyri//a, 7 Jun 1995, R. Kral 84937 (CLEMS, GH, MO, VDB). Wayne Co.: Clara, SE of jet. MS 63 & Big Creek, T7N, R7W, S89, open area, on sandy soil in creek, swift flowing water, 1 Sep 1993, C.T. Bryson 12920 (CTB, VDB); gravelly shallows of clear, swift stream by MS 63, at Clara, near roadside park, 20 Sep 1993, R. 85382 (CM, CTB, GA, GH, MICH, MS, MO, NCU, NY. TEX. US, USCH, VDB, VPI, VSC, WILLI, WIS). NORTH CAROLINA. Hoke Con Fore Bragg Milicary Tea semi-shady bank of Little River, stable Cretaceous clay soil, plants swept prostrate by flooding. First NC record! 27 Aug 1992, B.A. Sorrie s.n. (dupl. sent to VDB by Mr. Sorrie). Moore Co.: shallows and banks of swift lowing Buffalo Creek (trib. of Little River) by SR 1001 (Lobelia Rd.) 8.5 mi E of Vass, plants abundant this locality, 10 Aug 1993, R. Kral 82960 (FSU, GH, MICH, MO, MSC, NCU, NY, VDB-site given by B. Sorrie from his earlier record!). Thus we now have several records from five southeastern states of a plant which, up until late 1989, was known only from two specimens, sketchily documented, collected by Dr. Mohr from Mobile Co., Alabama, and thought by many to be quite possibly extinct. Two Georgia sites for R. crinipes were collected by Dr. John Bozeman but not so identified. In 1990, doing loans kindly sent by NCU and GA I was able to find and annote them: GEORGIA. Appling Co.: Big Satilla Creek swamp on GA 99 at Bacon Co. line S of Baxley, 9 Jun 1967, J. Bozeman 9313 (NCU). Turner Co.: Little River swamp on Co. Rd. $1989 S of Sy- camore and 1.9 mi W of US 41 at Anaha, 15 Jun 1967, J. Bozeman 9720 (GA, NCU). Both are excellent examples and provided incentive to relo- cate the populations. Attempts in 1991, 1992, 1993, and 1994 however failed. It must be said that the two Bozeman localities had suffered since the 1960s from heavy and destructive logging both in their immediate vicinity and in the surrounding watershed. Also, in the years since 1991, the history of weather has been one of extreme flooding alternating with extreme drought and heat during the growing season, making it difficult to visit the sites at an optimum time. But the condition of the bottoms and banks of the two stream systems, disturbed as they are, lead me to suggest our that these two populations are extirpated. Kral, Rhynchospora crinipes (Cyperaceae) and related species 405 5. Rhynchospora curtissii Britton ex Small, Fl. SE U.S. 195. 1903. Phaeocephalum curtissii House, Amer. Midl. Naturalist 6:201. 1920. Reece Polis A. ee var. ellipsoidea Kiik., Bot. Jahrb. Syst. 75:107. 1950. Type: FLOR A Co.: moist roadside, Milton, Curtiss 5929 (HOLOTYPE: NY; ISOTYPES: i. cee i R. fusca sensu Fernald, Bot. Gaz. 24:433. 1897, non (L.) W.T. Aiton. — Densely cespitose, e-rhizomatous, perennial 1—-3(—4.8) dm high. Cu filiform, ascending to excurved, subterete, coarsely few-costate ca. 0..3—0.4 mm thick, leafy toward and at base. Leaves erect to ascending or excurved, filiform, the principal ones mostly 1-2 dm, the narrow, conduplicate sheaths terete, multicostate, the scarious ventral side apically transverse, the blades few-nerved, 0.2—0.3 mm wide, at base strongly involute, upblade scarcely tapered, canaliculate, slightly compressed or 2-edged, blunt apically. Ter- minal and penultimate fascicles usually compound, mostly narrowly turbi- nate to ellipsoid or oval, open or dense, the lower ones simple, of few (-1) spikelets, subtending bract linear-setaceous, exceeding lower fascicles, ms mostly exceeded by, rarely longer than terminal fascicle; spikelets lanceoloid, red-brown, 4.5—7 mm, narrowly acute; fruiting bracts mostly 2-5, nar- rowly oblong-ovate or elliptic, 3-4.5 mm, acute, mostly convex, only the midcosta evident, this short-excurrent as mucro or aristula; stipe and re- ceptacle short, 0.1-0.2 mm, setose; perianth of 6, antrorsely barbellate, basally pilose bristles 3-3.7 mm, extending well past tubercle tip; stamens 2, anthers linear, ca. 2 mm; fruit body lenticular, narrowly oblong-obo- vate, 1.2—-1.5 mm margined, apically transversely troughed at junction with raised tubercle base, faces lustrous brown around a paler, usually glassy central disc, very finely lined longitudinally.transversely with wavy lines of fine pits; mature tubercle triangular or slightly concave-sided, edges setulose. Figure 6. Moist to wet sands and peats of bog edges, pineland pondshores, seeps and pine-cypress savanna, Atlantic and Gulf Coastal Plain, locally abun- dant, eastern North Carolina intermittently south through South Carolina, Georgia to northern Florida, west to western Florida, southern Alabama = and Mississippi. Figure 3. This low, wispy sedge forms dense tussocks as do R. rariflora, R. oligantha, and like them seems to prefer the black fine sandy peats of frequently burned, unbroken pine savanna or shallow bogs in that type. Like R. oligantha it becomes more scarce as virgin savanna is converted to field or improved pasture. Habitally it most resembles R. pleiantha but of course lacks the rhizomes; ecologically however, it is in quite another system and borders more on habitat occupied by R. harperi, R. filifolia of the complex. imens anes previously, t am citing only VDB vouchers and some recent Pore U.S.A. RIDA: Bay : 1/2 mi W Youngstown on exposed wet sands 4O6 Stipa 17(2) Fic. 6. Rhynchospora curtiss1i (Kral 7197). A. Habit sketch. B. Leaf sheath-blade junction, adaxial view (left), side view (right). C. Leaf blade sectors from base (bottom) to apex (top). D. Terminal fascicle complex. E. Spikelet with (left) subtending bract. F. Fertile scale, stamen apex with anther (right). G. Face view of mature fruit. Krat, Rhynchospora crinipes (Cyperaceae) and related species 407 of seepage bog, 18 Aug 1962, R. Kral 15665 (VDB). Escambia Co.: | miles S jct. US 90 and co 289A W of Pensacola aa peat of pine flatwoods, 7 Jul 1963, R. Kral 17626 DB). Franklin Co.: 4.5 mi S on FL 65 from ject. with Fort Gadsden Creek, frequent, 31 May 1990, 8S. & G. Jones 5127 (AES, VDB); pine flatwoods, 3 mi N of Eastpoint, 21 Jun 1965, R. K. Godfrey 65859(FSU, VDB) sandy peat at edge of small cypress dome in scrub, end of shell road, East Point, ENE end of FL a 2( on 1993, R. Kral 82845 with G. Moore Lares MICH, OSC, PH, SMU, VDB). Gulf t peaty soil at margin of gum pond, 7 mi S of Wewahitchka, 18 Jun 1958, R.LK. he 0 (FSU, VDB),; wet ditch, border- ing pine flatwoods, 2.5 mi E of Port Saint Joe, 18 Jun 1958, R.K. Godfrey 57120 (FSU, VDB); N side White City; sandy peat of clearing in slash pine-palmetto flatwoods, 27 May ie R. Kral 63775 (V nee Liberty Co.: a peaty soil, broad roadside pes border- g pine flatwoods, 11.5 mi N of Sumatra, 14 Jun 1960, R.K. Godfrey & J.N. Triplett ee (FSU, VDB), me seat of bull Hee cypress-pine, FL 65, Telogia ees portion), 3 Jul 1993, R. Kral 82810 (CM, CTB, FSU, GA, GH, MBM, UNA, VDB). Santa Rosa Co.: common in wet sands of roadway across savanna, between I-10 and Milton, 24 Jul 1974, R.K. Godfrey 73791 (FSU, VDB); sandy peat of pine savanna by FL 191 exit off I-10 Gast Exit), 29 Jun 1991, R. Kral 79282 (VDB & distribution); pine and cy- press flats along Yellow River W of FL 87, N of Holley & S of I-10, sane peat of wet savanna-like area, 27 Aug 1991, R. Kral 797 . 2(VDB& cay n); 4.5 mi E of Gulf Breeze, common on moist pine savanna, 11 Oct 1963, S. Dantel 3862 (FSU, VDB). Wakulla Co.: sandy-peaty ditch bordering oo flatwoo on | marks ee Refuge, anes Sopchoppy and Panacea, 9 Jue 1960, R.K. Godfrey 59705 (FSU, VDB); mesic ee eaf pine / wiregrass - saw pal na on SW side of FS Rd 314 os at int of FS Rd 4-H, ca 0.4 mi NW i t FH 13 at ca 2.4 mi NE of int FL 375, ca 1.6 mi NE of Smith Crk School, peaennes NE, 24 Jan 1 a. SR. Orzell & E.L. Bridges (TEX, VDB). o.: sandy peat of hillside seep in longleaf pine by FL 20, ca 7 mi W of Ebro, 29 Jun on z Kral 79290 (VDB & coin Washington Co.: N side of FL 20, 6. 5 mi E of Ebro; sandy peat of low pine savanna, cypress-pine-titi, 26 Aug 1995, R. Kral 85623 (BM, CM, GH, K, MICH, MSC, NCU, NY, OS, VDB) 6. Rhynchospora filifolia A. Gray in Torrey, Ann. Lyceum Nat. Hist. ew York 3:366. 1836. Phaeocephalum filifolium (Gray) House, Amer. Mid. Naturalist 181. 1920. Type: U.S.A. NortrH Carouina {without county, annotated in Gray's hand (HoLotype: NY, lectotypified by Gale (1944)}. Cespitose e-rhizomatous perennial (1—)2—8(—10) dm high. Culms filiform, wiry, mostly excurved or arching, obtuseangled to subterete, 0.3—0.7(-1.2) m thick, coarsely few-costate, leafiest at and toward base. Leaves erect, ascending or excurved, principal ones l—ca. 2 dm long, blades filiform, mostly 0.2—0.5 mm wide (broadest ones actually bracteal and to 2 mm wide), proximally flattened or broadly “V”-shaped to slightly involute, finely nerved but with midcosta strongly raised abaxially, upblade strongly nar- rowed, progressively more deeply canaliculate and filiform, becoming unisulcate, thickened, the very apex thick, blunt, but with scabrid edges. Fascicles of spikelets mostly compound, 1-3(—4), sparse to dense, narrowly turbinate or even short-cylindric to hemispheric, often lobed, 0.7—1.5 x 0.7—1.5 cm, the terminal one well exceeded by at least 1 subulate-seta- ceous bract, the lower ones progressively longer peduncled, peduncles erect 408 Stipa 17(2) or excurved, ae by progressively longer, leafier bracts. Spikelets SS to lanceoloid, 2.5—3.5 mm, narrowly acute; scales several, the fertile ones 2—3(—4), ovate, ore mm, acute, convex, only the strong midcosta oe this usually excurrent as a mucro or arista; stipe and receptacle stout, ca. 0.2—0.3 mm, sparsely to densely setose and setulose; perianth of 6 antrorsely scabrid bristles 1.5—2 mm long; fruit body mostly broadly obovoid, 0.8—1 mm, lenticular, margined, faces light to deep brown, lus- trous, finely longitudinally lined, sometimes very finely cancellace and with wavy lines of small pits, but always with a strongly contrasting pale, glassy center; tubercle triangular 0.5—O0.7 mm, its edges variably setulose-scabrid, base shallowly lunate, slightly decurrent. Figure 7. Moist to wet sands, sandy peats or peat of grass-sedge meadow, pine savanna, seeps and bogs, pondshores, ditches, streambanks and bars, pine flatwoods, Atlantic and Gulf Coastal Plain, and contiguous physiology, New Jersey south to southern Florida, then westward to eastern Texas; Greater Antilles, Mesoamerica. Figure 8. This species is the most ample ecologically of the species treated here, which doubtless explains its highly variable habit as well as its occurrence within the habitat of all of these taxa. But as regards its inflorescence, inflorescence bracts, spikelet color and dimension, perianth and fruit the many hundreds of specimens examined in this study are remarkably con- sistent; the bracts consistently exceed inflorescences; the spikelets are the shortest, often of broadest outline but with fewer fertile scales, these the shortest and broadest in outline; the fruit bodies are the shortest, distinctly the smoothest, always with the glassy pale central “disc” on each face. It is, as one would suspect, the weediest of the species, is a quick invader of disturbed wetland sites so long as they remain undrained. Sometimes, ina single savanna in southern Alabama or northern Florida, it can present a considerable range of morphs, the moister or shadier situations having the taller, broader-leaved individuals, the more exposed or slightly drier sites with che shorter, narrower culmed-and-leaved individuals, these also often with the more sparsely spikeleted fascicles. A review of the literature available on R. filifolia if followed by an exami- nation of many specimens over the range, will show that Dr. Gale’s taxo- nomic disposition appears to be closer to the real situation than is that of Dr. Ktikenchal. Since even a recording of specimens available here at VDB would be very lengthy, and since this distinctive species varies so little as to key characters, complete citations are not included here. Duplicates of many of my sets have been distributed as follows: Representative specimens: U.S.A. ALABAMA. Baldwin Co.: Kra/ 85310 (APSU, BM, CM, CTB, F, FLAS, FSU, GA, GH, K, MICH, VDB). Covington Co.: Kra/ 84588 (BRCH, Krat, Rhynchospora crinipes (Cyperaceae) and related species 409 nn fF SY S KP we Wes 4 RY ‘ f \. SUA A NTA <— SSE XY —. Dd ~~ Se SEES mm long, the bracts subulate, glabrous, 3 mm long, the pedicels 2 mm long. Flowers 5-merous; sepa/s subulate, tend- ing to be of unequal length, S—/ 1 mm Jong at anthesis, accrescent to 12 mm long and exceeding the mature capsule at maturity, the tips acute, the margins and midvein scabrous; corolla broadly funnelform with aaa lobes, fleshy, light pink outside with a green tinge at the base of the tube, two darker pink vertical lines on each lobe, lighter pink to white inside, 36-50 mm long, the throat 8-15 mm wide, the free portion of the lobes ovate, acute, /2— 13 mm long, with scabrous margins, the lobes spreading open at anthesis, sta- mens and style included; stamens epipetalous, borne |2—16 mm above the corolla base, the free filament 2 mm long, projecting inwards, the anthers 2—3 mm long, connivent around the style 1-2 mm below the stigmatic surface and dehiscing onto the pubescent portion of the style; pistil 24—27 mm long, the ovary globose, 1-2 mm long, green, glabrous, the style white to yellow, slender, 23-25 mm long, pubescent along the upper 6—9 mm, articulated 5-6 mm above the ovary, the stigma terminal, truncate to rounded, yellow, pubescent, less chan 1 mm wide. Capsule bilobed, slightly crested on the top, glabrous, green, 8 mm high, 6 mm wide, the persistent style segment erect, dark brown, 5-6 mm long, the persistent, subtending fruit base white, 6 mm long with emarginate tips. — Type: ALABAMA. Biss Co.: large Ketona dolomite ae on hillside above the Little Cahaba River from County Road 65, off Hwy. 25, 30 May 1996, Katherine Gould 145 with Sheryl Gould (HoLotypr: TEX; isotypes: FLAS, FSU, GA, oe UNA GouLb, A new, disjunct variety of Spigelia gentianoides (Loganiaceae) 419 Spigelia gentianoides var. alabamensis, habit, flowering plant. B. fruiting plant. C mature aren corolla cut open longitudinally to reveal stamens and pistil. D. upper por- tion of style and stigma showing relative anther position Additional specimens examined: Bibb Co.: N & E of Centreville [sic], dry, shallow soil accumulations on limestone outcrop, ] Jun 1992 y Allison 6688; Co. Rd 65 0.6 mi NW of Lictle Cahaba ses bridge at en Dog Bend, downstream, 33°03°33"N, 87°02’ 10°W, elev. 0.25 mi S$ on dirt road to river, 0.2 mi 280 ft, Ll Jun 1995, Ginzbarg aa (UNA). The most obvious difference between the two varieties is flower size: Spigelia gentianoides var. alabamensis has consistently longer corollas (36-50 mm vs. 25—30 mm) with broader throats (8-15 mm vs. 7-8 mm), longer lobes (12-13 mm vs. S—7 mm), longer pistils (24-27 mm vs. 17-19 mm, and longer sepals (8-11 mm vs. 4-6 mm). Furthermore, var. a/abamensis generally produces shorter inflorescences than var. gentianoides (2—4 flowers vs. 3-8). The overall appearance of the flowers in the two taxa is the same, except that the flowers of var. gentianoides barely open when mature, this 420 Stipa 17(2 ~~ appearing cleistogamous, while the flowers of var. a/abamensis open fully at maturity, with their corolla lobes reflexing. Leaf shape, though variable, tends to differ between the two popula- tions: var. gentianoides has leaves that are more broadly ovate with rounded bases, whereas var. a/abamensis produces leaves that are more lanceolate to elliptic, less frequently narrowly ovate, ovate or obovate, and with cuneate to rounded bases. Overall stem leaf size is the same between the two varieties. Other qualitative and quantitative traits differing between the two vari- eties 7 situ, such as plant height, internode length, branching frequency, leafand stem texture and leaf orientation, may be induced by environmen- tal conditions, such as degree of sun exposure and differences in soil com- position. The var. a/abamensis occurs in an open, glade habitat in shallow, gravelly soil, where it often grows in low clumps. It generally has a com- pact, leathery habit with short internodes and ascending leaves with the leaf margins curling upwards. In contrast, var. gevtianoides is usually found under the shade of a pine/mixed hardwood canopy or pine monoculture in humous- or duff-covered soil. In this environment it generally has one chin, weak stem with elongate internodes and thin, outspread leaves. However, yen grown in full sun and _ — the var. gentianoides takes on the former habit w sandy soil, as it does at Bok Tower Gardens’ Endangered Plant Program in Lake Wales, Florida. Populations of Spigelia gentianoides var. alabamensis are much larger in extent and number of individuals than the present-day populations of var. gentianoides, with some glades containing thousands of individuals. How- ever, the new variety appears to be entirely localized to these rare glade communities, and should therefore be considered a rare plant in need of protection. ACKNOWLEDGMENTS Tam grateful to James Allison for calling the new Spige/ia populations to my attention and providing feedback on this description, James Affolter for his personal observations and helpful comments, Angus Gholson for his field assistance in Florida, Steven Ginzbarg for providing plant material, Tammera Race for access to the Endangered Plant garden at Bok Tower Gardens and providing plant material, Billie L. Turner for academic advis- ing and editing, Gayle Turner for the Latin diagnosis, and an anonymous reviewer for comments on the manuscript. REFERENCES ALLISON, J.R. 1994a. A botanical “lost world” in central Alabama. In: J.S. Fralish, R.C. Anderson, J.E. Ebinger & R. Szafoni. Proceedings of the North American conference on barrens and savannas. Hlinois State University: Normal, IL. Pp. 323-327. ~ Ne —_ GouLp, A new, disjunct variety of Spigelia gentianoides (Loganiaceae) 4. a «1994b. A “lost world” in Bibb County, Alabama. Panga 4 Krat, R. 1983. Loganiaceae. In: A report on some rare, threatened, or andaa kee’ forest- related vasc ular plants of the South. USDA Forest Service, Southern Region, Tech. Publ. R8-TP2, Vol. 2, oe 877-880. Rocers, G.K. 1986. The genera of Loganiaceae in the southeastern United States. J. Arnold Arbor. 67:143-185. bo Sipa 17( BOOK NOTICE ANDREWS, JEAN. 1995. Peppers. The Domesticated Capsicums New Edition. (ISBN eae ae hbk.) University of Texas Press, P.O. Box 7819, Austin, Texas 78713-7819, (512) 471-4032, $65, 274 pp, 34 full page color plates, 24 ae photos, 20 b&w illustrations, 3 maps, 11 tables. a published in 1984, Peppers has become a chili icon—the classic and com- plete source for the history, biology, taxonomy, cultivation, medicinal and culinary uses of the domesticated Capsicum. This new addition has updates in each section, Artist and author, Andrews takes unique and creative license of s style by leaving the foliage nearly devoid of color in each ere which places more emphasis on the actual peppers. This makes the 34 full color plates not just botanical illustrations, but selective windows to the individuality of each Capsicum. Flaws on stem and fruit, the odd chewed leaf here and there brings the illustrations a naturalness juxtaposed to the unreal aspect of colorless leaves. You can almost feel the skin of the jalapefo with the characteristic crack- ing of its flesh as it ages on the stems. yer Ph.D. in art from the University of North Texas and it has ore, researched new addition. So beautifully dr. Andrews received stead her well in this, once again, even m illustrated, it’s worth looking at again and again.—Limy Heagy. Sipa 17(2): - . 1996 THE CURRENT STATUS OF CYPRIPEDIUM KENTUCKIENSE (ORCHIDACEAE) INCLUDING A MORPHOLOGICAL ANALYSIS OF A NEWLY DISCOVERED POPULATION IN EASTERN VIRGINIA TROY W. WELDY, HENRY T. MLODOZENIEC, LISA E. WALLACE, and MARTHA A. CASE! Department of Biology The College of William & Mary Williamsburg, VA 23187-8795, U.S.A. ABSTRACT In 1 i a eure of 120 very large yellow hee s slippers was discovered in a remote ravine on the Northern Neck peninsula of Eastern Virginia. Multivariate and univariate ie of oii al variation in the Vecinid population suggest that these indi- viduals belong to a rare species, Cypripedium kentuckiense, and not the more common yellow lady's slipper, C. parviflorum var. pubescens. These analyses also indicate that the dorsal sepal width and orifice length are two important characters that can be used to discriminate between C. kentuckiense and C. parviflorum var. pubescens. A few individuals at the Virginia site approached the size of C. parviflorum var. pubescens in some characters which may be due to historical introgressive patterns or genetic isolation. Lastly, a survey of records on C. Rentuckiense populations revealed that Arkansas contains the largest number of populations (70 out of 156). These records also indicate that C. kentuckiense is characterized by a major- ity of populations (58%) with less than 21 individuals. RESUMEN En 1995 se descubrié6 una poblacién = 20 individuos de zapato de venus (“yellow lady’s slippers” en inglés) en un | localizado en la peninsula llamada “North- = & ) Neck” del este del estado de Virginia, EEUU. Un analisis eae y univariante e la variaci6n morfoldgica en la poblacion investigada de Virginia sugiere que estos especimenes pertenecen a una especie rara, Cypripedium kentuckiense, en vez de la especie mds comtin de zapato de venus C. parviflorum var. pubescens. Este andlisis también indica que la anchura del sépalo dorsal y la longitud del orificio son dos caracterfsticas importantes que pueden ser usadas para ditioeun C. kentuckiense de C. ah a var. ees Unos especimenes del lugar de investigaci6n en Virginia se aproximan al tamano de C. par ia var. pubescens en algunas caracterfsticas que pueden explicarse debido a patrones histér ntrogresivos o aislamiento ac Por dltimo, una revision ne cae “de investigaciones sobre poblaci le C. kentucksense revelé que el estado de A el ndmero mas ceande de poblaciones (70 de 156). Estas investigaciones también wadlicas que C. kentuckiense esta caracterizado por una mayorta de poblaciones (58%) con menos de 21 especimenes. 'Corresponding Author Sipa 17(2): 4 435.1996 Sipa 17(2) bo aN INTRODUCTION The Cypripedium calceolus L. complex in North America is currently thought to comprise four species and three varieties (Sheviak 1992, 1994), i. includes the yellow lady’s slipper taxa [C. parviflorum var. parviflorum alisb., C. parviflorum var. pubescens (Willd.) Knight, and C. parvifloram var. a (Farw.) Sheviak} as well as lady’s slipper taxa chat contain white and creamy white labella (C. montanum Douglas ex Lindley, C. candidum Muhlenb. ex Willd., and C. kentuckiense C. Reed). Cypripedium parviflorum occupies the largest range of all species in the North American group, oc- curring in approximately 40 states and throughout most of Canada (Luer 1975). In addition to its large range, C. parviflorum displays high levels of morphological variation which has led to the delimitation of many specific and subspecific taxa (see review in Newhouse 1976). The delimitation of taxa within C. parviflorum is complicated by complex si ization pat- terns among its varieties and related species (e.g., Klier et al. 1991; Case 1993), regional variation in reproductive isolating mechanisms (Gz [9D ); and very high levels of genetic variation (Case 1994). The extensive mor- phological variation and complex breeding patterns of C. parviflorum and related taxa have caused them to be the subject of many taxonomic and evolutionary debates that have persisted for more than 200 years. One of the most recently described species in the N. American group is vecies most closely resembles C — the relatively rare C. kentuckiense. This s parviflorum var. pubescens in morphology and appears to be a taxon that was recently derived from C. parviflorum var. pubescens (Case 1994). Although C. kentuckiense was not validly described until 1981, its morphological con- cept may date back to an entity described by Rafinesque in 1828, named C. luteum var. grandiflorum Rat. However, the lack of a type specimen desig- nation for the Rafinesque name precludes a definitive association between this name and a biological entity (Atwood 1984). Prior to its valid descrip- tion by Reed, C. kentuckiense was considered part of one polymorphic North American lady’s slipper species (e.g., Correll 1950) or considered to be a distinct species [Soukup (1977) invalidly described this species under the name C. dau/tonii}. Shortly after its valid description, Atwood (1985) conducted a study to clarify the range of C. kentuckiense by examining herbarium and live speci- mens from the southeastern United States. He concluded that Arkansas had the largest number of localities (11), but suggested that this might be an artifact due to less collection effort in some states [e.g., Mississippi (1) and Alabama (1)}. He also found locations for C. Aentuckiense in Oklahoma (4), Texas (2), Louisiana (5), Tennessee (1), and Kentucky (4), but did not find this taxon represented in the Carolinas or Virginia. He suggested that CAsE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) 425 conservationists be on the lookout for this taxon in Missouri, Illinois, Indi- ana, Ohio, West Virginia, and Georgia. Since then, there have been numer- ous attempts to find C. Aentuckiense in these bordering states, but there are no documented records of any discoveries to date (Larry Morse pers. comm.). Currently, the global rank of C. entuckiense indicates that it is threatened throughout its range (G3), while the federal ranking (C2) indicates that it is a likely candidate for the list of federal endangered and threatened spe- cies. However, its C2 rank also implies that it will be necessary to obtain further biological information pertaining to potential threats to the species to determine whether or not it should be placed on the federal list (Depart- ment of the Interior 1993). At the state level, C. Aentuckiense is most often given S1 status (i.e., critically imperiled, extremely rare, and very vulner- able to extirpation; Table 1). In 1995, a population of C. Rentuckiense was discovered (Weldy 1995) on the Northern Neck peninsula of eastern Virginia (between the Rappahannock and Potomac rivers), approximately 285 mi from the near- est known C. entuckiense locality in northeastern Kentucky. The popula- tion contains approximately 120 individuals and is located along a sandy ravine bottom adjacent to a small stream (hereafter the population is called the Virginia population). The most notable aspects of the Virginia popula- tion are the large overall size of the individuals and their creamy yellow labella (Fig. 1). The latter characteristic is far more typical of C. kentuckiense than C. parviflorum var. pubescens, which usually has bright yellow labella. Although this population strongly resembles C. entuckiense, some indi- viduals approach the size of C. parviflorum var. pubescens, which is a smaller taxon. Therefore, the first objective of this paper is to quantify the morpho- logical variation within the Virginia population of C. kentuckiense in rela- tion to other C. kentuckiense and C. parviflorum var. pubescens populations. Our second objective is to review the number of populations, population sizes, and locations of all C. entuckiense populations. This review 1s in- tended to provide a phytogeographical perspective for the Virginia popula- tion and provide an updated review of the national status of C. kentuckzense. MATERIALS AND METHODS To obtain information on the current status of Cypripedium kentuckiense populations, element occurrence records were examined from Natural Heri- tage offices in states with known localities. These states included Alabama, Arkansas, Kentucky, Louisiana, Mississippi, Oklahoma, Tennessee, and Texas. In addition, Heritage offices in states Siu to those containing C. Rentuckiense populations were contacted for data on C. kentuckiense sites. These included Georgia, Illinois, Indiana, Missouri, North Carolina, Ohio, South Carolina, Virginia, and West Virginia. For each state, records were 126 Sipa 17(2 Fic. 1. Cypripedium kentuckiense in Virginia. Photograph by T. Weldy. obtained on the number of known populations and the number of indi- viduals in each population. When state records included a range for any given population size, the maximum population size was used in our re- view. If information from more than one survey was indicated for any given population, the most recent record that reported the number of individuals in the population was used. It should be noted, however, that in some cases the most recent record was an original record that was unverified by Heri- tage botanists. Therefore, the number of populations reported in this study may be an overestimate due to the inclusion of records that have not re- cently been verified. Populations for which no data were given on the num- ber of individuals present were included in the total count of populations but were noc included in any other statistics. Thirty-five percent of al populations reported did not contain census data on the number of indi- viduals present. Therefore, the numbers of individuals per state given in Table | are likely to be underestimates of the actual number per state. CASE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) 427 To establish the taxonomic identity of the Virginia population, multi- variate and univariate analyses of 13 morphological characters were con- ducted. Nine of these characters were used by Reed (1981) in his diagnosis of C. kentuckiense. The specimens used in this analysis included 14 indi- viduals from the Virginia population of C. kentuckiense, 5 individuals from 5 C. kentuckiense populations in Texas and Arkansas, and 35 individuals from 13 populations of C. parviflorum var. pubescens from Michigan, Mis- sourt, Georgia, Indiana, and Virginia. For each specimen, the following morphological characters were measured: plant height (to the top of the ovary), staminode length, length and width of the longest leaf, length and width of the petals, length and width of the dorsal sepal, length and width of the labellum, length and width of the labellum orifice, and the number of 180° turns in the lateral petals. Those characters in this analysis that were not used by Reed (1981) include the orifice dimensions, staminode length, and the number of 180° turns in the lateral petals. In addition, we noted labellum color in the field but did not include it in the quantitative anne All measurements were made on living specimens in natural popu- lations by L. E. Wallace, M. A. Case, or H. T. Mlodozeniec. Multivariate analysis of morphological characters consisted of a princi- pal components analysis (PCA) using NTSYS (Rohlf 1988) to evaluate the existence of natural groupings among all 54 specimens. A variance/covari- ance matrix was calculated from standardized measurements (i.e., these measurements were in units of standard deviation from the mean) and the first cwo principal axes were extracted. The axis coordinates for all the indi- viduals were subsequently plotted. For univariate statistical analyses, the existence of unequal sample variances, non-normal sample distributions, and a relatively small sample size of the western C. kentuckiense group pro- hibited the use of parametric tests. Therefore, a Kruskal-Wallis non-para- metric mean rank test (Sokal & Rohlf 1995) was applied to test for charac- ter differences among the three groups. These groups were defined as the Virginia population, the western C. kentuckiense individuals, and all C, parviflorum var. pubescens individuals. Subsequently, a Dunn’s non-paramet- ric multiple comparisons test was used to test all pairwise comparisons among the groups (Zar 1996). The latter was applied to those characters that displayed overall significant differences among groups in the Kruskal- Wallis test. Arithmetic means, standard errors, and ranges of the charac- ters were also calculated using EXCEL (Microsoft 1993). RESULTS The survey of element occurrence records revealed that there are 156 popu- lations of Cypripedium kentuckiense known. Arkansas had the largest number of populations as well as 66.4% of all reported individuals (Table 1). Ken- 428 Stipa 17(2 Do 4 tucky had the second largest number of populations (36) and Louisiana hac the third largest number (19). However, the mean population size in Loui- siana (6) is very small compared to the mean population sizes reported in other states (17-117). Therefore, Louisiana contains only 1.5% of all re- ported individuals. Only 9 populations of C. kentuckiense are known from Tennessee, but this state contains 13.4% of all reported individuals. This is due to the relatively large population sizes found in Tennessee. Lastly, the mayority (58%) of population sizes reported from all states ranged between 1—20 individuals (Fig. 2). Furthermore, 43% of all populations had only 1-10 individuals. The PCA revealed two distinct groupings on axis | which separates C. parviflorum var. pubescens from C. kentuckiense individuals (Fig. 3). The first axis explained 72% of the variation whereas the first two axes combined explained 81% of the variation. With the exception of the degree of petal turns which had a low negative correlation with the first axis (-O.11), all remaining characters had strong positive correlations with the first axis. These values ranged from 0.70 for leaf length to 0.96 for dorsal sepal width. The remaining characters all had correlation values greater than 0.81. Results of the univariate statistical tests parallel che multivariate re- sults. With the exception of the degree of petal turns which was not significantly different among groups (p > 0.05), highly significant differ- ences (p < 0.001; Kruskal-Wallis test) were found among groups for all other characters. In pairwise group comparisons for these twelve significant characters (Dunn’s non-parametric multiple comparisons test), C. kentuckiense from Arkansas and Texas were never significantly different from C. kentuckiense from Virginia. In contrast, each of these two C. kentucksense groups were highly significantly different from the C. parviflorum var. pubescens group (Table 2). In general, the upward range limits of character measurements for C. par viflorum var. pubescens overlap slightly with the lower range limits of characters recorded for the two C. kentuckiense groups. For a few charac- ters (e.g., dorsal sepal width and orifice length) the measurement ranges of C. parviflorum var. pubescens relative to the C. kentuckiense groups were well separated and non-overlapping. These multivariate and univariate statisti- cal results reflect the large size of most C. Rentuckiense individuals relative to C. parviflorum var. pubescens individuals. DISCUSSION The morphological evidence presented in this paper indicates that the Virginia population is best interpreted as a disjunct population of C, kentuckiense, and not as a new taxon or population of large C. parviflorum var. pubescens. Our multivariate and univariate data analyses are consistent with the hypothesis of two morphological groups. In all group comparisons where CASE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) 429 Tape 1. Distribution and density statistics for Cypripedium kentuckiense populations throughout its known range. The information as includes: the state where populations are located (first col- umn); the state occurrence rank for C. kentuckiense [second column (SI = critically imperiled in state, extremely rare, and very vulner aie to extirpation; SU = possibly in peril in state, but status uncer- tain; §3 = rare or uncommon in state)}; the month and year the data were obtained (third column); the total number of known populations per state and percent of all populations found within each state (f fourth cn the sort pues bs aris Apehiiad ag state and percent of all individu- als found I Ich tion, and range of population sizes calculated from occurrence records (sich aa. a five percent of all occurrence records included census data on the number of individuals per population. Summary statistics are in the totals row. State Rank Date # Pops. & # Individuals & Meanzsd & (% of Total) (% of Total) (Range of Pop. Sizes) AK S3 4I95 70 (44.9) 2,884 (66.4) 60+94 (1-450) KX: S3 10/95 36 (23.1) 380 (8.8) 17+22 (1-100) LA Sl 10/95 19 (12.2) 67 (1.5) 6+5 (1-15) T™N Sl 10/95 9 (5.7) 584 (13.4) 117+108 (14-300) TX Sl 3/95 8 (5.1) 110 (2.5) 18415 (G-42) OK Sl a/95 7 (4.5) 148 G.4) 21+25 (3-60) AL Sl 10/95 4 (2.6) a a MS SU 3/95 2 (1.3) 50 (1.2) __ VA = _ 1 (0.6) 120 (2.8) Totals = am 156 (100) 4,343 (100) 42+75 (1-450) or Mean overall significant differences exist, no significant differences were found between the C. Kentuckiense and Virginia population comparisons whereas C. kentuckiense and the Virginia population were always significantly differ- ent from C. parviflorum var. pubescens (Table 2). Furthermore, the dimen- sions of characters measured for each taxon are generally within the ranges previously reported (Table 2). However, the dimensions of eight characters measured for C. kentuckiense individuals fell outside the ranges previously published for this taxon in Reed (1981) or Gleason and Cronquist (1991). These characters and their corresponding maximum or minimum values were height (53 cm; Virginia population), leaf width (14 cm; Virginia popu- lation), petal length (14 cm; Virginia population), petal width (1.0 cm; Virginia and western C. kentuckiense), dorsal sepal length (10.5 cm; western C. kentuckiense), dorsal sepal width (5 cm; Virginia population), labellum length (6.3 cm; western C. Rentuckiense), and labellum width (4.5 cm; Vir- ginia population). It is important to note that these values substantially increase the previously published ranges for these characters and that the Virginia population as well as the more western C. kentuckiense populations contributed to these increases in character ranges. A substantial increase in Sipa 17(2) XY ao CO Percent of Populations 1-20 21-40 41-60 61-80 81-100 100+ Number of Individuals per Population Fic, 2. Distribution of population sizes for 102 (65%) Cypripedium kentuckiense population records reporting sizes. the ranges of characters for C. kentuckiense was also found by Sheviak in his treatment of Cypripedium for Flora North America (Sheviak pers. comm.; Table 2). The results of the present study and Sheviak’s results indicate that prior sampling of C. kentuckiense has not been sufficient to determine the morphological ranges of these characters. Therefore, we recommend that floristic researchers be aware of the changing concept of character variation in this taxon. Moreover, our analysis indicates that dorsal sepal width and orifice length may be the most important characters for discriminating between C. kentuckiense and C. parviflorum var. pubescens. These were the only two characters in our study that did not overlap between these two species (Table 2). Qualitative floral traits of the Virginia population are also consistent with traits found in more western sites. First, the Virginia population is composed of individuals with creamy yellow labella, dark maroon petals and sepals, and dorsal sepals with prominent green and maroon striationsnear the sepal base (Fig. 1). These characteristics were always present in the five C. kentuckiense populations visited in Texas and Arkansas, but they were absent or rare in the C. parviflorum var. pubescens populations visited (Mlodozeniec pers. obs.). Furthermore, the habitat of the Virginia popula- tion (1.e., sandy substrate in the flood plains of creeks) is consistent with aN OD CASE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) xis 2 (9 %) (% TL) | SIXV Fic. 3. Principal components analysis of 35 Cypripedinm parviflorum var. pubescens nae rat he circles), 5 C. kentuckiense individuals from Arkansas and Texas (black triangles), and 14 C. kentuckiense from Virginia (white circles The amount of variation explained by axis 1 and axis 2 is given in p saenehes ses. The morphological characters used in this analysis are given in Table 2 432 Sipa 17(2) Tasie 2. Arithmetic meansstandard errors and ranges (in parencheses) of pee characters obtained from 54 living specimens, and combined ranges of characters reported by Gleason and Cronquist (1991 )! Sheviak (pers. comm.)-, and/or Reed (1981)°. Measurements are in cm. Statistica groups include Cypripedium parviflorum var. pubescens from all aps itions (PUB), C. kentuckiense from Texas and Arkansas (KENT), a Virginia population of C, auld way 2 ifferent super- ie letters among groups within a Samp nQ.0 Gre al character indicate >in meat rank scores among pairwise § < 0.005 (*) or p < ‘01 (a *) eves (Dunn' s non- parametric multiple chraisanlsoct test fer iiusle al- Wallis mean rank te Character & ranges of measurements easurements from this study M yreviously reportec (individuals measured indicated by n) y Leaf width KENT; (4.3-15)° PUB (n=35) k VEG O+0.4 (2.8-10.5) KENT (n=5 ) VA (n= 14) ae (5.741.5 567.041.6 >60.841.5 PUB (20— go)! ee an (27.8- 60. 5) (63.0—70.0) (53.0—68.0) eaf length “E14 640.4 b19.5+40.4 Roe: UB (6— a - KENT (13-2: (9.0—21.0) (18.0—20.0) (15.0—20.0) neue (8.0—14.0) Staminode length “ET 240.03 b1.9+0.2 1640.06 (O.8—1.7) (1.5-2.5) (1.5—2.0) ae length 4**7 040.3 b11.540.3 b11.4+0.3 PUB (5-8)!; KENT (8—15.6)! (4.4-10.5) (10.5-—12.5) (9.5-14.0) Petal width “0.7+0.03 b1.0+0 b1.0+0 KENT (0.7-1.5)2 (O.5—1.0) orsal sepal length a**5 540.2 bg.440.3 68. 840.2 PUB (3-8)!; KENT (6.1-12.67 ( 5) (9.0-10.5) (7.5—10.0) Dorsal sepal width *2.24 4.540 b4.1+0.1 KENT (2.4-6.5)° (1.5-2.9) (3.5-5.0) La eae cea ae a#*3 80.1 5.9+0.2 b5,.540,1 PUB (3-5)!; KENT (4.1-6.5 5—5.8) (5.5-6.3) (4.5—6.0) Labellum width a**2_2+0.0 b3.9+0.06 b3.5+0.1 KENT (3.5)? (1.5-3.0) (3.7-4.0 (3.0-4.5) Orifice length “0.9 40.03 b3.9+0.06 b3 540.1 KENT (2.7-3.7)° (0.5—1.3) (3.7-4.0) (3.0—4.5) Orifice width “1040.04 b2.2+0.2 1.8 40.08 (O.5—1.5) (1.5—2.5) (1.4—2.5) Number of 180° turns in petals "3,740.2 a3.2+0.5 *2.7+0.4 (1-7) —)) (0-5) the habitat that is typically oo for C. kentuckiense in the eastern por- tion of its range (Whitlow | Even though no significant differences exist in the morphological char- acters measured among the Virginia site and the more western sites, the range of character measurements were generally much larger in the Vir- ginia population than in the more western sites (Table 2). This, in part, CASE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) 433 could be due toa relatively small sample size of the western C. kentuckiense. Alternatively, the Virginia population may be composed of individuals that differ slightly from populations in the remainder of the species’ range. A long period of genetic isolation or introgression with C. parviflorum vat. pubescens might have contributed to this pattern. Although non-overlap- ping groups were formed by the PCA, one Virginia C. kentuckiense indi- vidual clustered very close to an individual of C. parviflorum var. pubescens (Fig. 3). This pubescens individual was from a population that was 95 mi from the C. kentuckiense site. The similar PCA clustering region of these two individuals might indicate historical patterns of introgression. How- ever, Weldy (1995) did not find C. parviflorum var. pubescens at the Virginia site or in adj acent ravines. In addition, high levels of year-to-year morpho- logical variation within the same individual of C. kentuckiense have been observed by Sheviak (pers. comm.). Similar non-genetic contributions to phenotype might also explain why some individuals of C. kentuckiense ap- proach the dimensions of C. parviflorum var. pubescens. Research is currently in progress to address hypotheses concerning the population genetic char- acteristics of C. kentuckiense and C. parviflorum var. pubescens in Virginia. It is interesting to note that the Virginia population contains approxi- mately 120 individuals. This large population size is relatively uncommon for this species (Fig. 2) and might be considered especially unusual for a population at the fringe of its species’ range. Most documented C. entuckiense populations contain fewer than 21 individuals. Although small population sizes can be caused by many contributing factors (e.g., Harper 1987), we hypothesize that small C. entuckiense population sizes are, in part, due to low rates of population growth. Life history characteristics that could con- tribute to low rates of population growth include a low rate of seed germi- nation, high seed-to-adult mortality, and a long development time from seed to flowering adult. Furthermore, high seed dispersal rates are expected in C. kentuckiense due to its highly reduced wind-dispersed seeds. This com- bination of characteristics could account for the observed pattern of widely- dispersed, small, isolated populations. Lastly, slow population growth rates appear to occur in related species such as C. parviflorum var. pubescens. In this taxon, estimates indicate that a seed can require up to 16 years of growth before flowering (Curtis 1954). In other Cypripedinm taxa such as C. acaule Aiton, pollinator limitation may be an important factor in causing low levels of seed set (Davis 1986; Helenurm & Barret 1987; Gill 1989). Polli- nator limitation may also play a role at the Virginia site. Out of 120 indi- viduals examined in 1996, no capsules were present from the previous year and floral visitation during 1996 was very low (Case & DeWitt pers. obs.). Although a number of C. kentuckiense sites have been found since 1985, it is our opinion that this taxon should remain a potential candidate for the — 434 Stipa 17(2) federal endangered and threatened species list (1.e., category 2 or, with more documentation of threats to existing populations, elevation to category 1). Only 156 populations are known with the majority (68%) of the popula- tions occurring in only two states, Arkansas and Kentucky. In some states, such as Louisiana, this species seems to be particularly at risk. Approxi- mately 10 Louisiana populations (not included in this study) have been extirpated primarily due to logging and development (Julia Lark pers. comm.). The remaining states listed in Table 1 each contain less than 6% of all populations each. The infrequent and scattered occurrences of C, kentuckiense in states ranging from Virginia to Texas may be the remnants of more widely distributed and abundant ancestral populations. In this case, it will be particularly important to categorize the demographic, genetic, and anthropogenic threats to the extant populations, especially in light of the large number of small, apparently isolated populations. Potential fu- ture discoveries of populations in new regions, such as the mountains bor- dering Virginia and West Virginia might be expected. These discoveries could provide important new insights into the distribution and genetic structure of this rare orchid. ACKNOWLEDGMENTS We wish to thank Todd J. Bierbaum and Donna M. E. Ware for their helpful comments throughout this research. In addition, we thank Todd J. Bierbaum, Donna M. E. Ware, Lawrence K. Magrath, and Charles J. Sheviak for valuable critiques of a previous draft of this manuscript. Howard Fraser kindly provided the Spanish translation and we are thankful. In addition, we express our sincere appreciation to Heritage botanists, university and park personnel, and the many non-professional botanists who helped us obtain information and locate Cypripedium populations. Also, we thank Merck/AAAS Undergraduate Science Research Program and the College of William & Mary for their financial support of this research. REFERENCES Atrwoob, J.T. Jr. 1984. In defense of Cypripedium kentuckiense C. F. Reed. Amer. Orchid Soc. Bull. 53:835-841 1985. The range of Cypripedium kentuckiense. Amer. Orchid Soc. Bull. sb, M.A, 1993. High levels of allozyme variation within eek calceolus (Orchidaceae) and low levels of divergence among its va s. Syst. 18:663-67 994. Extensive variation in the levels of genetic Se and degree of Amer. J. Bot. 81:175-184. CorretL, D.S. 1950. Native orchids of North America. Chronica et oe Company, Waltham, MA. Curtis, J.T. 1954. Annual fluctuation in rate of flower production by native Cypripediums ~ relatedness among five species of Cypr pedium (Orchidaceae). CASE ET AL., Status of Cypripedium kentuckiense (Orchidaceae) 435 during two decades. Bull. Torrey Bot. Club 81:340-—352. Davis, R.W. 1986. The pollination biology of Cypripedium acaule (Orchidaceae). Rhodora 88:445-450. ene eae OF THE INTERIOR. 1993. Federal Register, Pare 1V 58(188):51160. ILL, 89. Fruiting failure, pollinator inefficiency and speciation in orchids. In: D. Otte mi A. Endler (Eds.). Speciation and its consequences. Sinauer, Sunderland. Pp. 458-481. GLEASON, H.A. and A. Cronquisr. 1991. Manual of vascular plants of northeastern United States and leas Canada. New York Botanical Garden, New York. Harper, J.L. 1987. Population biology of plants. Academic Press, San Diego HELENURM, 2 ee S.C. BARRETT. 1987. The reproductive biology of boreal forest herbs II. Phenology of flowering and fruiting. Canad. J. Bot. Sci paliege Kuier, K., M.J. Leoscuke, and J.F. WenpbeL. 1991. Hybridization and introgression in white and — ladyslipper eichiide (Cypripedium de and C. pubescens). J. Hered- ity 82:30 18. Luer, C.A. oo The native orchids of the United States and Canada excluding Florida. The New York Botanical Garden, New Yor Microsorr. 1993. EXCEL, Version 5.0. Micnosoe Corporation. Newhouse, C.J. 1976. Pollination biology in seven taxa of Michigan Orchidaceae and a eae) of Cypripedium calceolus in oe based on living plants and herbarium speci- s. M.S. thesis, Michigan State University, East Lansing. ea CE 1981. Carian pane oie Reed, a new species of orchid in Kentucky. Phytologia 4 426-4 Rou, EJ. 19 ees pc, numerical taxonomy and multivariate analysis system. Exeter S Shewale: cane SHEVIAK, C.J. 1992. oral hybridization between C Ce montanum and its yellow- lipped laa Amer. Orchid Soc. Bull. 61:546-559. «1:29. — parviflorum Salisb. I: an small-flowered varieties. Amer. Orchid Soe: Bull. a 664-666 OKAL, R.R. and EJ. ROHLrF. ie Biometry. W.H. Freeman and Company, New York. SoukupP, V. 1977. One daultonti, Soukup: sp. nov. A new lady’s-slipper from north- eastern Kentucky. Mid-American 56:9— WeLpy, T.W. 1995. The vascular flora of the Corrotoman River watershed, Lancaster County, Viren: M.A. thesis, The College of William & Mary, Williamsburg, Virginia WuitTLow, C.E. 1986. Cypr. see ire Raf. from southeastern United Grates Be it what it may. Orchid Diges 43. Zar, J.H. 1996. Sitar ae Prentice Hall, New Jersey. Sipa 17(2) BOOK NOTICE , DouGias J. FuruyMa, and FRANcEs C. JAMEs, eds. 1995. Annual Review of Ecology and Systematics. Volume 26. (ISBN 0-8243-426-3, hbk). Annual Reviews Inc., 4139 El Camino “a P.O. Box 10139, Palo Alto, CA 94303-0139. $47.00, 757 pp. FauTIN, DAPHNE Gall eleven chapters of Volume 26 are devoted to a special section on Sustainability Issues. From the Preface: “Several chapters were written by authors invited to discuss sustainability from the perspectives of disciplines not typically included in ARES. Topics and authors for the section were selected in part to demonstrate some of the ways in which this word is employed re underlying assumptions and prin- ciples used in discussions of and proposals for sustainability. This section will have achieved a diversity that extends tot one of its aims if readers come away recognizing some of these differences and we need to se explicit in their own usage and about the underpinnings of their positions.’ maining 19 chapters cover various aspec Women in Systematics, Molecular evid 4 —_ P ts of ecology and systematics. Ex imples inc ia ence for Natural Selection, The Role of Nitrogen in the Response of Forest Net Primary Production to Elevated Atmospheric Carbon Dioxide, Architectural Effects and the Interpretation of Patterns of Fruit and Seed Development and Plant-Vertebrate Seed Dispersal Systems in the Mediterranean: Ecological, Evolution- ary, and Historical Determinants. Indexes include: Subject; Cumulative Index of Contrib- uting Authors, Volumes 22—26; and Cumulative Index of Chapter Titles, Volumes 22-26. Stipa 17(2); 436. 1996 CONDUCTIVE TISSUE IN CERATOPHYLLUM DEMERSUM (CERATOPHYLLACEAE) EDWARD L. SCHNEIDER and SHERWIN CARLQUIST Santa Barbara Botanic Garden 1212 Mission Canyon Road Santa Barbara, CA 93105, U.S.A. ABSTRACT a Using scanning electron microscopy (SEM) and light microscopy, we attempted to es- tablish if xylem and phloem occur in Ceratophyllwm stems; roots are lacking in the genus. In larger stems, a central core of elongate cells contains scattered sieve tube elements with simple sieve plates; the sieve tubes are associated with companion cells. Sieve tubes are not organized into phloem strands. The central core of main and branch stems contains elon- gate cells, some of which lack starch and tannins. These cells lack any evidence of the annular or helical thickenings of secondary wall material characteristic of primary xylen in other vascular plants, and therefore xylem cannot be claimed for Ceratophyllum. ee e of xylem in Coan is interpreted as secondary, a loss related to adaptation to the submersed habi RESUMEN 0 mpleando la microscopia electrénica de barrido y la microscopia 6ptica, hemos intentac re si existe xilema y floema en los tallos de Ceratophyllum, no existen raices en € género. En los tallos mas grandes, un nucleo central de células alargadas elementos cribosos dispersos; esros elementos cribosos tienen pk acas cribosas simple SV estan asociados con células ene Los elementos cribosos no se organizan en paquetes de floema. E] nucleo central de los tallos puna y laterales contiene células alargadas, y en algunas falta el almidén y los taninos. Estas células no tienen engrosamientos espiralados o anillados en la pared secundaria caracteristicos del xilema primario en otras plantas vasculares, y no puede decirse que haya xilema en Ceratophyllum. La ausencia de xilema en tat eda se interpreta como secundaria, una pérdida relacionada con la adaptacién a rg INTRODUCTION Ceratophyllaceae have recently attracted attention because of shifts in concepts of their evolutionary relationships. Earlier phylogenetic schemes placed them nearer such families of aquatic dicotyledons as Cabombaceae (Ito 1987) or Nelumbonaceae (Thorne 1987); Les (1988) reviewed place- ments by various authors. Cladistic analyses based on macromorphological data have placed Ceratophyllaceae near Nymphaeaceae, in a basal position in dicotyledons (Les 1988). Cladistic analysis of rbcL data (Les et al. 1993, Qiu et al. 1993) confirmed this idea, and locates Ceratophyllaceae as the outgroup of all other angiosperms. Conceding that Ceratophyllaceae may SipA 17(2):; 437-443. 1996 438 Sipa 17(2) contain both specialized and primitive features, knowledge of any feature in the family is important in view of the phyletic placement now accorded Ceratophyllaceae. Klercker (1885) showed that Ceratophy//um has sieve-tube elements with associated companion cells (“cellules adjunctives”), but reported no ale cells. Jones (1931) found sieve-tube elements and companion cells; he claimed presence of phloem parenchyma, and also believed that “some of the sieve tubes show evidence of transformation into air spaces. This de- generation 1s particularly evident in stems in which the aerenchyma is strongly developed.” Jones further claimed that “the xylem is so degener- ated that there is in reality very slight evidence of this tissue. There are two kinds of cells composing the xylem. One is larger than the other in diam- eter and has thicker walls. It is probably a reduced vessel. The other is of the xylem parenchyma type. There is abundant evidence that the reduced vessel-like cells are becoming parenchymatous.” ee cited presence of tannins, chloroplasts, and starch in the “reduced vessel-like” cells surround- ing the central canal. In his legend for a transection of a mature stem, Jones's conclusions are somewhat ambiguous. He believed that there is “transformation of some xylem vessels into parenchyma cells,” but also considers that “it is perhaps more appropriate to designate all the tissue inside the phloem as pith since the cells appear to be assuming the func- tion of parenchyma and in a few cases, even that of chlorenchyma. Further- more they are quite similar to parenchyma cells in appearance. ” Jones (1931) failed to find any evidence of lignin in cells. The purpose of our study is to reinvestigate the nature of conductive tissue in Ceratophyllum, using both light microscopy and SEM. Is the ph- loem clearly referable to that concept, as Jones claimed? Does the central tissue of stems (either main or branch stems) deserve the designation of xylem, or is it merely pith? Is there any evidence that cells are “reduced” vessels or tracheids? Is reduction or absence of xylem related to adaptation to the aquatic habitat, as Jones (1931) claimed? MATERIALS AND METHODS Our materials of Ceratophyllum demersum L. were collected from two sites, both in Texas, during September, 1995. The primary collection site was Aquarena Springs, the headwaters of the San Marcos River in San Marcos, Texas. In this site, where clear water of constant temperature (21° + 1°C) emerges from a limestone aquifer, C. demersum is common, with several individuals anchored at depths of more than 5 m. Additional collections were obtained from Toledo Bend Reservoir, Sabine County in eastern Texas. Stems were fixed in 50% ethanol. Portions were dehydrated according SCHNEIDER AND CaRLQUuIST, Conductive tissue in Ceratophyllum demersum 439 to the tertiary butyl alcohol method (Johansen 1940), embedded in paraffin, and sectioned on a rotary microtome. Some sections were stained with a safranin-fast green series corresponding to Northen’s modification of Foster's tannin acid-ferric chloride method (Johansen 1940). Other sections were mounted on aluminum stubs much as paraffin sections would be affixed to glass slides; these sections were then cleansed of paraffin with xylem, sput- ter coated, and then observed with a Bausch and Lomb Nanolab SEM. ANATOMICAL RESULTS Transections of the main stem (Fig. 1) have a cortical region delimited by a circle of rather large cylindrical air canals, appreciably larger than the parenchyma cells of the cortex. Internal to the cylinder of air canals is what Jones (1931) considered the vascular core. Although we did not find any histological differentiation of the cells at the periphery of the core, Jones (1931) identified an endodermis in labels of his figures. Presumably he considered the narrowest cells at the periphery of the vascular core to be endodermis (Fig. 1). Within the vascular core, there are air canals (Fig. 1, center). Longisections show that these are air canals rather than enlarged cells. One of these may be placed centrally in the core (Fig. 1). This desig- nation is in agreement with Jones (193 1), who, however, thought of these stelar air canals as modified sieve tubes: “the erstwhile sieve tubes are being modified for this purpose.” In general, the sieve-tube elements are located in the outer portion of the vascular core, as one might expect in a dicotyle- donous stem. If one views a longisection of a Ceratophyllum stem, one sees that at nodal regions, cells of the central core are irregular in orientation, and the verti- cal air spaces are interrupted by a nodal plate (Fig. 2). The nodal region as seen in transection (Fig. 3) illustrates that strands of elongate cells extend from the vascular core into the branches that occur in verticillate fashion at the nodes. Examining with SEM the cells that lead from the central core into the branches, one finds that most of these cells are elongate, or prosenchymatous, in shape (Fig. 4). Many of these cells lack the starch and tannins seen with SEM in other neighboring cells; cytoplasmic remnants can be observed with SEM, however (Fig. 4, above) Neither in che main stem nor in the lateral stems did we see the features ordinarily used to identify primary xylem annular, helical, or reticulate bands of secondary wall superimposed on a primary wall. With SEM, one cannot find small pores in primary walls, invisible with light microscopy, that indicate presence of vessels by virtue of the porose walls (e.g., Barclaya, Schneider and Carlquist 1995). Xylem can be said to be present only by 440 Sipa 17(2) QLLTEVTTTTLATTTIAT TET AL __ tt he SE TUQMUS AQT ATHUTEUAPEEE TET ev anas : a wy: Fics. 1-2. Sections of mature main stem of Ceratophyllum demersum. Fig. 1. Transection of stem, showing cortex delimited from vascular core by a circle of air canals; smaller air canals are evident within the vascular core. Arrows indicate presumptive endodermis be- tween the large air canals. Fig. 2. Median longitudinal section of stem; the main air canals are interrupted at the nodal plate, middle of photograph; smaller air canals adjacent to the nodal plate extend into branches. Figs 1-2, divisions = 10 pm in magnification scales. designating the central portion of the core as xylem, as Jones (1931) did, although even he had reservations about doing so. Because neither light any structural characteristics of tracheary ele- pee microscopy nor SEM revea ments in this region, we prefer to consider that Ceratophyllum lacks xylem. Sieve-tube elements and companion cells (Figs. 5, 6) are present in the vascular core, usually more commonly around the periphery than in the center. The sieve plates are simple (Figs. 5, 6), with clearly defined pores that are densely placed and more uniform in size than primary pit field pores (Figs. 5, 6). Sieve-tube elments are generally narrower in diameter than the parenchyma cells they accompany, and are not organized into strands or groupings that one expects in phloem strands of stems. For this reason, we do not use the term “phloem parenchyma” as Jones (1931) did. Endress (1994) reports phloem, but no xylem, in floral structures. Fics. 3—G6. Sections from mature stem of Ceratophyllum demersum. Fig. 3. Transection of nodal plate, showing strands of elongate cells extending into bases of |: ‘eral branches. Fig. SEM photograph of elongate cells from longisection of ee branch; remnants of cyto- plasm are present on wall surfaces. Fig. 5. SEM photograph of sieve plate of a sieve-cube element from a transection of the main stem. Fig. 6. Sieve tube element (sieve plate in face view) surrounded by parenchyma cells, from transection of vascular core of the main stem. Fig. 4, scale as in Fig. 1; Figs. 4-5, scale bars = 10 pm. Fig. 6, scale divisions = 10 pm. Sipa 17(2) aN X NO DISCUSSION AND CONCLUSIONS The nature of sieve-tube elements in i aaa is of potential phylo- genetic interest, because the sieve-tube elements and their associated com- panion cells in Ceratophy/lum correspond to the most specialized phyloge- netic condition according to the criteria of Zahur (1959) for woody dicotyledons and Cheadle and Whitford (1941) for monocotyledons. These authors have considered that sieve-cube elements with little differentiation between end wall and lateral wall in morphology of sieve areas, and with no companion cells derived from the same initial as the sieve-tube element, are primitive in dicotyledons and monocotyledons. The genus Austrobaileya has been cited as exemplifying this condition (see Metcalfe 1987). How- ever, transverse partitioning ina sieve-tube element may result in end walls bearing simple sieve plates in a dicotyledon with vessels with long scalari- form perforation plates in xylem, as in Cercidiphyllum (Zahur 1959), and thus sieve-plate morphology may not be a reliable key to degree of phyletic advancement. Also, one can find simple sieve plates with conspicuous pores in peduncles of Victoria (Schneider 1976) and other Nymphaeaceae; this may be a byproduct of conductive rates, rather than phyletic status (Carlquist 1975). However, whatever the criteria used, the sieve-tube elements of Ceratophyllum do not show a condition that one would call primitive for dicotyledons. The status of sieve-tube elements is placed first in this discussion be- cause the sieve-tube elements of Ceratophy/lum do not suggest an incipient stage in the evolution of that cell type. Therefore, one would be tempted to consider the absence of xylem in Ceratophyl/um to be secondary, a loss or reduction related to adaptation to the submersed habit rather than indica- tive of a primitive status among angiosperms. If vegetative structures of Ceratophyllum are typically submersed, there is little selective value for con- duction of water from one part of the plant to another. However, there is, in flowering plants, a selection for phloem because the location of photo- synthesizing organs is almost always at a distance from sites for active stor- age of photosynthates—the fruits (or, in some plants, tubers or other stor- age organs). Although there is no xylem in the species of Ceratophyllum, studied herein there are elongate cells other than sieve-tube elements in the vascular core of stems. Such elongate cells could conceivably serve for moderate degrees of water conduction. Of prime importance in discussing the probability chat xylem was present ancestrally but has been lost is the primary xylem of Brasenia (Schneider and Carlquist 1996). In Brasenia, annular or helical thickenings of second- ary wall material are present at tips of tracheary elements. However, on lateral walls of the tracheary elements of Brasenia, these thickenings are reduced to inconspicuous ridges, visible only with SEM, that lack SCHNEIDER AND CARLQUIST, Conductive tissue in Ceratcophyllum demersum 443 lignification. Brasenia thus offers an example of a stage in reduction of xylary cells with relation to adaptation to the aquatic habitat. Although xylem reduction has been suggested for aquatic plants by a number of authors, likely stages in reduction (other than reduction in quantity of tracheary elements) have not been demonstrated for tracheary tissue of genera other than Brasenia. REFERENCES CaRLQUIST, S. 1975. Ecological strategies of xylem evolution. University of California Press, Berkeley and Los Angeles Cueab_e, V.I. and Wuitrrorp, N.B. 1941. Observations on the phloem in the Monocotyledoneae. 1. The occurrence and phylogenetic specialization in structure of the sieve tubes in the metaphloem. Amer. J. Bot. 28:623—627 Enpress, P. 1994. Evolutionary aspects a the floral structure in Coaiilion: Pl. Sysc. 83. Evol. Suppl. 8:175- Iro, M. 1987. Phylogenetic systematics of the Nymphaeales. Bot. Mag. (Tokyo) 110:17- a). JOHANSEN, D.A. 1940. Plant microtechnique. McGraw Hill, New York. Jones, E.N. 1931. The jar uae and biology of crear in demersum. Stud. Nat. Hist. lowa Univ. 13:11— KLercker, J.-E.-F. 1885. Sur : anatomie et Bie développement de Ceratophylum. Bih. Kungl. Svensk. Vet.-Akad. Handl. 9(10):1 Les, D.H. 1988. The _ and affinities of the Ceratophyllales. Taxon 37:326—3 Les, D.H., D.K. Garvin, and C.F. Wimper. 1991. Molecular evolutionary ie af an- cient aquatic angiosperms. Proc. Nat. Acad. Sci. USA 88:10119-10123. Merca.re, C.R. 1987. Anatomy of the dicotyledons, ed. 2, vol. 1. Clarendon Press, Ox- ford. Qiu, Y.-L., M.W. Cuase, D.H. Les, H.G. Hitis, and C.R. Park. 1993. Molecular snmlosenducs of the Manel lidae: a eladietic analysis of the nucleotide sequences of the plastid gene rbcL. Ann. Missouri Bot. Gard. 87:587-606. ee R, : 7 ale The floral anatomy of Victoria. Bot. J. Linnean Soc. 72:11 SCHNEIDER, E. nd S. Cariquist. 1995. Vessels in the roots of Barclaya rotundifolta (Nymy nen Amer. J. Bot. 82:1343-134 SCHNEIDER, E.L., and S. Cartquist. 1996. Vessels in Brasenia eae new perspec- tives on vessel origin in primary xylem of angiosperms. Amer 83:1236-1240. TuHorng, R.E. 1992. Classification and geography of the flowering ae oA Rev. 58:225- 348. —148. ZAHUR, M.S. 1959. Comparative study of secondary phloem of 423 species of woody 4 cotyledons belonging to 85 families. Cornell Univ. Agric. Exp. Stat. Mem. 358:1—16 eS ‘ aes Sipa 17(2) NEW SPECIES OF ARDISIA (MY RSINACEAE) FROM ECUADOR AND PERU JOHN J. PIPOLY Hl Botanical Research Institute of Texas 509 Pecan Street Ft. Worth, TX 76102-4060 U.S.A. ppipoly @ brit.org ABSTRACT Routine determination of specimens received as gift for determination resulted in che discovery of four new species of Ardisia, from Ecuador and Peru. Ardisia premontana Pipoly, A. zakii Pipoly, A. flavida Pipoly, and A. websterii Pipoly are described, illustrated, and their phylogenetic relationships are discussed. RESUMEN | determinar pliegos de herbario ae como regalo para su determinaci6n, se encontraron cuatro especies nuevas pertenecientes al género Ardisia, y procedentes de Ec- uador y Pert. Se describen e ilustran Ardisia ee Pipoly, A. zakii Pipoly, A. flavida Pipoly, y A. websterii Pipoly, y se discuten sus relaciones filogenéticas. INTRODUCTION The genus Ardisia Swartz is pantropical and contains approximately 400-500 species (Chen Cheih & Pipoly 1996). There are two geographic regions with high concentrations of species, Panama and adjacent Co- lombia, and the Malesian area (Stone 1982, 1989, 1990, 1992). The ge- nus is currently defined by the combination of mostly bisexual flowers, pluriseriate ovules (many times appearing uniseriate because of high anthotaxis), quincuncial, imbricate, or contorted corolla, relatively long style and often punctiform or apiculate stigma. Clearly, the entire tribe is badly in need of revision on a worldwide scale, because generic delimita- tion is often controversial and inconsistent even within a region. In the Neotropics, Aublet (1775), Urban (1922), Ducke (1930), and Lundell (1963, 1964, 1981a, 1981b, 1981c, 1981d, 1982) have segregated vari- ous groups from Ardisia as distinct genera, while in the Paleotropics, several genera have been synonymized (Stone 1993a, Larsen & C. M. 1995, Miller & Pipoly 1993, Taton 1979). Finally, Stone (1993b) de- scribed one new Asian subgenus, Scherantha. At this time, a phylogenetic analysis of the tribe will be necessary to resolve the usually disparate classifications recognized in the Americas, Africa, Asia and the Pacific. Therefore, for the taxa I have described from those areas, (Miller & Pipoly Sina 17(2): 445-458. 1996 446 Stpa 17(2) 1993; Pipoly 199 1a, 1991b, 1992, 1994a, 1994b, 1995) I recognized as few segregates as possible (such as Crenaridisia Ducke, Gentlea Lundell, Synardisia Lundell, and Solonia Urban for the Neotropics) and define the rest of Ardisza broadly, by informally recognizing a number of infrageneric groups. Routine determination of specimens sent as gift for determina- tion from the many Andean exploration programs of the Missouri Bo- tanical Garden resulted in the discovery of the following novelties, de- scribed herewith. Ardisia premontana Pipoly, sp. nov. (Fig. 1) Propter laminas magnas chartaceas ellipticas vel oblongas atque ad apices breviacuminatas, inflorescentiam terminalam bipinnatipaniculatamque, pedicellos obsoletos vel subobsoletos, margines sepalinos hyalinos, necnon antera nese super ba- sin dorsifixa, A. a/bovirenti valde arcte affinis, sed ab ea ramulis dense rufo-furfuraceo lepidotis (non glabris) angulatis vel subteretibus (nec teretibus), ik canaliculatis (non marginatis) 1.5—2.5 (nec 0.8—1.5) cm longisque, inflorescentia 17—19 (non 5—7.5) cm longa, calyce cupuliforme ees cotyliforme), lobis calycinis carinatis (non planis) glandulari-granulosis (nec glabris) ad apices obtusis (nec late rotundatis) secus margines glabris subintegris erosisque (nec incegerrimis glandulari-ciliolatis), corolla 4—4.5 (non 5—6) mm longa irregulariter dividentique, lobis corollinis ovatis cucullatisque Nae ellipticis planisque), filamencis antheris oer (non) we ay caters anthe ad apices acutis (non aliquantam emarginatis), denique | 11 (non sysides) praeclare distat. Tree to 10 m tall. Branchlets angulate to subterete, 5-7 mm diam., densely rufous furfuraceous lepidote, early glabrescent. Leaves alternate; blades chartaceous, elliptic or oblong, 23.5—30.6 cm long, (8—)9—10.5(— 11.7) cm wide, apically short-acuminate, the acumen ca. 0.5 mm long, basally acute to cuneate, slightly asymmetric, decurrent on the petiole, midrib canaliculate above, prominently raised below, smooth and sordid above, sparsely and minutely rufous lepidote below, glabrescent, densely but inconspicuously red punctate and punctate-lineate, secondary veins 36-48 pairs, notable but not prominent above and below, the margin entire, irregluar, flat, glabrous; petiole canaliculate, 1.5—2.5 cm long, densely and minutely furfuraceous lepidote tomentellous, glabrescent. Inflorescence terminal, pyramidal, bipinnately paniculate, 17-19 cm long, 11-24 cm wide, the rachis deeply angulate, densely rufous glandular- granulose, the branches spicate; inflorescence bract unknown; peduncle |—-1.5 cm, floral bracts membranaceous, linear, 1—1.2 mm long, 0.2-0.3 mm wide, cucullate, apically obtuse, densely glabrous above, glandular- granulose below, sparsely red punctate, the margin entire, glabrous; pedicels subobsolete to obsolete. F/owers 5-merous, chartaceous, yellow, 4—4.5 mm long; calyx narrowly and deeply cupuliform, 2.2—2.5 mm long, the tube 1-1.5 mm long, the lobes symmetric, ovate, 1—-1.2 mm long, iN oy | Pipoty, New species of Ardisia (Myrsinaceae) ie ye ty Lid ! ie io i 1G. 1. Ardisia premontana Pipoly. A. Habit. B. Abaxial leaf surface, showing minute rufous lepidote scales. C. Flower, showing obsolete pedicel, cupuliform calyx with ovate lobes and campanulate corolla. D. Stamens, showing anthers with acute, apiculate apices. E. Pistil. FE Placenta, showing biseriate ovules. A-F, drawn from holotype. 148 SpA 17(2) 0.6-1 mm wide, apically obtuse, medially carinate, minutely glandular- granulose, densely and prominently red punctate and punctate-lineate medially, the margin Supcninite except erose apically, hyaline, glabrous; co- rolla campanulate, 4—4.5 mm long, tube 1.8—2 mm long, the lobes i irregu- larly divided, ovate, 2.2—-2.6 mm long, 0.8—1 mm wide, hyaline, apically acute, somewhat cucullate, glabrous throughout, the margins entire; sta- mens 3.5—3.8 mm long, the filaments free, terete, glabrous, hyaline, 2.6— 2.9 mm long, the anthers elliptic, 1.4-1.6 mm long, 0.7—0.8 mm wide, apically acute, apiculate, basally obtuse, dehiscent by wide longitudinal slits, the connective epunctate; pistil obnapiform, 1.5—1.7 mm long, the ovary globose, 0.6—0.8 mm wide, densely and prominently red punctate, the placenta turbinate, ovules 3—4, biseriate, the style 0.9-1.2 mm long, the stigma truncate. Frv/t unknown. Type: PERU. HuAnuco: La Divisoria, Tingo Marfa- Pucallpa road near Loreto border, ca. 09°05' 8, 75°50' W, 1,150—1,250 29 Mar 1977 (fl), A. Gentry, D. Daly & S. Cruz {S804 (HOLOTYPE: MO; IsoTyPEs: a 7 USM). Paratype: ECUADOR. Morona- oe 1AGO: Bomboiza, 17 km SE ng ae 03°27! S, 78°34' W, 700 m, Jul-Oct 1985 (fl), J. Zaruma 315 (MO, NY, QCN = 5 Distribution —Known only from the eastern slopes of the Andes of south- ern Ecuador and central Peru, at 700—1,250 m elevation. Ecology and conservation status. —Ardisia premontana occurs in the species- rich premontane forests of the lower eastern slopes of the Andes, at the rim of the Amazon Basin. The Tingo Marfa area is well-known for its many endemic species, yet still remains underexplored. Because Ardisia premontana is known only from two specimens, no determination of its conservation status 1s possible at this time. Etymology.—The specific epithet refers to the habitat of Ardisia premontana, premontane moist forest. Ardista premontana is most closely related to Ardisia albovirens Mez (in- cluding A. wigrovirens MacBride), by virtue of its large, chartaceous, elliptic or oblong leaves with acute apices, the terminal bipinnate panicles, obso- lete to subobsolete pedicels, hyaline sepal margins, and anthers dorsifixed just above the base. However, Ardisia premontana is clearly separated from A. albovirens by the densely rufous furfuraceous lepidoce and angulate or subterete branchlets, the canaliculate, longer petioles, longer inflorescence, cupuliform calyx with carinate, glandular-granulose lobes with obtuse api- ces and glabrous subentire and erose margins, the shorter corolla irregu- larly divided with ovate, cucullate lobes, the filaments longer than the an- thers, the anthers apically acute, and the obnapiform ovary. Ardisia premontana is best placed in subgenus Ardisia, because of the longitudinally dehiscent anthers, pyramidally paniculate, terminal inflorescence with spi- cate branches, symmetric calyx lobes and truncate stigma. — Prpoty, New species of Ardisia (Myrsinaceae) 449 Ardisia zakii Pipoly, sp. nov. (Fig. 2) Quoad ramulos crassos usque ad 1 cm diametros, folia magna coriaceaque, petiolos usque ad 3 cm lon 7OS, A, carchianae simulans sed ab ea ramulis angulatis lon ritudinaliter ’ fa) 2 ~ alatis (non teretibus), foliis alternis (non pseudoverticillatis), petiolis teretibus ial ween es non canaliculatis), laminis desuper perpuncticulosis scrobiculatisque (non epunctatis laevibusque) subter manifeste minuteque atro-punctatis atque atro- cere laeiel (nec beak pallies punctatis), secus margines revolutis (nec planis), ad bas obtusis (nec acutis), inflorescentia rameali (non terminlai), secus rhachides epunctatas (non atro-punctato-lineatas), lobis calycinis depresso-ovatis auriculatisque (non lineari- lanceolatis symmetricisque), ovario pentangulo (non tereti), necnon in sylvas montanas nebulosasque (non premontanas) incolens, perfacile cognoscitur Tree 8 m tall. Branchlets sharply angulate, ca. 1 cm diam., with longi- tudinal wings along the angles, glabrous, the pith hollow. Leaves alter- nate; blades coriaceous, very widely elliptic, 28-31 cm long, 15-16.5 cm wide, apically widely rounded with a small acute tip, basally obtuse, decurrent on petiole to base, nitid and glabrous above and below, perpuncticulose and scrobiculate above, smooth, black punctate and punc- tate-lineate below, the midrib flat above, prominently raised below, sec- ondary veins 26-30 pairs, tertiary veins prominent above and below, margin entire, revolute, glabrous; petiole almost terete, prominently marginate, 2.5—3 cm long, glabrous. Inflorescence ramigerous, pyramidal, paniculate, 9-16 cm long, 7-10 cm wide; peduncle 3—4 cm long; rachis glandular-papillate, glabrescent; the branches spicate; inflorescence bracts unknown, floral bracts unknown; pedicel obsolete. F/owers 5-merous, co- riaceous 5.2—5.9 mm long; calyx cupuliform, 1.8—2 mm long, the tube ca. 0.6-0.8 mm long, the lobes asymmetric and auriculate, depressed- ovate, 1—-1.2 mm long, 1.2—1.4 mm wide, apically broadly rounded, ba- sally rugose, the margin scarious, minutely erose apically, sparsely eis dul Colats, glabrescent; corolla rotate, 5.2-5.9 mm long, tube 5-angled, 3—3.3 mm long, the lobes deltate, 2—2.6 mm long and wide, reflexed 180° at anthesis apically acute, densely but inconspicuously pel- lucid punctate-lineate, the margin irregular, Sate glabrous; stamens 3.5-4 mm long, the filaments flat, hyaline, 2.2-2.5 mm long, glabrous, the anthers lanceolate, 1—1.2 mm long, 0.8—1 mm wide, apically apicu- late, basally cordate, dehiscent by wide longitudinal slits; pistil lageniform, 5—5.3 mm long, the ovary ellipsoid, 5-angled, 3—3.5 m long, 1.5 mm diam., the style curved, 1.5 mm long, the ovules 5-8 in 2-3 rows, the stigma subcapitate, 3-lobed. Fruit globose, 5-7 mm diam., black at maturity, inconspicuously pellucid punctate. Types: ECUADOR. Picuincua: Carretera Quito-San Juan Chiriboga-Empalme, Km 59, 16 km NW of road, 1,700—2,000 m, 23 Sep 1986 (Al, fr), Vi Zak 1298 (HOLOTYPE: MO; tsorypes: BRIT, QCNE). 450 Stipa 17(2) FiG. 2. Ardista zakir Pipoly. A. Habit, showing winged branchlet. B. Detail of abaxial leaf surface, showing prominent tertiary veins. C. Portion of inflorescence, showing spicate branches. D. Flower bud, showing depressed-ovate calyx lobes and corolla lobes with hya- line margins. E. Corolla at anthesis, showing angled tube and reflexed lobes. F. Pistil, showing angled ovary and subcapitate stigma. G. Fruit. A-G, drawn from holotype. Prroty, New species of Ardisia (Myrsinaceae) 451 Distribution.—Known only from the type. Ecology and conservation status. —Ardisia zakii occurs in remnant cloud forests on steep slopes. With increasing forest destruction as roads are fur- ther developed, this species should be considered threatened. Etymology.—It is with great pleasure that I dedicate this species to Vlastimil Zak, former collector and participant in the Missouri Botanical Garden’s program of botanical exploration in Ecuador. Ardisia zakii is most closely related to Ardisia carchiana Lundell, but is easily recognized by the angulate, longitudinally alate branchlets, margin- ate, terete petioles, the leaf blades with obtuse bases, adaxially perpuncticulose and scrobiculate, abaxially black punctate and punctate- lineate, the margins revolute, the pentangular ovary and the depressed- ovate, auriculate calyx lobes. The auriculate calyx lobes and lanceolate, lon- gitudinally dehiscent anthers indicate placement in the group of Ardisza often segregated as Auricularidisia Lundell. The group has its center of diversity in Panama, with high concentrations of species also in Colombia. Ardisia flavida Pipoly, sp. nov. (Fig. 3) Ol | (eke as ae fOlia lamina d “fr perpuncticulosa costa canclculers petiolos neque ad 2 5 cm lonees: tanoresceacams pyramido- ncaa pedicelos obsoletos, calycem cupuliformem, lobos calycinos secus rgines erosos glabrosque, corollae lobos oblongos vel ellipticos, filamentia teretia, A. ae valde arcte affinis, sed ab ea ramulis glabris (non ferrugineo-stellato-tomentosis), petiolis subceretibus marginatisque (non canaliculatis), floribus flavidos (non eborinos), lobis calycinis 1-1.2 (non 0.8—1) mm longis minute rufo-lepidotis (nec rufo-puberulis) lobis ep carinatis (nec planis) ad apices planis (nec cucullatis) pellucido- (nec atro-)- punctato-lineatis secus margines crenulatis (nec erosis), filamentis glabris (non dense rufo- papillosis), antheris ovatis (non linearis) ad apices apiculatis (nec emarginatis), denique stigmate capitato (non puntiforme) statim separabilis. Shrub 4 m tall. Branchlets angulate, trigonal, 5-7 mm diam., glabrous. Leaves alternate; blades coriaceous, oblanceolate to elliptic, 19-25 cm long, 9—11.8 cm wide, apically widely rounded, shortly subacuminate, 3-5 mm long, basally obtuse, decurrent on the petiole, nitid and sparsely perpuncticulose above, pallid and inconspicuously black punctate below, midrib canaliculate above, prominently raised below, the secondary veins 25-36, margin entire, flat, scarious, glabrous; petiole subterete, margin- ate, 1.5—2.5 cm long, flat above, glabrous. Inflorescence terminal, pyramidal paniculate, 10-21 cm long, 8-16 cm wide at base; peduncle obsolete to 4 mm long; rachis sharply angulate, densely and minutely rubiginous glan- dular-granulose, the branches spicate; inflorescence, branch and floral bracts apparently early caducous, unknown, pedicels obsolete. nes ae ceous, yellow, 4—4.5 mm long; calyx deeply cupuliform, 3—3.4 mm long, unequally divided, the tube 1.3—1.5 mm long, the lobes eee asymmet- Sipa 17(2) dX A i) Fic. 3. Ardista flavida Pipoly. A. Habit, showing terminal inflorescence and angulate branchlet. B. Detail of abaxial leaf surface, showing inconspicuous black punctations. C. Flower cluster at tip of inflorescence rachis branchlect. D. Open corolla, showing minutely Pistil, crenulate margins, and apiculate anthers. E. Corolla lobe, showing medial ridge. F. showing placenta and apparently uniseriate ovules. A-F, drawn from holotype. Pipo.y, New species of Ardisia (Myrsinaceae) 453 ric, oblong, 2—2.2 mm long, 1.6—1.8 mm wide, apically obtuse to rounded, densely and prominently black punctate and punctate-lineate, densely and minutely rufous lepidote, hyaline, margin entire, scarious; corolla cam- 3—3.3 mm long, the tube 0.8—1 mm long, the lobes oblong to es panulate, elliptic, 2-2.3 mm long, 0.9-1.1 mm wide, apically acute, hyaline, medi- ally carinate, pellucid punctate and punctate-lineate, the margins minutely crenulate, glabrous; stamens 2—2.2 mm long, the filaments flat, 0.9-1.2 mm long, inserted at corolla base, hyaline, the anthers ovate, 1-1.3 mm long, 0.6-0.8 mm wide, subversatile, apically apiculate, basally cordate, dehiscent by large longitudinal slits, the connective epunctate; pistil obnapiform, 2—2.3 mm long, the ovary globose, 1—1.2 mm long and diam., the placenta depressed-globose, ovules 4, pluriseriate in a high spiral, thus appearing uniseriate, the style 0.8—1.1 mm long, the stigma capitate, 4- lobed. Fruit unknown. Type: ECUADOR. Esmeratpas: Canton San Lorenzo, 10 km $W of Lita, going up to the El Cristal Sector; 00°48' N, 78°30' W, 800 m, 10 Sep 1990 (fl bud), D. Rubio & C. QOuelal 664 (HOLOTYPE: MO; IsoryPEs: BRIT, QCNE). Distribution.—Known only from the type. Ecology and conservation status.—Ardisia flavida occurs at the limit of low- land forest with premontane forest. The forests of Esmeraldas form part of the Chocé Floristic Province, contiguous with the Cordillera Occidental of Colombia, an area that receives some of the highest rainfall known in tropical ecosystems. The coastal forests of Ecuador have very few old growth forest stands left, and thus, this species, along with its habitat, should be consid- ered endangered. Etymology.—The specific epithet refers to the yellow color of the flowers, a rarity among members of the genus. The chick, angulate branchlets and large, alternate leaves with perpuncticulose blades and canaliculate costa, the petioles up to 2.5 cm long, and pyramidal bipinnately paniculate terminal inflorescense, obso- lete pedicels, cupuliform calyx with lobes erose and glabrous along the margins, oblong to elliptic corolla lobes, and terete filaments, indicate that Ardisia flavida is most closely related to Ardisia monsalveae of Colombia. However, the glabrous branchlets, subterete petioles, yellow flowers, the longer, minutely rufous lepidote calyx lobes, the carinate, pellucid punc- tate-lineate corolla lobes with flat apices and crenulate margins, the gla- brous filaments, ovate, apiculate anthers and capitate stigma permit easy separation. The terminal panicles with spicate branches, and apiculate an- thers dehiscent by wide longitudinal slits indicates that Ardisia flavida 1s best placed in subgenus Ardisia, despite the fact that the style is not long and does not have a punctiform stigma. Clearly, the entire tribe is in great need of revision. 454 Sipa 17(2) Ardisia websterii Pipoly, sp. nov. (Fig. 4) Quoad ramulos stellato- tomentosos, filamenta longa, anthera lineari-lanceolata ad api- ces emarginata, necno n pedic A, monsalveae accedens, sed ab ea trunco uniaxiali (non multiaxial1), vestiment lino fe jue rufo- (non ferrugineo-) stellato-tomentoso, foliis pseudoverticillatis (non alternis), laminas ad bases semiauriculatis (non acutis), inflorescentia axillari (non terminali), lobis calycinis suborbicularibus (non ellipticis) ad apices late rocundatis (nec obtusis), lobis corollinis lineari-lanceolatis (non ellipticis) ad apices planis (nec cucullatis), filamentis glabris (non rufo-papillosis), denique stigmata capitata (non punctiforme) praeclare distinguitur Monoaxial tree (Corner’s Architectural Model sensv Hall et al. 1978) to 3 m tall. Branchlets terete when fresh, drying angular, 8-12 mm diam., densely and minutely rufous stellate-tomentose, the tomentum persistent. Leaves pseudoverticillate; blades chartaceous, widely ob] oe (48—)5 2—60 cm long, 17—22 cm wide, apically acuminate, the acumen 3—4 cm long, gradu- ally tapering to a truncate, semiauriculate base, aay slightly elevated but canaliculate above, prominently raised below; sordid and glabrescent above, pallid and sparsely stellate pubescent below, tomentose along the secondary veins, inconspicuously pellucid punctate and punctate-lineate, fea the margin roughly sinuate-dentate, the tooth vascularized; petiole canaliculate, obsolete to 5 mm long, glabrous above, densely stellate to- mentose below. [vflorescence lateral (axillary), at times produced in whorls between major whorls of hea, a columnar thyrse 9—15(—20) cm long, 3— 6 cm wide; peduncle 1.5—3 cm long; inflorescence bract foliaceous, mem- branaceous, oblong, 5 cm long, 1-1.2 cm wide, apically broadly rounded, gradually tapering to cuneate base, sparsely stellate tomentose above and below, densely and prominently black punctate and punctate- lineate, the margin entire, glabrous; secondary branch bracts foliaceous, membranaceous, oblong, 1.5—2.5 cm long, 0.4—-0.6 cm wide, apically rounded, basally cuneate, otherwise like inflorescence bract; floral bracts membranaceous, linear, 3.2—3.5 mm long, 0.8—1 mm wide, apically nar- rowly acute to subulate, cucullate, hyaline, densely and prominently black punctate, very sparsely stellate tomentose below, the margin entire, gla- brous; pedicel obsolete. F/owers S-merous, membranaceous, pale pink, 5—6 mm long, hyaline; calyx cupuliform, 2.5—3.5 mm long, irregularly divided, tube 1.4—1.7 mm long, the lobes suborbicular, |—1.8 mm long and wide, apically broadly rounded, densely punctate and punctate lineate medially, very sparsely lepidote, the margin hyaline, minutely erose, glabrous; co- rolla rotate, 5.5—6 mm long, the tube 2—2.4 mm long, the lobes linear lanceolate, 3.1—-3.4 mm long, 0.7—1 mm wide, apically acute, hyaline, gla- brous, the margin entire; stamens free, 6-6.6 mm long, exserted or anthers appearing versatile, the filaments free, 4.6—5 mm long, inserted at corolla base and adnate to 0.3 mm, hyaline, glabrous, the anthers linear-lanceolate, Prpoty, New species of Ardisia (Myrsinaceae) 455 we Nee ‘fa Fic. 4. Ardisia websterii Pipoly. A. Leaf, showing sinuate-dentate margin. B. Detail of abaxial leaf surface, showing stellate-comentum. C. Inflorescence, showing leaflike bract and co- lumnar thyrse morphology. D. Portion of inflorescence, showing floral bracts. E. Flower bud, showing cupuliform calyx and punctations. F. Corolla lobe and stamen, showing relative lengths of each. A—F, drawn from holotype. 456 Sipa 17(2) 2.1—2.8 mm long, apically narrowly acute to a minutely emarginate tip, seal deeply sagittate, dorsifixed ca. 1/3 from base, dehiscent pI large longitudinal slits, the connective epunctate. Pistil obturbinate, 4.8—5.5 mm long, the ovary subglobose, 1—1.2 mm long, 0.7—0.9 mm wide, densely black punctate, glabrous, the style tortuous, 3.5—3.8 mm long, the pla- centa subglobose, the ovules 3-5, appearing uniseriate, the stigma subcapitate, 3-4 lobed. Frait unknown. Type: ECUADOR. Picuincna: Canton Quito; Parroquia ae Maquipucuna Pro- tected Forest; 5 km airline SE of Nanegal, E side of rio Tulambi, 00°06! N, 78°37.5' W; 1,350-1,400 m, 5 Sep 1993 (Hl), G. Webster 30307 (HOLOTYPE: E es IsoTyPEs: DAV, QCNE). Pararypes: ECUADOR. Picuincna: Canton Quito; Parroquia Nanegal; Maquipucuna Protected Forest; 6 km airline SE of Nanegal, along trail from Rfo Umachaca to Loma Sta. Lucia, 00°07! N, 78°37' W; 1,675 m, 6 Sep 1993 (Al), G. Nee etal. 30361 (BRIT, DAV, QCNE), 5 km airline SE of Nanegal, above Rio Tulambi, 1,550 m, 00°7.5 ‘N, 78°38.5 'W, 1,550 m, 31 Aug. 1993 (A), G. Webster et al. 30009 (DAV, QCNE). Distribution.—A rdisia websterii appears to be endemic to the Maquipucuna Forest Reserve, Ecuador, at 1,350—1,675 m elevation. Ecology and conservation status. —This species is a conspicuous element of the montane rainforest understory, growing on streambanks, just above high water marks. The Maquipucuna Forest Reserve seems well-protected at this time, so Ardisia websteri? may be considered not under threat. Etymology.—This species is named for Grady L. Webster, curator of the John M. Tucker Herbarium of the University of California at Davis and pre-eminent authority on the systematics of neotropical Euphorbiaceae. Ardisia websterti with a monoaxial trunk and pseudoverticels of inflorescences, exhibits Corner’s Architectural Model (Hallé et al. 1978), At first glance, it has the aspect of several members of Cybianthus subgen- era Wegeltia and Comomyrsine, but the glabrous petals, long, free, exserted stamens, and anthers with deeply sagittate bases indicate placement in the genus Ardisia. While Ardisia websterii does not easily fit into any known subgenus of Ardisia, description of a new one is postponed until more neotropical species of the genus have been examined. Ardisia websterit ap- proaches A. monsalveae in several aspects, most notably because of the stel- late tomentum, long filaments, linear-lanceolate anthers with minutely emarginate tips and the obsolete pedicels. However, Ardisia. websterii is easily distinguished from A. monsalveae by the rufous vestiture, the pseudoverticillate, basally auriculate leaves, axillary inflorescence, subor- bicular calyx lobes with broadly rounded apices, the linear-lanceolate co- rolla lobes with flat apices, the glabrous filaments and capitate stigma. Clearly, more study is needed of the entire large-leaved group of Ardisia in the Andes. Prpoty, New species of Ardisia (Myrsinaceae) 457 ACKNOWLEDGMENTS The Missouri Botanical Garden’s exploration programs in Ecuador and Peru, headed by Dr. David Neill and Ing. Rodolfo Vasquez, respectively, continue to produce fine quality specimens from remote, poorly known and often dangerous areas. Their gifts sent to BRIT for determination made the present study possible, along with those of Dr. Grady Webster, of the University of California at Davis. The fine line illustrations were prepared y Ms. Linda Ellis with her usual knack for simplicity and accuracy. REFERENCES =. AUBLET, J. 1775. Histoire des plantes de la Guiane Francoise. Pierre-Francois Didot. Paris. CHEIH, CHEN and J. PipoLy. 1996. Myrsinaceae. Iv; Wu Zheng Yi and P. H. Raven, eds. Flora of China. Vol 15. Science Press, Beijing and Missouri Botanical Garden, St. Louis.[with Chen Cheih}. Pp. 1-38 Ducks, A. 1930. Plantes al o peu communes de la région amazonienne. Archiv. Jard. o ee 5:99-18 HALLE A. OLDEMAN, aa P.B. TOMLINSON. 1978. Tropical trees and forests. Springer- Velen. aes rk: LARSEN, K. and C.M. Hu. 1995. Reduction of Tetrardisia to Ardisia. Nord. J. Bot. 15:161— 162, LUNDELL, C.L. 1963. Studies of the American Myrsinaceae-I. Wrightia 3:77—90. 1964. Studies of the American Myrsinaceae-I]. Wrightia 3:97-114. .1981la. Neotropical Myrsinaceae-IV. Phytologia = a 1981b. Neotropical Myrsinaceae-V. Wrightia 7: . 1981c. Studies of American plants-X X. Phy ee 48: | 31-136. 1981d. Neotropical Myrsinaceae-VI. Pl ONE Ie 49: 341-354. 1982. Neotropical Myrsinaceae-VIU. Wrightia 7:38—50. Miter, J. saat. Pipoty. 1993. A new species of Ardisia (iisinacere) from Madagascar. Novon 3:63— Pipoty, J. 199 1a. rere gees a new species of Myrsinaceae from Panama. Ann. Missouri cle Gard. 78:5 pee ee Auta ines i género Ardisia Swartz (Myrsinaceae) en Colombia. Caldasia irs . 284 ek eee eee pes ieee oo a new species from the Antioquian Choc6 of an Novo 9: «21:99 4a. New species a e. (Myrsinaceae) from the Cordillera Occiden- tal of Colombia ae Ecuador. Novon 4:38—44 —__. :1994b. Further notes on the genus Ardisia (Myrsinaceae) in Madagascar. Sida 16:361—364. «1995. Dos nuevas especies del aa es alana de la provincia floristica aceana de Colombia. Caldasia 17(82-85):419— STONE, B. 1982. New and noteworthy Malesian My rsinaceae, - Malaysian. For. 45:100- 121 1989. New and pie aes Malesian Myrsinaceae, III. On the genus Ardisia in Borneo. Dice. Acad. Nat. Sci. Phila. 141:263-306 —_____.. 1990. Studies in Malesian Myrsinaceae, V. Additional new species of Ardisia Sw. Proc. Acad. Nat. Sci. Phila. 142:21-58. 458 Stipa 17(2) «992. A revision of the genus Tetrardisia Mez (Myrsinaceae). Malayan Nat. J. 461-11. 1993a. Reduction of the genus Parardisia (Myrsinaceae). Nord. J. Bot. 13:55— pee 993b. New and noteworthy Malesian Myrsinaceae, VI. Scherantha, a new subgenus of Ardisia. Pacific Sci. 47:276-294 Taron, A. 1979. Saree a etude du ae Ardisia Sw. (Myrsinaceae) en Afrique tropicale. Bull. Jard. Bot. Nat. Belg. 49:81-12 Ursgan, I. 1922. ae antillanum IV. Repert. ae Nov. Regni Veg. 18:22. NEW SPECIES OF GEISSANTHUS (MY RSINACEAE) FROM THE HYLAEA/ANDEAN INTERFACE OF ECUADOR AND PERU JOHN J. PIPOLY II Botanical Research Institute of Texas 509 Pecan Street Fr. Worth, TX 76102-4060 U.S.A. ppipoly @ brit. org ABSTRACT Routine determination of — received as gift for determination aes in the discovery of three novelties in Geéssanthus. Geissanthus vanderwerffi Pipoly, G. spectabilis Pipoly and G. ae Pipoly are described, illustrated and their eee rela- tionships are discussed. RESUMEN Al determinar muestras recibidas como cepales pe su determinacién, se encontraron a tres especies nuevas, By perteoceien ntes a Se describe, se ilustra y se discute el parentesco de Gerssanthi wun Pipoly, @ Gece bale Pipoly y G. challuayacus Pipoly. INTRODUCTION The genus Geissanthus was circumscribed by Hooker (Bentham & Hooker 1876) to include 10 taxa from the Andes, which he did not list. The only reference to a taxon was made indirectly, by listing Fendler 759, a specimen of Geissanthus fragrans Mez. Mez (1902) distinguished Geissanthus from the other genera of the tribe Myrsineae by its free stamens, dorsifixed anthers, and the calyx closed in bud, opening later into irregular lobes. In that treatment, 25 species were recognized, of which 17 were new. We must assume, therefore, that at least eight of the taxa described as new by Mez had actually been seen previously by Bentham, but remained undescribed. Subsequently, miscellaneous new species were added to the genus by Mez (1905, 1920), MacBride (1934) and Cuatrecasas (1951). Agostini (1970) was the first worker since Mez to discuss generic de- limitation among taxa assigned to Conomorpha A. DC., and Stylogyne A. DC. Agostini distinguished Gesssanthus from Stylogyne and what was at that time the Cybzanthus Martius complex of genera (including Conomorpha A. DC., Weigeltia A. DC., Comomyrsine Hooker) based on a combination of several characters, including: the terminal inflorescence; sessile or subsessile flowers; calyx lobes closed in bud, then rupturing into 2-8 unequal lobes Sipa 17(2): 459-470. 1996 460 SIDA 17(2) on the same inflorescence; corolla lobes fused 1/3 or more of their length; and a capitate stigma. Using those criteria, he transferred three taxa from Conomorpha, and one taxon from Stylogyne, to Geissanthus. My previous studies (Pipoly 1993) and the present one reconfirm that Geissanthus may best be defined by its unique calyx, closed in bud and opening into 2-8 usually unequal lobes, the corolla with linear, oblong or rarely ovate lobes that are distally recurved at least 180° in anthesis, and the subversatile or versatile anthers which are latrorsely dehiscent by wide or narrow longitudinal slits. Sty/ogyve may easily be separated from Geissanthus by its contorted corolla, with the lobe tips highly twisted in bud. Cybianthus (including Comomyrsine Hooker, Conomorpha A.DC., Correlliana D'Arcy, Grammadenia Bentham, Microconomorpha (Mez) Lundell, and Weigeltia A. DC., see Pipoly 1987, 1992) may be separated from Geissanthus by its axillary inflorescences, stamens connate by their filaments to form a tube, the staminal tube adnate to the corolla tube, and the glan- dular granules present at least at the junction of the corolla lobes and tube. Using these criteria to determine generic limits, Pipoly (1993) transferred three species into Gerssanthus and described an additional four species. Those seven taxa, and the three novelties described here, bring the total number of species recognized in Gezssanthus to 45. Because I disagree with the place- ment of Stylogyne ambigua (Martius) Mez in Gerssanthus (Agostini 1970), the genus is entirely Andean except for populations of Gesssanthus perpuncticulosus (Lundell) Pipoly occurring in the Darién of Panama. Mem- bers of the genus are mostly known from moist, wet and pluvial premontane, montane, and cloud forests, and less frequently from subpaéramo thicket formations. They are moderate trees, usually less than 15 m tall, with very conspicuous terminal, paniculate inflorescences. Quantitative studies of Andean forests I have conducted in Colombia indicate that they generally occur in populations smaller than 20 individuals per hectare, at least in the mid-level stratum of the montane and cloud forests. My qualitative obser- vations from Ecuador and Peru have thus far confirmed the statistics ob- served in Colombia. Determination of gift specimens received from the Missouri Botanical Garden, in connection with their floristic studies in Ecuador and Peru, resulted in the discovery of the following novelties, described herewith. Morphological terminology follows Pipoly (1993) and Lindley (1848). Geissanthus vanderwerffii Pipoly, sp. nov. (Fig. 1) Quoad ramulos angulatos glabrosque, folia alterna, petiolos marginatos, lamina secus margines integerrima, inflorescentiam folias longitudine aequantiam vel subaequantiam, corollam campanulatam, necnon lobos corollinos retrocurvatos, G. ecwadorensi valde arcte affinis, sed ab ea laminis cortaceis (non carcilagineis) desuper scrobiculatis (nec laevibus) Pipoty, New species of Geissanthus (Myrsinaceae) ~ Ny ra . ~ — tr. a ro ag as oy acs é OW Rr] he i il x ‘o Seg — gh \, = Es > us vanderwerffii Pipoly. Habit, showing the angled branchlets and pedicel- Fic. 1. Gesssanth baxial leaf surface detail, showing the dense and minute pellucid puncta- bt late flowers. B. A tions. C. Flower with corolla removed, showing the pedicel, and the cupuliform calyx with entire lobes. D. Young fruit. E. Opened corolla, showing the apiculate anthers with latrorse slits. A-C, E, drawn from holotype; D, drawn from vanderwerff 8965. 462 Stipa 17(2) subcer glabris (nec lepidocis) dense minuteque pellucido-punctatis (nec epunctatis), inflorescentiis polygamis (non bisexualis), pedicelis 3-5.6 mm longis (non obsoletis), flores —9 (non 3—5.6) mm longis, calyce cupuliformi (non cotyliformi) in fructo spissescenti (nec in statu codem reliquo), lobis ee ovatis vel deltatis (non suborbicularibus) rufo- papillosis (nec glabris) 2.5—3.5 (nec 1.8—2.2) mm longis secus margines opacis integrisque (nec hyalinis erosisque), lobis corollinis ad apices anguste vel late acutis (non rotundatis), denique antheris ad apicem acutis (non rotundatis) perfacile cognoscenda. Tree to 6 m tall. Branchlets angulate, 4.5—10 mm diam., glabrous. Leaves alternate; blades coriaceous, widely oblong to obovate, (9.8—)11—15.5(— 18.2) cm long, (4.4—)6-7.2(-8.5) cm wide, apically obtuse, widely rounded or rarely truncate, basally obtuse to truncate, decurrent on the petiole, dull, glabrous, minutely scrobiculate above, densely and minutely pellucid punc- tate below, the midrib flat to slightly impressed above, prominently raised below, the secondary veins mere pairs, the margin entire, revolute, gla- brous; petiole marginate, 1.8—2.5 cm long, glabrous. Izflorescence terminal, pyramidal, bipinnately or mixed paniculate, (7.5—)9-12 cm long, 8-15 cm wide at base, polygamous, the Howers staminate and bisexual, mono- morphic; peduncle 0.5—1 cm, anglulate, glabrous or very sparsely and mi- nutely rufous papillate, the branches with flowers corymbose; inflorescence, branch and floral bracts apparently early caducous, unknown; pedicels cy- lindric, 3—5.6 mm long, sparsely pellucid punctate-lineate, glabrous or very sparsely and minutely papillate. F/owers chartaceous, white, 7—9 mm long; calyx deeply cupuliform, 5.5-7 mm long, 3-6 lobed, irregularly di- vided, the tube 2.8-4.9 mm long, sparsely rufous papillate, pellucid or orange punctate and punctate- linear the lobes widely ovate to deltate, 2.5—-3.5 mm long, 2.1—3.5 mm wide, apically narrowly to broadly acute, the margin entire, opaque, glabrous; fruiting calyx incrassate obconic, 6.6— 6.5 mm long, 6—6.3 mm diam.; corolla campanulate, 7—9 mm long, 5- lobed, the tube 3.5—4.2 mm long, the lobes, ovate, 3.5—4.8 mm long, 2.8— 3.5 mm wide, apically acute, reflexed 180° at maturity, densely and promineatly orange or pellucid punctate, glabrous, the margin entire, slightly involute, glabrous; stamens 5 in staminate flowers, 7.5—8 mm long, the filaments free, 6-6.5 mm long, subterete, the anthers ovate, 1.8—2.5 mm long, 0.9-1.4 mm wide, apically obtuse with prominent apiculum, basally deeply cordate, dehiscent by wide latrorse longitudinal slits, rhe connective epunctate; stamens of bisexual flowers like those of staminate but 4.2—4.5 mm long, the filaments free, 2.1—2.4 mm long, widened ba- sally 1-1.2 mm, flat, adnate 0.8—1 mm above corolla tube basae, translu- cent, epunctate, glabrous, the anthers 2.1—-2.4 mm long, 1.4 —1.7 mm wide; pistil of bisexual flowers obnapiform, 5.3—-5.8 mm long, the ovary 2.6-3 mm long, 3—3.5 mm diam., densely pellucid punctate, glabrous, the placenta depressed-globose, the ovules 3—5, uniseriate; the style 2 om — 463 ~~ Pipoty, New species of Geissanthus (Myrsinaceae 2.7 mm long, pellucid punctate-lineate, glabrous, the stigma capitate, 4- lobed; pistillode similar to pistil but 3.5—4 mm long, the ovary 1.2—1.5 mm long, the ovules obsolete, the style 2.3—2.9 mm long, the stigma capitate, unlobed. Frat globose, 6-8 mm diam., inconspicuously pellucid punctate. Type: ECUADOR. Zamora CHINCHIPE: along old trail from Nudo de Sabanilla to eas st m, 6 May 1987 (bisexual fl), H. van der Werff & W. Palacios 9374 (HOLO- type: US; ypes: BRIT, MO, QCNE). sews pes: ECUADOR. ZaMORA-CHINCHIPE: Canton Zamora; Parque Nacion eae: near el Tambo, 40 km NW of Zamora on road to Loja, 03°58'S, 79°C “a 2,210 m, 13 July 1993 (fr), A. Gentry 79920 (MO, QCNE); Yangana-Valladolid Rd., km 21, oo 79°09'W, 2.650—2,750 m, 2 Dec 1988 (stam. fl), J. Madsen, C. Bloch & H. Christensen 75794 (AAU, BRIT, QCA). Distribution.—Endemic to the south-central Andes of Ecuador, in open elfin forest or scrub, at 2,200—2,750 m elevation. Ecology and conservation status. —Getssanthus vanderwerffii forms a conspicu- ous element of the cloud forest vegetation, even though it is a component of the understory (van der Werff, pers. comm.). Because sizeable popula- tions of this species occur in a national park located in a somewhat remote area, it is not currently under threat. Etymology.—It is with great pleasure that I dedicate this spectacular new species to Henk van der Werff, head of the Taxonomy Department at the Missouri Botanical Garden. Henk is a prodigious, indefatigable field worker and pre-eminent authority on the systematics of Neotropical Lauraceae. The angled branchlets, alternate leaves with entire blades, marginate petioles, the inflorescence equalling or subequalling the leaf length, and the campanulate corolla indicate that Gerssanthus vanderwerffii is most closely related to Geissanthus ecuadorensis Mez. However, the coriaceious leaf blades scrobiculate above, densely and minutely pellucid punctate and glabrous below, the polygamous inflorescence with longer pedicels and larger flowers, the cupuliform calyx incrassate in fruit, and with ovate to deltate, rufous- papillate lobes that are entire and opaque along the margins permit easy recognition of G. vanderwerffii. In addition, the narrowly to widely acute corolla lobe apices are distinctive. The incrassate (thickening with matu- rity) fruiting calyx is unique within the genus. Geissanthus spectabilis Pipoly, sp. nov. (Fig. 2) ropter ramulos ad apices angulatos furfuraceo-lepidotos, folia pseudoverticillata, lamina chartacea anguste vel ae »blanceolata ad apices acuminata, secus margines integerrima plana, a labraque, petiolos canaliculatos desuper glabros uate us emeea lepidotos, inflorescentia terminalia pyramidalia, ores membranaceos, lol | apices acutos secus eee ntegros glabrosque, corolla campanulata, necnon aoe corollinos lineares G. longistamineo valde arcte affinis, sed ab ea ramulis pentangularibus (nec irregulariter angulatis mature teretibus) sine granulis glanduares (nec glandulari-granulosis), laminis = 2 464 Sipa 17(2) lcm 1 ———$——__ ? eae P ~ a —," i‘ Ya "Os - —T ~ Ki YA) i) ; +. j SS ty \ Cis 5 cm \ <= y, B N sae: Fic. 2. Gerssanthus spectabilis Pipoly. A. Habit, showing the angulate branchlets and pseudoverticillate leaves. B. Abaxial leaf surface detail, showing scattered rufous furfura- ceous lepidote scales. C. Flower, showing acute calyx lobes and linear corolla lobes. D. Dissected corolla lobe and attached stamen, showing prominent anther apiculum. E. Fruit. A-D, drawn from holotype; E, drawn from Smith 4172. Pipoty, New species of Geissanthus (Myrsinaceae) 465 ad bases acutis (non abrupte truncatis) subter obscure pellucido-punctatis punctato- Hncaesd ue (nec Sseaeaas atro punctato-lineatis), panicula tri- (non bi-) pinnata, pedicels soletis (non obconicis 0.2—1.3mm sal egy floribus 3.5—4.5 (non 5.8—6.3) mm longis, ais mentis ants (non ee 2.3 | 4—4.5) mm longisque, antheris parce dorsifixis (non versatilis) lanceolatis (nec sbleag) 1—1.2 (nec 2.2—2.4) mm longis, ad apices apiculatis (nec obtusis) denique pistillo obnapiformi a obturbinato) statim separabilis. Tree to 6 m tall. Branchlets angulate, 7-20 mm diam., pentagonal in cross section, prominently ridged along the angles, densely and minutely ferrugineous furfuraceous-lepidote, glabrescent. Leaves pseudoverticillate; blades chartaceous, narrowly to widely oblanceolate, (22.5—)29.4—36.5(— 41) cm long, (7.5—)8-11.5 cm wide, apically short acuminate, the acumen 1—2 cm long, basally acute, decurrent on the petiole, drying dull and smooth above, pallid, inconspicuously pellucid punctate and punctate-lineate as well as sparsely ferrugineous furfuraceous-lepidote below, the midrib im- pressed above, prominently raised below, the secondary veins 20—31 pairs, the margin entire, flat, glabrous; petiole canaliculate with a slight margin from decurrent leaf base, 1—1.5 cm long, glabrous above, sparsely furfura- ceous-lepidote below. Inflorescence terminal, pyramidal tripinnately panicu- late, (20—)24—38(—44.5) cm long, (1 3—)24—33 cm wide at basally, entirely bisexual; peduncle 0.5—1 cm long, the rachis angulate, densely and mi- nutely furfuraceous-lepidote, glabrescent, the branches spicate; inflorescense, branch and floral bracts apparently ree caducous, unknown; pedicels ob- solete. F/owers membranaceous, 3.5—4.5 mm long; calyx deeply seek 2.5—3.5 mm long, (3—)4—5-lobed eoalate divided, tube 1—2.5 mm long, ne ovate to deltate, 1-1.3 mm long, 0.6-1 mm wide, apically acute, glabrous or sparsely lepidote, densely and prominently black punctate, the margin entire, opaque, glabrous; corolla campanulate, 3.5—4.5 mm long, S-lobed, the tube 1—1.3 mm long, the lobes reflexed 180° at maturity, linear, 2.5—3.5 mm long, apically. subulate: hyaline, epunctate, glabrous, the margin entire, flat; stamens 2.8—3.3 mm long, the filaments free, flat, epunctate, glabrous, 2—2.3 mm long, adnate to corolla tube 0.8—1 mm, widened at basally, the anthers lanceolate, 1-1.2 mm long, 0.4—0.5 mm wide, apically rounded with a prominent apiculum, basally rounded, slightly dorsifixed, dehiscent by wide latrorse slits, connective epunctate; pistil obnapiform or globose, 1.5—2 mm long, the ovary globose, ca. | mm long and diam., densely black punctate and punctate-lineate, style slender, 0.5— 1 mm long, the stigma capitate, 4-lobed, placenta globose, ovules uniseriate, 4, Fruit globose, black at maturity, 5-7 mm diam. when dried, pellucid or red punctate. Type: PERU. Junin: Prov. Chanchamayo, Fondo Romero, Pampatigre, above Sta. Ana (SE of La Merced), 11°00'S, 75° 10'W, 1,500—1,700 m, 7 Mar 1985 (bisex. fl, fr), B. Stern & C. Todzia 2336 (HoLtoryee: MO; tsorype: USM). 466 Stipa 17(2) PARATYPES: PERU. Pasco: Pepe fe along road between Oxapampa and Villa Rica, Km 7 SE from Miraflores crest, 10°37'S, 75°20'W, 11 Oct 1982 (fr), R. Foster 9145 (EB, MO, US, USM); 5 km SE of Oxapampa, Oswaldo Miiller property, LO0°36'S, 75°23'W, 1,850 m, 31 Jan 1983 (fl, fr), D. N. Smith 3166 (MO, US, USM), 25 May 1983 (fr), D. N. Smith 4172 (MO, US, USM). Distribution.—Getssanthus spectabilis on the eastern slopes of the Peruvian Andes, in the Departments of Pasco and adjacent Junin, 1,500—2,400 m elevation. Ecology and conservation status. —Geissanthus spectabilis grows in tall “ceja de selva” forest, the cloud forest region of Andean Peru that faces Amazonia, The region is known for its high endemism (Gentry 1993) and contains approximately 57% of the total vascular plant species known for the coun- try. Gerssanthus spectabilis is a conspicuous element of the forest, although locally common. The late D. Smith indicated (Smith, pers. comm.) that this tree is common at the junction of the taller “ceja” with that of the shorter portion, especially at the margin of the forest along watercourses in small ravines. Given the restricted distribution it is probable that this spe- cies is threatened. Etymology.—The specific epithet describes the aesthetically appealing aspect of the plant. ae spectabilis 1s closely related to G. longistamineus (A. C. Smith) Pipoly by virtue of its angulate, furfuraceous lepidote branchlets, pseudoverticillate, chartaceous, narrowly to widely oblanceolate leaves with acuminate apices and entire, flat margins, the terminal and pyramidal inflorescences, the chartaceous flowers with calyx lobes acute, entire and glabrous along the margin, and finally, the corolla campanulate with linear lobes. However, Geissanthus spectabilis may immediately be separated by its 5-angled stems, leaf blades with acute bases, obscurely pellucid punctate and punctate-lineate below, the tripinnate panicles, shorter, sessile flowers with flat, shorter filaments, che shorter anthers slightly dorsifixed, lanceolate, and apiculate, and finally the obnapiform pistil. Geissanthus challuayacus Pipoly, sp. nov. (Fig. 3) Species haec ob lamina chartacea subter rufo-furfuraceo-lepidota, inflorescentia bisexuales, flores membranaceos, corolla campanulata, anthera lanceolata, G. francoam affinis, sed ab tls), ea ramulis aliquantum teretibus (non quadrangularis) glabris (non furfuraceo-lepic laminis secus margines integris (nec serratis), subter aac lepidotis dissite praeditis (non superpositus sic a velvetem similis) indutis, petiolis 1.7—2.5 (non (2.2—)2.5—3) cm longis, corolla 4—4.5 (non 3-3.2) mm longis, lobis sae oblongis (nec linearis), staminibus 4—4.2 (non 2.1—2.2) mm longis, denique stigmate 4-lobato (nec puntiforme) praeclare dignoscenda. Tree to 10 m tall. Branchlets somewhat angular, almost terete, 5-9 mm diam., glabrous. Leaves alternate; leaf blades chartaceous, oblanceolate, Pipoty, New species of Geissanthus (Myrsinaceae) 467 5 oma fll Fic. 3. Geissanthus challuayacus Pipoly. A. Habit, showing the almost terete branchlets. B. Abaxial leaf surface detail, showing sparse and minute lepidote scales. C. Inflorescence branchlet tip, showing obsolete pedicels. D. Partially opened Hower bud, showing asymmetric rup- turing of calyx. E. Dissected corolla lobe and stamen, showing flat filament adnate to corolla lobe. F. Pistil, showing capitate, lobed stigma. G. Longitudinal section of ovary, showing placenta and ovules. A, drawn from Hurtado 1787, B—G, drawn from holotype. 468 Stipa 17(2) (16.8—)22—28 cm long, (4.7-)7-9(-11.8) cm wide, apically cuspidate- acuminate, the acumen 1—1.5 cm long, basally obtuse or rarely acute, de- current on the petiole, dull and glabrous above, pallid and very sparsely and minutely rufous furfuraceous lepidote below, the midrib canaliculate above, prominently raised below, the secondary veins 16—26 pairs, essen- tially flat above, slightly raised below, margin entire, flat; petiole margin- ate, 1.7—2.5 cm long, glabrous. Inflorescence terminal, pyramidal bipinnately paniculate, 11.5-20 cm long, 12-28 cm wide at basally, bisexual, the branches spicate; peduncle subsessile, 3-5 mm long, glabrous; inflorescence, branch and floral bracts apparently early caducous, unknown; pedicels ob- solete. F/owers membranaceous, greenish-white, 4—4.5 mm long; calyx deeply cupuliform, 3.2-3.5 mm long, hyaline,3—5-lobed, irregularly di- vided, the tube 1.8—2.5 mm long, angular, the lobes ovate, 1.4—1.6 mm long, 1.2-1.4 mm wide, apically acute, densely and prominently black punctate and punctate-lineate, the margin irregular, entire, glabrous; co- rolla campanulate, 4—4.5 mm long,4—5-lobed, hyaline, regularly divided but highly variable, the tube 1.2—2.5 mm long, the lobes oblong, 2—3 mm long, 1-1.2 mm wide, apically narrowly acute to subacuminate, cucullate, densely black punctate and punctate-lineate apically, the margin regular, entire, glabrous; stamens 4—4.2 mm long; the filaments free, flat, 2.5— mm long, hyaline, adnate to corolla tube 0.5 mm from base, the anthers lanceolate, 1.3—1.5 mm long, 0.3—0.5 mm wide, apically acute, at times with a minute apiculum, basally deeply cordate, dorsifixed ca. 1/3 from base, dehiscent by wide latrorse slits, the connective epunctate; pistil obturbinate, 2.2-3 mm long, 0.7—1 mm diam., the ovary globose, 1 mm long, néllweid punctate, glabrous, |—1.2 mm ica the placenta subglobose, the ovules 4, uniseriate, exposed, the sty — e truncate, 1.5—2 mm long, the stigma capitate, 4-lobed. Fruit subglobose, 5-7 mm diam., glabrous, pel- lucid punctate. TYPE: cg a Napo: Canton Archidona; § slopes of Volcan Sumaco, Hollfn-Loreto Rd, Km 1 Challua Yacu Village, 00°43'S, 77°40'W/, 1,200 m, 20-25 Mar 1989 (bisex. fl), W eee 4006 (HOLOTYPE, BRIT; tsorypes, MO, QCNE, US). Pararyprs. ECUADOR. Napo: Cantén Archidona; $ slopes of Volcdn Sumaco, Hollin- Loreto Rd, Km 25, Challua Yacu Center, 00°43'S, 77°40'W, 1,230 m, 10-19 Nov 1989 (fr), E Hurtado & A. Alarado 891 (MO, QCNE); 00°43'S, 77°36'W, 1,200 m, 20 Mar 1989 (fl), EF Hurtado & X. Rufz 1787 (QCNE, MO, US); Hollin-Loreto Rd, 5 km W of Guamant, slopes of Volcan Sumaco, 00°43°S, 77°38'W, 1,200 m, 6-7 Sep 1988 (fr), D. Neill et al. 8574 (MO, QCNE, US) Distribution.—Thus far, known only from the Volcan Sumaco area, along the Hollin-Loreto Rd., Province of Napo, Ecuador, from 1,200—1,230 m elevation. Ecology and conservation status. —Geissanthus challuayacuus grows in lower Pripoty, New species of Geissanthus (Myrsinaceae) 469 montane forest, on the eastern slopes of the Andes, facing Amazonia. It 1s presumed that, given the forest destruction present in the Napo region of Ecuador, this species may be considered threatened. Etymology.—The specific epithet is the genitive form of the village name, Challua Yacu, Napo, Ecuador, and is thus declined as a Latin noun of the Fourth Declension. The chartaceous leaves, furfuraceous lepidote below, bisexual inflorescences, membranous flowers with campanulate corollas and lanceolate anthers indicate that Geissanthus challuayacuus is most closely related to G, francoae Pipoly, but may be separated by the somewhat terete, glabrous branchlets, leaf blades with entire margins, sparsely lepidote undersurface, shorter petioles, longer corolla with oblong lobes, longer stamens, and 4- lobed stigma. ACKNOWLEDGMENTS Specimens for the present study was provided by the Missouri Botanical Garden as gift or loan for determination. Material from Peru was collected through grants from the Andrew W. Mellon Foundation, that also subsi- dized the excellent illustrations so skillfully executed by Ms. Linda Ellis. I thank Jon Ricketson (MO) and B.L. Turner (TEX) for reviews of the manuscript. REFERENCES Acostini, G. 1970. Notes on Myrsinaceae. I. Generic assignment of Conomorpha sodtroana Mez, Ardisia ambigua Mart., and related species. Phytologia 20:401—403 Hox, J.D. 1876. Myrsineae. In: G. ae & J. D. Hooker, eds. Gunes Plantarum. yell Reeve & Co. London. Pp. 639-65( pe J. 1951. Notas a la flora de iene XI. Rev. Acad. Colomb. Ci. Ex. 8:297-328. Gentry, A. 1993. Overview of the Peruvian flora. Pages xxix—xl. In: L. Brako & J. Zarucchi (editors). Catalogue . the Flowering Plants and Gymnosperms of Peru. Missouri Botanical Garden. St. , Missouri. LInbLey, J. 1848. Theos caer of botanical terms. {[Exerpt from Hlustrated Dic- tionary of Botanical Terms, by J. Lindley, With an Introduction by A. Eastwood. 1964, School of Earth ee ay ee Univ., Stanford, California. ] Macsripe, J.F. 1934. New or renamed spermatophytes, mostly Peruvian. Candollea 5:346-402. Mez, C. 1902. Myrsinaceae In: A. Engler, ed. Das Pflanzenreich. 9(V, 236):1—437. Leipzig. Verlag von Wilhelm Englemann. «1905. Additamenta monographica. IH. Bull. Herb. Boissier sér. 2, 5:527-538 «1920. Additamenta monographica. HI. Repert. Spec. Nov. Regni. Veg. 16:410-425. Pipoty, J. II]. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenta (Myrsinaceae). Mem. New York Bot. Gard. 43:1—76. 470 Stipa 17(2) — «1992. The genus Cybianthus su Ann. Missouri Bot. Gard. 79:908—957. —_______.. 1993. The genus Gesssanuthus (Myrsinaceae) in the Chocé Floristic Province. Novon 3:463—474. dgenus Conomorpha (Myrsi in Guayana. A NEW SPECIES OF CAREX (CYPERACEAE) FROM URUGUAY AND A NEW NAME IN THE GENUS GERALD A. WHEELER Department of Plant Biology University of Minnesota . Paul, MN 55108-1095, U.S.A. w ™ ABSTRACT Carex herteri (sect. Digitatae) is newly described from the Department of Florida in south- ern Uruguay and is known only from the holotype. This very distinctive, but diminutive species is characterized by having large, glabrous perigynia that are distinctly beaked, few perigynia per spike, large achenes that are strongly apiculate, and ciliate leaf margins. Additionally, the South American plants previously called C. phalaroides var. crassiflora (sect. Digitatae) are here raised to species rank. The name C. gibertir is offered for this species and a lectotype is selected. RESUMEN Se describe Carex herters (sect. Digitatae) del Depart to de Florida en sur de Uruguay y slo se conoce el holotipo. Esta especie, muy diferente aunque diminuta, se caracteriza por tener utriculos grandes y glabros, con un pico distinto, pocos utriculos por espiga, aquenios grandes fuertemente apiculados, y margenes foliares ciliados. Adicionalmente, as plantas sudamericanas denominadas previamente C. phalaroides var. crassiflora (sect. Digitatae) se elevan aqui al rango especffico. Se propone para esta especie el nombre C. gibertit y se selecciona un lectotipo. NTRODUCTION It is estimated that over 200 species of Carex L. (Cyperaceae) occur in South America (Wheeler 1996). Of those growing in the southern half of the continent, slightly under 20 have been reported from Uruguay (Osten 1931; Chebataroff 1942; Herter 1953; Pedersen 1969; Wheeler 1987, 1988). In this paper, a new species of Carex is reported from the Depart- ment of Florida in southern Uruguay. In addition, a new name is offered here for plants previously called C. phalaroides Kunth var. crassiflora Kiik., which occur in southeastern South America. DESCRIPTIONS AND COMMENTS Carex herteri G. A. Wheeler, sp. nov. (Fig. 1) Herbae laxae caespitosae, parvae; culmi ca. 2 cm alti; vaginae basales glabrae, brunneae. Folia ca. 7, plerumque b asilaria, superare culmi; laminae ca. 1.5—4 cm longae, 0.8— latae, marginibus ciliatis; ligulae 0.5—1 mm longae. Inflorescentiae ca. 1.5 cm longae; 3 spicae ca. 3, ice 3—4-florae, inferiore breve pedunculatae. Perigynia 4.4—4.8 mm Sipa 17(2): 471-477. 1996 aN 72 SIDA 17(2) Fic. 1. Carex herter’, habit with achene and perigynium to the right and far right of the plant, respectively (plant from the Department of Florida, Uruguay), from Hb, Osten 19091 (S partim), holotype. gar = lcm. longa, 2—2.3 mm lata, acriter trigona, glabra, pallide viridis marginibus levis; rostra 0.8— 1.2 mm longa. Achenium 3.3—3.5 mm longum, |.8—2 mm latum, acriter trigoni, brunnei cum flavis anguli. Stigmata 3. Plants loosely cespitose from short rhizomes, low-growing. Fertile culms ca. 2 cm tall, with glabrous, brownish basal sheaths. Leaves ca. 7, basal, exceeding the culms; blades ca. 1.5—4 cm long, 0.8—2 mm wide, flat or channelled proximally, the margins ciliate (but only sparingly so near the apex); leaf sheaths very short, glabrous; inner band of leaf sheaths hyaline or pale brown, glabrous; ligules 0.5—l mm long, rounded. Inflorescences ca. 1.5 cm long, the terminal spike slightly larger than the lateral ones, WHEELER, Carex (Cyperaceae) from Uruguay 473 spikes strongly overlapping; lowermost bract scale-like, 3.5—5 mm long, ca. 2.5 mm wide, sheathless, with a ciliate awn ca. 0.8 mm wide and up to 2.5 cm long, the upper ones somewhat reduced. Spikes ca. 3, androgynous. Terminal spikes ca. 1 cm long; staminate portion ca. 8 mm long and 2.5 mm wide, ca. 7-flowered; pistillate portion ca. 6 mm long and 4.5 mm wide, ca. 3-flowered. Lateral spikes 5-7 mm long, on smooth peduncles up to 4 mm long; staminate portion inconspicuous, ca. 2—3-flowered; pistil- late portion ca. 4-6 mm long and 4—5 mm wide, ca. 3—4-flowered. Pistil- late scales exceeding the perigynia (at least the prolonged tips), the bodies 3.6—5 mm long, 1.8—2.6 mm wide, ovate-lanceolate, glabrous, pale green or stramineous center with broad, white hyaline or pale brown margins, 3- veined, the midvein prolonged into a cuspidate or sparingly aristate tip up to ca. 12 mm long. Staminate scales with bodies 3.4—4.2 mm long and 1.3-1.9 mm wide, ovate-lanceolate, glabrous, pale green or stramineous center with broad, white hyaline or pale brown margins, 3-veined, the midvein prolonged into a cuspidate or sparingly aristate tip up to 4 mm long. Perigynia 4.4-4.8 mm long, 2—2.3 mm wide, erect or somewhat spreading, strongly trigonous with slightly concave, elliptical to rhombic sides, glabrous, pale green or stramineous, 2 distinct veins and the rest obscure (however, several short veins are usually visible near the base), not inflated, the margins sharply angled and smooth, tapered to a stipitate-like base (ca. 0.6—-1.2 mm long), more or less abruptly contracted into a broad beak; beaks 0.8—1.2 mm long, whitish green (especially distally), the apex entire or minutely bidentulate, the teeth (when present) weak and up to 0.1 mm long. Achenes 3.3—3.5 mm long, 1.8—2 mm wide, sharply trigonous with slightly concave, rhombic sides, closely enveloped by the perigynium, brown with yellowish angles, apiculate, sessile or nearly so. Stigmas 3. Anthers unknown. Serendipity undoubtedly plays a role in scientific discovery. For instance, W. G. Herter apparently was not aware that he had discovered a new spe- cies when collecting plants of the commonly-occurring Carex phalarotdes. On the herbarium sheet containing the holotype of C. Aerter7 are seven plants of C. phalaroides and a single plant of the new species (Fig. 1). Type: URUGUAY. Dept. Floripa: Estancia Santa Cruz, in campis, Sep 1926, legit. W. G. Herter, Hd. Osten 19091 (S! partim). The holotype is a single plant mounted in the far upper right hand corner of the herbarium sheet, where it is so marked. Carex herteri is known only from the type locality (Fig. 2). The plant was collected “in campis” (on a plain) and has fully-developed perigynia and ripe achenes. The new species is named in honor of Wilhelm G. Herter (1884-1958), who made the type collection and who also was an indefati- gable worker on the flora of Uruguay. ‘ Cn @ C. gibertii ee © C. herteri 7 a “ey — Fic. 2. Map of southern South America showing the distributions of Carex esbertii and C. hertert. As is evident in Fig. 1, the perigynia of Carex hertert appear to be over- sized for the diminutive stature of the plant. Indeed, the combination of large perigynia with few of them per spike, large achenes that are strongly apiculate, and leaves with ciliate margins, readily separates this species from all other Uruguayan carices. It appears to be most closely related to WHEELER, Carex (Cyperaceae) from Uruguay 475 members of the C. phalaroides species complex (sect. Digitatae (Fries) Christ subsect. Radicales Kiik., sensu Kiikenthal 1909), a group of mostly South American carices chatiecenved by having androgynous spikes, cuspidate scales, and sharply trigonous achenes. Although the aspect of C. herter7 and C. phalaroides is strikingly similar, they differ in several features. For ex- ample, the former differs from the latter by having: ciliate leaf margins; fewer perigynia per spike; much larger perigynia that are glabrous and distinctly beaked; larger achenes that are strongly apiculate; and differ- ently-shaped scales. Also see Table 1 Notably, numerous sheets of Carex phalaroides have been examined from eastern Argentina, southern Brazil, southeastern Paraguay, and Uruguay, but thus far only the holotype of C. erter7 is known. It seems remarkable that a species with such large and distinctive perigynia as C. herters has hitherto been overlooked. However, the diminutive stature of both C. herters and C. phalaroides, and their physiognomic similarity to each other, un- doubtedly has contributed to the former species having been overlooked in the past. Another factor may be the early fruiting date of this species. The plants are known to bear ripe achenes in early October, indicating that the species probably flowers in late July and August. Also, though as yet unveri- fied, C. Aerteri may be a local endemic with only a few existing populations. A NEW NAME IN SOUTH AMERICAN CAREX Carex slave G. A. Wheeler, nom. et stat. nov. BASIONYM: Carex phalaroides inth var. crassiflora Kiik., Verh. Bot. Ver. Brandenburg 47:209. 1905. Type: URU- . AY. [Deparr. Mewteuises 1 onevideo: {Cerro de Montevideo, in saxosis, 4 Oct 1864,] Gibert 593 (Lecrorype, here designated: S!; lsoLecroryPE: MVM?). Be- cause the holotype at Berlin (B) was destroyed, the isotype at S$, which was anno- tated by Georg Kiikenthal (in 1904) as “Carex phalaroides Kunth var. crassiflora Kiik. var. nov.,” is chosen as lectotype Carex gibertii is an uncommonly collected species, thus far known from southern Brazil, Uruguay, and northeastern Argentina. Growing in dry sites, it flowers in late September and October and mature fruit has been collected from November through March. The epithet crasszflora is not avail- able for use at species rank because of the earlier C. crassiflora Kiik. Hence, I here offer the name C. gibertii for the a after the Uruguayan bota- nist José Ernesto Gibert (1818-1886), who made the type collection. The plant has been illustrated in Chebataroff (1942, Lam. I, C—D). Additional specimens examined: (Carex giberti/): ARGENTINA. Prov. Buenos Aires: Vela, 10 Mar 1910, Stwckert 21543 (LIL). BRAZIL. State of Rio Grande do Sul: Pelotas, 9 Nov 1955, Sacco 404 (F). Wiens Depart. San José: Mauricio, 940, Chebataroff 4016 (SI) on barrancas, Nov | A476 Stipa 17(2) TABLE 1. A selected morphological comparison of Carex giberti/, C. hertert, and C. phalaroides in south- eastern South America. Character Carex phalaroides Carex gibertit Carex herteri fertile culm height (cm) 2-21 7-13 ca. 2 leaf margins smooth or smooth or ciliate & sparingly ciliolate minutely serrulate (but sparingly so near apex) spike width (mm) 3—5.5(—6) 6-10 4-5 (average 5) (average 8) number of perigynia (5—)L0—40 (5—)8—35 34 r spike perigynium 2.6-3.2 3.2-4.6 4.4-4.8 length (mm) (average 3) (average 3.9) perigynium L.1-1. .6-2 2-2.3 width (mm) (average 1.3) (average 1.7) perigynium pubescent pubescent glabrous perigynium beakless or beakless or 0.8-1.2 beak length (mm) nearly so nearly so piscillate scale average length (mm) 3 { 4.5 (excluding awn) achene 1.5-1.9 2-2.8 3.3-3.5 length (mm) (average 1.7) (average 2.4) & & § achene 0.9-1.2 1.5-1.9 1.8—2 width (mm) (average 1.1) (average 1.6) achene apex short apiculate, short apiculate, strongly apiculate, and tip length (mm) <0.2 <0.2 “a. 0.8 anther the majority <2.6 the majority >2.6 unknown length (mm) Like the new species described earlier, Carex gibertii is a member of the C. phalaroides species complex. Kiikenthal (1905, 1909) and subsequent workers (Osten 1931, Chebataroff 1942) have called these plants C. phalaroides var. crassiflora, but there appears to be a real discontinuity in morphology between the two. Indeed, examination of type material, along with other specimens, reveals that the two entities are closely related yet taxonomically distinct species. Carex gibertii differs from C. phalaroides by having larger spikes, perigynia, achenes, and scales. In Table 1, I have cal- culated mean values of several characters for both species, and, in each case, the corresponding means are so disparate that two species are strongly sug- gested. For example, perigynium mean length and width in C. gzbertis are 3.8 mm and 1.7 mm, respectively, whereas those in C. phalaroides are 3 mm WHEELER, Carex (Cyperaceae) from Uruguay 477 and 1.3 mm. Also, no intergrades between these two species have been seen. Moreover, the two apparently grow in different habitats. Carex giberti has been collected “in saxosis” (rocky soil) and in “barrancas” (ravines), whereas C. phalaroides frequents moist meadows and grasslands, pastures, and roadside embankments. Carex gibertii differs from C. herteri by having taller culms, leaves lacking marginal hairs, more perigynia per spike, smaller achenes and scales, and slightly smaller perigynia that are pubescent and shorter beaked. A mor- phological comparison of 13 characters for the three species discussed in this paper is given in Table 1. ACKNOWLEDGMENTS I want to thank the curators and directors of the following herbaria for the loan of specimens of the Carex phalaroides species complex: B, B BAB, BM, C, F, G, GH, K, LIL, MICH, MIN, MO, NY, P, S, SGO, SI, UC, UPS, US, and WIS. REFERENCES Cuepatarorr, J. 1942. Lista del las especies de Ciperaceas mae ahora conocidas del Uru- guay. Comun. Bot. Mus. Hist. Nat. Montevideo I(3):1— Herter, W.G. 1953. Flora del Uruguay V. Glumiflorae II. Revises Sudamer. Bot. 9:129- 173 KUKENTHAL, G. 1905. Die von + Ule gesammelten brasilianischen Carices. Verh. Bot. Ver. Brandenburg 47:204 eee: NOD: Cypencene Caricoideae. In: Bogle A., ed. Das Pflanzenreich, IV. 20, Heft 38. Leipzig: Wilhelm Engelmann. Pp. 1-824. OsTEN, C. 1931. Las eens del Uruguay. Anales Mus. Hist. Nat. Montevideo (ser. II) 3:109-256 Pepersen, T.M. 1969. Carex. In: Cabrera, A., ed. Flora de la provincia de Buenos Aires. Buenos Aires: Colec. Cient. Instituto Nacional de Tecnologia Agropecuaria. Pp. 318—- 38. Weenies G.A. 1987. A new species of Carex (Cyperaceae) from western South America and a new combination in the genus. Aliso 11:533—537. «1988. Taxonomic notes on Carex (Cyperaceae) of austral South America. Aliso 12: 97- 102. _ «1996. Three new species of Carex (Cyperaceae) from Argentina and a range extension for C. ecuadorica. Hickenia 2(41):189-—200 478 Sipa 17(2) NATURAL HYBRIDIZATION AMONG THREE SYMPATRIC BAPTISTA (FABACEAE) SPECIES IN NORTH CENTRAL TEXAS MATTHEW A. KOSNIK,' GEORGE M. DIGGS, JR. and PEGGY A. REDSHAW Department of Biology Austin College Sherman, TX 75090-4440 U.S.A. BARNEY L. LIPSCOMB Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4060 U.S.A. ABSTRACT A north central Texas Baptista (Fabaceae, Faboideae, Thermopsideae) hybrid complex containing three parents (B. australis, B. sphaerocarpa, and B. bracteata) and the three F, hybrids is investigated using eet spectrophotometric, and isozyme electro- phoretic methods. Two of the three hybrids (B. x bushii and B. x bicolor) have been previ- ously named, but the B. australis x B. sphaerocarpa hybrid (B. x variicolor) is here described for the first time. This is the first report of sympatric occurrence of these three Baptisia species and all the possible F, hybrids. Morphological evidence for introgressive hybrid- ization involving B. australis and B. sphaerocarpa is presented. RESUMEN Un complejo de hibridos del norte de Texas, Baptista (Fabaceae, Faboideae, Thermopsideae) que contiene tres padres (B. australis, B. sphaerocarpa, y B. baci y tres hibridos F, se investiga por métodos morfolégicos, espectrofotométricos, y d de isozimas. Dos de los tres hibridos (B. x bushit y B. x bicolor) habian sido bemibrados previamente, pero el hibrido de B. australis < B. sphaerocarpa (B. x varticolor) se describe aqui como nuevo. Esta es la primera cita de la existencia simpatrica de estas tres especies Baptisia y todos los hibridos posibles de F,. Se presenta la evidencia morfolégica de hibridaci6n introgresiva que afecta B. australis y B. sphaerocarpa. INTRODUCTION The genus Baptista, commonly called Wild Indigo, is a perennial, her- baceous, and conspicuously flowered legume genus with about 25—30 spe- cies occurring in the eastern United States. In Texas they occur in remnant native habitats and are common weeds in overgrazed pastures (Turner 1959). ‘Present address: Department ease ew SIEOGES, oa of Chicago, 5734 South Ellis Avenue, Chicago, IL 60637, makosnik Sipa 17(2): 479-500. 1996 480 Stipa 17(2) Taxonomic difficulties have been common in the group due to extensive hybridization and introgression among species (Correll & Johnston 1970). This taxonomic confusion and the proliferation of secondary compounds within the genus led to early use of chemical taxonomic methods (e.g. chromatography by Turner & Alston 1959; Alston & Turner 1962; Alston & Turner 1963), and later detailed investigation of flavonoid distributions within the genus (Markham et al. 1970; Dement & Mabry 1975). Because of this work, a great deal is known about Baptista both morphologically and chemically. Three Baptisza species occur in the area of north central Texas studied, B. austvalis (L.) R. Br. ex W.T. Aiton var. minor (Lehm.) Fern. (a blue-violet- flowered species, Fig. 1), B. sphaerocarpa Nutt. (a yellow-flowered species, Fig. 1), and B. bracteata Muhl. ex Ell. var. lewcophaea (Nutt.) Kartesz & Gandhi (a cream-flowered species, Fig. 1). Nomenclature follows Kartesz (1994). Baptisia australis reaches the southern extreme of its range in the north central Texas area, B. bracteata reaches its western boundary in the area, and B. sphaerocarpa reaches its northern extreme in central Oklahoma (Larisey 1940a). The area where all three ranges overlap is limited to north central Texas, southern Oklahoma, and western Arkansas (Fig. 2). Hybridization and multispecies hybrid complexes are well documented in the genus Baptisia, both through traditional morphological methods and through extensive chemical analysis. In her monograph on the genus Baptisia, Larisey (1940a) recognized eight Baptisia hybrids and cited the relatively frequent occurrence of hybridism, particularly in areas where the borders of the geographically larger species overlap. At several north cen- tral Texas locations in Grayson County and neighboring Fannin County, several Baptisia species coexist and have been suspected of hybridizing (Kosnik 1996). The B. australis x B. bracteata hybrid, with an intermediate or light blue-violet flower (Fig. 1), has been described previously (B. x bicolor Greenm. & Larisey), and the B. bracteata X B. sphaerocarpa hybrid, with an intermediate or light yellow flower (Fig. 1), has also been described previously (B. x bushiz Small). However, the proposed B. australis X B. sphaerocarpa hybrid, with a flower color varying from brick-red and yellow to orangish to bluish-violet and yellow (Figs. 1 and 3), has not been previ- ously described in the literature. Baptisia bracteata, B. australis, and B. sphaerocarpa have also each inde- pendently been observed hybridizing with other members of the genus. Baptisia australis X B. bracteata hybrids have been observed in Kansas (Hitchcock 1894). Baptista sphaerocarpa X B. laevicaulis hybrids have been de- scribed using morphological and chemical methods in southeastern Texas by Turner and Alston (1959) and Alston and Turner (1962). Baptisia sphaerocarpa x B. leucantha hybrids were observed in Oklahoma by Engelmann (1878), KOSNIK ET AL. idization in Baptisia (Fabaceae) Fic. |. Photographs of parental sae ies and hybrids. Clockwise from upper left: Baptisia . B. x varticolor, B. x bicolor, B. bracteata, B. x bushi, and B. sphaerocarpa 482 Sipa 17(2) petite B. australis var. minor ae B. bracteata var. leucophaea ———_ B. sphaerocarpa MiG. 2. Distribution map for the parental species. (Modified from Larisey 1940a). and in southeastern Texas they were described morphologically by Larisey (1940b) and morphologically and chemically by Alston and Turner (1962). Baptista sphaerocarpa X B. bracteata hybrids were observed in Louisiana by Harper (1938). However, it is only in north central Texas that all three of these species and all three possible F, hybric eon s have been observed in mixed populations. Introgressive hybridization (Anderson 1949), or the move- ment of genes from one species into another via fertile hybrids, has also been suggested for Baptisia (e.g., Alston & Turner 1963; Correll & Johnson Lo 7), METHODS Collection localities. —Plant material was collected at 11 sites in Grayson and Fannin counties, Texas. At three additional sites hybrids were observed but not collected. Table | summarizes collections and observations. Coor- dinates for each site (Table 1) were determined using a Garmin 45 Global Positioning System. Voucher specimens are deposited at the Botanical Re- search Institute of Texas (BRIT) in Fort Worth. Morphological Measurements. —Measurements of a number of characters were made in the field, prior to collection and pressing. Vegetative height was measured from the ground to the highest leaves (0.5 cm increments). Plant height was measured from the ground to the maximum height (0.5 cm increments). Raceme angle was measured with a weighted protractor (2.5° increments, O° considered vertical and 90 considered horizontal). Total lower length (from the base of the flower to the tip of the keel or wings), total KosNik ET AL., Hybridization in Baptisia (Fabaceae) 2 Fic. 3. Photographs of B. X varticolor (B. australis x B. vaerocarpa). Note variable flower color. 484 Stipa 17(2) Tabie 1. Sites and collections. Planes are indicated: B = B. australis, BY = B. x variicolor, Y = B. iphaerocarpa, YC = B. x bushit (sphaerocarpa * bracteata), C = B. bracteata, CB = B. x bicolor (bracteata * australis). Collection number indicates the number of herbarium specimens collected at each loca- tion, an & indicates that the plane was observed but noc collected for pressing. Location Position Collection/Observation Name Latitude Longitude B BY ‘ YC C CB FAN-AG 33° 31! 96° 20! 3 5 10 \ 10 7 G36-CY 33° 44! 96° 49! 10 5 10 GEO-B 33° 30! 96° 31' 10 HAG-C 33° 45! 96° 48 3) MEM-CY 33° 46! 96° 38! 5 > 5 PRE-AG 33° 48! 96° 38! 6 2 Xx 2 2 vi} PRE-BY 33° 47! 96° 36! 7 | 2 HAG-Y 33° 45! 96° 48' LO 1417-Y 33° 41! 96° 39! 5 PWR-B 39° SL! 96° 28' 10 POT-AG 33° 47! 96° 38! XxX | ORIGIN 33% 32! 96° 22' x xX B36-CY 33° 44! 96° 48' Xx x X FAN-CY 33° 313 96° 20' X Xx X Total 36 8 42 14 a2 I> — flower height (from the bottom of the keel or wings to the tip of the ban- ner), maximum horizontal width of the wings, and maximum banner width were measured with digital calipers (0.01 mm increments). All flower measurements were made on the lowest fully developed, intact flower on the best developed inflorescence, with at least one fully developed flower above it. Measurements of all other vegetative characters were made after press- ing using digital calipers (0.01 mm increments). These included pedicel length (of the measured flower); floral bract length, maximum width, width at attachment; petiole length; central leaflet length, maximum width, dis- tance to the maximum width (from the leaflet base); and stipule length, maximum width, width at attachment, and distance to maximum width (from the stipule base). The leaflet measurements were taken on the center leaflet of the first undamaged, well developed leaf below the raceme. Stipule measurements were taken from the same leaf, unless the leaf did not have a stipule, in which case the stipule closest to that leaf was measured (this was often the case with B. sphaerocarpa). Fruit characters were chosen to examine for evidence of introgressive KosNIk ET AL., Hybridization in Baptisia (Fabaceae) 485 hybridization between B. australis and B. sphaerocarpa because of the very different fruit shapes and sizes of these taxa and thus the increased prob- ability of finding unambiguous evidence of introgression. Individuals from the hybrid population were labeled as to flower color during the spring flowering season and fruits were subsequently collected when mature. Fruit measurements were taken using digital calipers (0.01 mm increments) from completely dried fruits collected in the field in June and July from FAN- A6, GEO-B, HAG-Y, and PWR-B (Table 1). These included maximum fruit length (not counting stipe or beak), height, width, and length of beak (from where fruit body narrows dramatically apically to tip of beak). Mea- surements were taken on the lowest well developed, well preserved fruit on an infructescence The Royal Horticultural Society Colour Chart (1966) was also used in an attempt to standardize color measurements. The lowest, mature, un- damaged flower on the raceme was color matched before it dried. Statistical Analyses. —Morphological measurements were analyzed using principal component analysis, multiple discriminant analysis (a multivari- ate statistical technique sometimes called canonical analysis), and analysis of variance and run with Minitab 10.5 Xtra (1995) and BioStat IJ (Pimentel 1995) on a Macintosh Centris 650. Because missing data are not allowed in these analyses, only the characters listed in Tables 2 and 3 were used. Prin- cipal component analysis attempts to unravel patterns of variation within the entire dataset, while multiple discriminant analysis attempts to mini- mize within group variation while maximizing between group variation. Discriminant analysis computes synthetic characters by weighting dataset characters to maximally separate the groups. For the discriminant analysis individuals were assigned to one of six groups based on flower color. Spectrophotometric Analysis. —Flowers from four to nine individuals of each of the six taxa were examined spectrophotometrically. Flowers from three sites (FAN-A6, MEM-CY, and PRE-AG) were selected to represent the maximum variation seen within the populations. Petals from a fresh, un- damaged, mature flower were ground completely in 2 ml ofa 1:4, 1% HCL: 100% EtOH solution; the samples were then centrifuged for three min- utes to separate large particulate material. One ml of the resulting extract was diluted with an additional 2.5 ml of solvent. An ultraviolet-visible light absorbance spectrum was then measured using a Beckman DU-70 scanning spectrophotometer; absorbance scans were run at 1200 nm/min. from 200 nm to 700 nm. Absorbances for peaks at 422 nm, 449 nm, and 540 nm (+2 nm) were recorded. Enzyme Electrophoretic Analysis. —Leaf samples were obtained from 21 in- dividuals including B. australis, B. sphaerocarpa, and all three F, hybrids at FAN-AG in July 1990. Baptisia bracteata was not examined. Starch gel 486 Sipa 17(2) enzyme electrophoretic procedures generally followed Soltis et al. (1983). The tris-HClI grinding buffer-PVP solution described by Soltis et al. (1983) was used with a PVP 40,000 concentration of 12% (wt/vol). Starch gel concentration was 12.5%. The following enzymes were resolved: aspartate aminotransferase (AAT or GOT), glyceraldehyde 3-phosphate dehydrogenase (G3PDH), isocitrate dehydrogenase (IDH), malate dehydrogenase (MDH), 6-phosphogluconate dehydrogenase (G6PGD), shikimate dehydrogenase (SKDH), and triosephosphate isomerase (TPI). Buffer systems follow Soltis et al. (1983): G3PDH and SKDH were resolved on a modification of gel and electrode buffer 1; GOT and TPI on a modification of buffer 8 (modification de- scribed by Soltis & Soltis 1987); 6PGD on a modification of buffer 9; and MDH and IDH on a morpholine system (electrode buffer: 0.04 M citric acid, pH 7.5 by addition of N-(3-aminopropyl)-morpholine; gel buffer: dilute electrode buffer 1:25). Staining procedures followed Soltis et al. (1983 RESULTS Morphological Analysis.—Based simply on flower color, the three parents and their intermediate putative hybrids are easily distinguished. A variety of morphological characters also suggest hybridization. Simple two charac- ter plots of raceme angle versus flower size (height) or leaflet size (length) distinguish six basic morphological groups which support the hybridiza- tion hypothesis (Fig. 4). The three hybrids are generally intermediate mor- phologically between the parents. These graphs also show considerable overlap among hybrid taxa. Principal component analysis separates the taxa in a manner similar to the two character plots. The plot of the first principal component versus the second principal component illustrates that variation within all six taxa (3 parental species and 3 hybrids) is less than the variation between taxa (Fig. 5)—in other words, groups can be observed though there is some overlap. Individuals of the three parental species form discernable clusters toward the periphery of the plot, while all the suspected hybrid taxa are generally intermediate between the purported parental species. Character weightings are given in Table 2. Discriminant analysis separates the taxa with greater resolution than the principal component analysis, but still does not completely separate all individuals of all the taxa (Fig. 6). The discriminant analysis supports 94% of the original Hower color grouping. The 6% of the individuals which are ambiguously grouped by the discriminant analysis on the basis of the mor- phological data are clearly distinct in other characters, such as flower color. The linear discriminant functions are given in Table 3. Kosnik ET AL., Hybridization in Baptisia (Fabaceae) + + + + + © B. australis ee) eee oe ~ (Blue) + ++ + + + + + B. bracteata 100 4 a oe (Cream) ; + + ® go - 4 B. sphaerocarpa D (Yellow) & © a ® 60-4 ros O B.x bicolor 5 Bou © (Blue x Cream) w a - oo oO & 40- o © © Bx bushi O° A : Oo (Cream x Yellow) oa 20-4 fe) fe) ° 4 B.x variicolor a ee. (Blue x Yellow) x 0O4_x x AK HX os o 4 8 oman aD 08 ° 10 12 14 16 18 20 2 24 Flower height (mm) 4h + 7 © B. australis 120 + a (Blue) + + + + 100 - = i: 4 et: + 8B. bracteata 2 + + (Cream) a + + : ® go- Oo + B. sphaerocarpa eS “ 6 (Yellow) < 2 60- x © Oo 8B. x bicolor E ° } oa } ° (Blue x Cream) % Oo 6 © © 404 a B © © B.x bushi o AO © S ‘ (Cream x Yellow) oo 20- o 9° 4 B.x variicolor oe S - Oise x X (Blue x Yellow) 04 _ ee ee is eee 10 20 30 40 50 0 70 80 Leaf length (mm) Fic. 4. Two character plots of raceme angle versus flower height and leaf length. There are additional characters, suggestive of hybridization, that were not included in the principal component analysis or the discriminate analysis because they could not be accurately and consistently quantified. These include hairs (too numerous to count effectively on B. bracteata), flower bracts (length, maximum width, and distance to maximum width cannot e measured on plants without persistent bracts), and flower color (not continuously quantifiable). Baptisia bracteata is densely pubescent, has large persistent floral bracts (10-30 mm long), and cream flowers, while B. sphaerocarpa and B. australis are glabrous, have smaller, early deciduous bracts (O—5 mm long), and have vividly colored flowers (bright yellow in the case of B. sphaerocarpa or blue-violet in the case of B. australis). The hybrids, B. Xx bushii (B. sphaerocarpa X B. bracteata) and B. X bicolor (B. australis B. 488 Sipa 17(2) 6 l UX ; x ca x Sx x OB. australis ] (Blue) 4 a 2 xx = 7 Qe x + 8B. bracteata € 27 & ge (Cream) C ] o fe) . fe) . ros x B. sphaerocarpa 3 - 3 attr & (Yellow) © ome) a . ] . Poh © p to © Oo B.xb oF SE, Mo (Blue ao i aia) oO = 1 Oo Fs ++ ; = 27 i oo | gay © B. x bushii ‘ g + tet oe (Cream x Yellow) ] ” + + 1 A B.x variicolor “— zl re é (Blue x Yellow) : +t 4 + : -6 t T 1 T T T T T T T T TT Tv T t uv T T T T T -€ -4 -2 ) 4 € Principal Component 2 Fic. 5. Principal components analysis. bracteata), are intermediate between their respective parents in all of these characters. Baptisia sphaerocarpa and B. australis are easily distinguished by flower color and plant form. Baptisia sphaerocarpa has numerous flexible, vegetative branches creating a bushy form, commonly multiple racemes per plant, and intense yellow flowers. Baptisia australis has fewer, much more rigid branches, typically only one raceme per plant, and a blue-violet flower color that varies greatly in saturation between plants. Baptisia variicolor is generally vegetatively intermediate, but often is superficially more simi- lar to B. sphaerocarpa; they have vegetative branches creating a bushy form and often have multiple racemes per planc. The flower color of B. X varticolor is strikingly variable. It ranges from intermediate to a patch-work combi- nation of the parental flower colors, with different hybrid individuals hav- ing flowers ranging from brick-red and yellow to orangish to bluish-violet and yellow. This variation can be seen in Figs. | and 3. The Royal Horti- cultural Society Colour Chart measurements are an indication of the varia- tion in color (Table 4). Spectrophotometric Analysis,—The absorbance spectra indicate three diag- nostic wavelengths (Fig. 7). Baptisia australis has a large peak at 540 nm, B. sphaerocarpa has distinct peaks at 422 nm and 449 nm, and B. bracteata has low absorbance at all of the diagnostic wavelengths. Baptista variicolor clearly shows both parental peaks, while B. x bicolor and B. x bushii have peak aes, KosnIk ET AL., Hybridization in Baptisia (Fabaceae) 489 Tasie 2. Principal component character weightings. Variable Component | Component 2 Plant total height 0,272 -0.205 Plant vegetative height 0.275 -0.063 Raceme angle -0.276 0.264 Flower total length -0.236 -0.326 Flower max. wing width -0.036 0.123 Flower total height -0.29] -0.265 Flower max. banner width -0.257 -0.300 Petiole length -0.284 0.184 Pedicel length 0.108 -0.046 Leaflet length -0.002 0.479 Leaflet max. width 0.198 0.344 Leaflet distance to max. width -0.042 0.438 Stipule length -0.332 0.114 Stipule max. width -0.346 0.096 Stipule width at attachment -0.348 0.045 Stipule distance to max. width -0.288 -0.050 absorbances intermediate between those of the parental species (Fig. 7). Absorbance data for the diagnostic wavelengths are given in Table 5. Electrophoretic observations. —Preliminary electrophoretic observations on B. australis, B. sphaerocarpa, and the three F; hybrids show low levels of variation both within and between taxa. Eight of ten loci, Aat, Idh, G3pdh, Mdb-1, Mdh-3, 6P gd, Skdh, and TPI-2, were monomorphic with all indi- viduals of all taxa expressing exactly the same alleles (one individual of B. australis did not express the Mdh-! allele). For Mdb-2 all individuals were monomorphic for the Mdh-2 allele except one individual of B. australis and one individual B. x bicolor which also expressed Mdh-2a. For Tpi-1 all individuals expressed Tp/-16 with most individuals being homozygous. However, one B. x bushii individual and one B. x bicolor individual were heterozygous Tpi-la and Tpi-1b, while five B. australis individuals and two B. x bicolor individuals were heterozygous Tpi-16 and Tpi-Ic. General observations. —It is important to note that the spring flowering times of these species overlap by several weeks. Baptisia bracteata flowers first, followed by B. australis and B. sphaerocarpa, which flower at roughly the same time with some variation depending on the particular site. A significant number of B. bracteata individuals are still lowering when B. australis and B. sphaerocarpa come into flower. Baptisia sphaerocarpa tends to 490 Sipa 17(2) 12 + ] 4 fy fi , fat | y ++ ‘ 8-5 _ + © B. australis | + +4 (Blue) ~ J + — J + 8B. bracteata = (Cream) ae oo c 7 > OP ‘ XB. sphaerocarpa . 4 oo B&D 4 ee (Yellow) = 7 ae oO E © eo) ; = 0 8 S a—y— 0 B.x bicolor 8 J © Sa (Blue x Cream) O | - 8 : 7 20 © B.x bushii x 2 ope 8 (Cream x Yellow) -44 Xx Ro Re aA wn P76 8 ox 4 B.x variicolor et (Blue x Yellow) x ce ee eee ee eS So -6 4 -2 C 2 4 5 Discriminant Variate 2 Fic. 6, Discriminant analysis. flower somewhat longer, generally being the last of the species for which flowers can be observed. Hybrid flowering times in general are intermedi- ate between those of the parent species. Over the course of field work dur- ing the 1996 flowering season, not only were the parental species simulta- neously flowering, but also large bumble bees were observed indiscriminately visiting all three of the parents and hybrids (Fig. 3). While several dozen hybrid individuals were observed, the number of individuals of the parental species are estimated in the hundreds or thou- sands. Baptisia sphaerocarpa in particular is quite abundant, being a vegeta- tional dominant in overgrazed pastures. This species, which is apparently inedible to livestock, can have thousands of individuals in a single large field. At the sites containing more than one parental species, the expected hybrid(s) were always found, and the hybrids were always in much lower abundance than the parents. At the Fannin County site (FAN-A6) contain- ing all six taxa, B. australis is fairly rare, while B. sphaerocarpa is the most common. Baptista varicolor and B. X bushii are the two most common hy- brids at that site. At the Grayson County site (PRE-AG) containing all six taxa, B. sphaerocarpa is rare, while B. australis is the most common. Baptisia bicolor and B. X variicolor are the more common hybrids at that site. Soils differ significantly among sites; in the area studied, B. australis grows on Blackland clay soils, while B. bracteata and B. sphaerocarpa grow on sandy substrates. The hybrids involving B. australis occur either in the Preston Anticline in northern Grayson County where the soil types change rapidly allowing parental coexistence, or in western Fannin County in areas — Kosnik eT AL., Hybridization in Baptisia (Fabaceae) 491 TaBLe 3. Linear discriminant functions. Variable australis bracteata sphaerocarpa X bushit — X bicolor X varticolor Constant -172.04 -202.93 -137.77 -155.08 -169.55 -141.85 Plant total height 0.30 0.04 0.02 -0.13 -0.01 -0.01 Plant veg. height 0.09 0.32 0.67 0.47 0.43 0.61 Raceme angle -0.06 1.06 -0.09 0.36 0.37 -0.10 Flower length 4.10 3.05 3.61 3.24 3.95 3.05 Flower wing width -0.58 0.30 0.62 0.24 -0.42 -0.10 Flower height 4.4] 4,55 2.58 4.42 4.09 3.01 Flower banner width 3.74 2.31 2.85 2.87 3.56 S52 Petiole length -0.83 0.07 -0.86 -0.5 -0,79 -0.71 Pedicel length -0.09 -3.38 0.01 -1.70 -0.61 O.11 Leaflet length 0.29 0.67 0.59 0.72 0.30 0.44 Leaflet width 1.88 1.03 2.06 1.71 1.74 1.89 L dist. to max. width 0.38 0.42 0.47 0.44 0.65 0.46 Stipule length -1.40 -0.57 -1.39 -1.24 -1.44 -1.39 Stipule max. width -0.50 0.90 -0.80 -0.89 -0.21 -1.13 S attachment width 4.88 5.85 2.88 6.77 6.11 4.53 S dist. to max. width 0.26 -4.69 -0.87 -1.71 -1.41 -0.07 of intermediate or mixed (a term used locally by early settlers) soil; plant communities closely parallel the geology of the area. Baptisia bicolor 1s known from only two sites within the study area, while the B. australis x B. sphaerocarpa hybrid is known from three sites in the study area, but has also been reported from southern Oklahoma (Shirley Lusk, pers. comm.). Both B. sphaerocarpa and B. bracteata grow on sandy soils, and the plant distribu- tions observed in the field suggest that the habitat separation between these two species is subtle. The hybrid of these two parental species (B. x bushi?) is by far the most common hybrid in terms of both number of individuals and number of sites at which they occur, which is to be expected since the parental species coexist more readily. The B. X bushii sites range from grazed pastures (G36-CY and B36-CY) to native prairie (MEM-CY Evidence for introgressive hybridization.—Figure 8 shows that fanie lengths and widths of the two monospecific populations of Baptisia australts (GEO-B and PWR-B) were different from those of individuals of the same species from a site with B. sphaerocarpa and known hybrids (FAN-A6). Mea- surements of fruit length and width of B. australis from the hybrid site were intermediate between those of B. australis and B. sphaerocarpa (HAG- Y) from sites with only a single species. In contrast, fruits of B. spherocarpa 49? Sipa 17(2 — TasLe 4. Royal Horticultural Society Color comparisons. Plant species Color matches B. australis DIA, DIB, 92A, 92C, 93A, 93B, 94A, 94B, 95A, LO2ZA B. & varticolor GLA, 17A, 1A, 92A, 92C, 4A, 94B, 9A, 14A B. sphaerocarpa YA B. x bushi DA, YB, 14A, 14B B. bracteata 10C, LOD B. x bicolor SOA, YOB, 92A, 93A, 94A, Tasit 5. Absorbance at diagnostic wavelengths. Plant Sample 422 nm 449 nm 540 nm Type Size Ave St. Err Ave St. Err Ave St. Err B. australis 6 18 02 .20 03 85 16 B. & variicolor 9 AS OS aa) OS 88 08 B. sphaerocarpa 6 78 OS 65 04 05S O01 B. & bushi ) 37 06 ole) 04 O35 O] B. bracteata 6 8 O38 plis) 02 03 Ol B. x bicolor 4 17 02 16 02 4 10 from the hybrid site (FAN-A6) showed almost complete overlap in size with fruits of B. spherocarpa from a monospecific site (HAG-Y). These differences were confirmed by one-way analyses of variance that tested for differences in fruit length and fruit width as a function of group (the four groups: B. australis from monospecific sites, B. australis from hybrid site, B. sphaerocarpa from monospecific site, B. sphaerocarpa from hybrid site). Multiple com- parisions tests (Fisher's) showed that there were no significant differences between the B. sphaerocarpa populations from the monospecific and hybrid sites, but all other pairwise comparisons were significantly different. DISCUSSION Two character plots, principal component analysis, and discriminant analysis all point to a conclusion similar to the one intuitively derived when these plants are observed in the field—that is, some plants are hy- brids intermediate between the respective parental species. Principal com- ponent analysis also shows that within taxon morphological variation is less than between group variation. Based on raceme angle and flower color it is possible that backcrosses between F; hybrids and parental species (B. X bushi crossed with B. bracteata and B. x bicolor crossed with B. australis) KosNIK ET AL., Hybridization in Baptisia (Fabaceae) 493 occur, possibly resulting in introgressive hybridization. At the Grayson County site containing all six species, several individuals are suggestive of this possibility. Two unusual B. bracteata individuals with racemes at a 45 degree angle to the ground were observed, as well as some B. australis indi- viduals with multiple racemes and an atypically flexible stem structure. A B. australis population (from a site also containing B. sphaerocarpa and known hybrids) with fruit length and width values intermediate between typical B. australis and B. sphaerocarpa is suggestive of introgression (Fig. 8). Evidence from additional characters, including genetic markers, is needed to confirm this hypothesis. In contrast, the lack of fruit differences between B. sphaerocarpa from the hybrid and monospecific sites suggests that genes from B. astralis are not moving into B. sphaerocarpa. Discriminant analysis separates the groups well, but some individuals are ambiguously placed. Plants were originally grouped to taxon based on flower color. The discriminate analysis concluded that 6% of the individu- als were not originally placed in the proper group (obvious stray points in Fig. 8). These plants may be backcrosses of F, hybrids parental species, products of introgression, or abnormal individuals. However, the fact that 94% of the individuals are correctly classified, the intermediate placement on the plot of the hybrids relative to the parents, and the intermediacy in non-quantitative characters (e.g. pubescence and bract morphology) leaves little doubt that this is a hybrid system. Spectrophotometric data also support what the eye observes (Fig. 7). Pigment levels in both hybrids involving B. bracteata (B. x bushii and B. x bicolor) are intermediate (an average of the two parents) and in B. X variicolor flower color is additive (a sum of the two parents). The extensive flower color variation seen in B. X variicolor reflects to a certain extent the varia- tion seen in B. australis, but clearly exceeds that level of variation. In some cases the keel is only yellow, while the banner is a deep brick-red or bluish- violet, while in other cases the flower is uniformly orangish or blue-violet- yellow. It is important to note that despite this dramatic variation in ob- servable color, all B. x varticolor flowers exhibit nearly identical absorbance spectra (all have the same peaks with only minor differences in the actual absorbance). Baptisia australis also has variable flower pigmentation; in some flowers the eet | is almost white, while the banner is a variable blue-violet, while in others the whole flower is a variable blue-violet. This variation in color saturation is also observed in the variable absorbance at the 540 nm peak. Baptisia bracteata and B. sphaerocarpa show little variation in flower coloration. The ce data suggest that compounds from both parents might be found in the rids and that the north central Baptisia hybrid swarm may be an elle candidate for detailed chemotaxonomic investigation. In Sipa 17(2) n st ee Baptista X vaviicolor 2 7 Blue x Yellow) a ' \ 100 Fr ' ‘ Cee ee eae ee 1 ’ acute Baptisia aystralis | : Baptisia sphaerocarpa ria \ (Blue) ‘ ; (Yellow) Loo ' 1 t 1 1.00 ; 1 r : \ a O75 =) ‘ \ 1 ! ‘ Lg cn / : ' 1 ! : 1 ! : / : / H ‘ ' 4 t ‘ y ‘ f \ ee \ O75 & ' \} we s i 0.75 i ‘ ar ! ‘ 0.50 ON 1 : \ q ¥ és \ \ i \ r H ‘ \ fi \ 1 i “5 \ \ ' ee i a ‘ ‘ \ 1 \ ' 0.50 ied * ; NaN. ity : " ‘ 0.50 ' - vy Se a a oe ‘ i iN 3 ia ‘\, O38 » Po / ie Ta ‘ i; ! ‘\, : Be An \ \ a a \ . \ \ bey \ \" 0.25 i AW cS 0.25 \ oe \ a a a rT \ 4 ¥ 400 450 S00, 550 600 650 Pas ae \ ‘ a as a. L L FE 1 400) iso 500, SSO 600 O50 400, 450, S00 550 600 650 Baptisia X bicolor Baptista X bushii (Cream x Blue) (Cream x Yellow) 0.75 0 oN nh 0.50 # \ 0.50 BA> i ' \ ‘ 1 i \ 1 ; / 4 ‘ a a Fi i ' vy * ’ \ \ \ ‘ ; ; j 025 RS ti 1 O.25 Fa \ ! ‘s . Ps : ‘ oe ’ } \ er ies A ‘ A a ne 4 ie \ v Soe A -- XS n 1 1 _ > ) n 1 n I 400 450 S00. SS0 600 650 JOO 450 S00 550 600 650 Baptisia bracteata (C ream) 075 is 450, 500. 550, 600 Fic. 7. Absorbance spectrum of flower extracts. Absorbance on the Y axis, wavelength (nm) on the X axis. Solid lines are mean absorbances (Table 5), dashed lines are the mini- mum and maximum absorbance spectra obtained. Kosnik ET AL., Hybridization in Baptisia (Fabaceae) 495 70 4 e 60 4 ® e : : e e® = B. australis only population ae ‘ : ; 7 ‘. (BI hi ly population) = e = 3 © eo e oe o B. australis in hybrid population = J ee eo e (Blue in hybrid population) 2 40 ~ - ‘i 2 0 00 e x B.x variicolor o ae ° CO 13 (Blue x Yellow hybrid) é 2 00° ° ° Fe io ell ly pop on ° ® \ af Me eal 20 7 a vag a 4B. sphaerocarpa in hybrid populatior a : A a a (Yellow in hybrid population) tof * a 0) T T T T T T T T T T T 8 10 12 14 #16 #18 #$%2 22 24 26 28 30 = 32 Fruit Width (mm) Fic. 8. Scatter diagram of fruic length and fruit width for B. australis from pen ake sites, B. australis from sites with B. sphaerocarpa and known hybrids, B. variicolor, sphaerocarpa from monospecific sites, B. sphaerocarpa from sites with B. australis and ces hybrids. Blue-flowered individuals (B. australis) from the hybrid population are interme- diate in fruit shape between B. hie and B. australis from sites with only a single species; this is suggestive of introgression their detailed study of Baptisia flavonoid chemistry, Markham et al. (1970), and later Dement and Mabry (1975), divide Baptisia into subgroupings based on the distribution of flavonoid pigments throughout the genus, placing B. sphaerocarpa and B. australis in separate subgroups within the genus because of large (relative to other members of the genus) chemical differences. In contrast to this, our preliminary electrophoretic data (eight of 10 loci monomorphic and identical) point to a close relationship be- tween B. sphaerocarpa and B. australis. This similarity is consistent with the apparent ease with which hybrids occur. Overall, hybridization, morpho- logical similarity, and electrophoretic data all suggest a close relationship between the species. These findings possibly suggest that the chemotaxo- nomic subgroupings may not be reflective of phylogeny. In either case this hybrid complex warrants further investigation. TAXONOMIC TREATMENT Herbaceous perennials with woody crown and tough roots; leaves very short-petioled, palmately compound with 3 leaflets (or uppermost with only 1 or 2); stipules large to small, persistent or falling early; flowers termi- nal, large, solitary or in erect to horizontal or drooping racemes or solitary in the leaf axils; corollas variously colored; stamens 10, separate; fruit rounded or subcylindric, becoming woody, often with a beak. The plants are avoided by grazing animals; some species are reportedly toxic to livestock. 496 Sipa 17(2) Baptisia nuttalliana Small occurs in north central Texas to the south of the study area but is not involved in the hybrid complex discussed here. It an be distinguished by the yellow flowers occurring singly in the upper leaf axils (at least some) or in very short terminal racemes with 1—4 flowers The only other Texas Baptisia species is B. alba (L.) Ventenat var. macrophylla (Larisey) Isely [=B. lencantha Torr. & Gray}, known only from sandy soil in east and southeast Texas. It is distinguished by its erect inflorescences with white flowers subtended by early deciduous bracts KEY TO NORTH CENTRAL TEXAS BAPTISIA PARENTAL AND HYBRID TAXA Petals cream; inflorescence held distinctly below the level of the leaves; veg- etative structures densely pubescent . Petals blue-violet to yellow, orange, or brick-red, but not cream; inflorescence at least partially erect and held above the level of t B. bracteata — ne leaves or solitary in the upper leaf axils; vegetative structures glabrous to moderately pubescent 2. Petals blue-violet (keel sometimes whitish); stems at base thick (+5 mm diam.); plants usually growing individually; inflorescences one to a ew per plant Petals intensely co moderately oP haee inflorescence vertical (strictly erect); plant glabrous and sane . Petals moderately to pale B. australis ylue- ae inflorescence angled about 45° ee slightly pubescent, not glaucous B. x bicolor 2. Petals yellow or multicolored; stems at base thin (<5 mm diam.); plants usually growing in dense clusters; inflorescences usually numerous per planc. 4. Petals yellow, all + + the same color; foliage glabrous to moderately pubescent 5. Petals vividly yellow; raceme nearly vertical (strictly erect); vegeta- tive portions of the plant glabrous ws) sphaerocarpa 5. Petals pale yellow; raceme angled about 45°; vegetative portions of y & j he plant moderately pubescent oN 3. bushit Petals brick-red and yellow to orangish to blue-violet and yellow, different petals of the same flower often of different colors; foliag glabrous B. X vartitcolor Baptisia australis (L.) R. Br. ex W.T. Aiton var. minor (Lehm.) Fern., Rhodora 39:312. 1937. Baptisia minor Lehm., Ind. Seimin. Hort. Hamb. Baptista minor vat. eens . Baptista texana Buckl., 16. 1827. . Missouri Bot. Gard. 27 ape 1940. c. Acad. it. Sci. Philadelphia 13:452. 1861. Baptisia versicolor eddies: Bot: Cab. 12: pl. 1144. 1826 Baptista vespertina Small ex Rydb., Fl. Prairies & Plains Central N. Amer. 456. 1932. Plants usually growing individually, tall (usually 46-74 cm), of strictly erect habit, with a single, thick (>5 mm diam.), glaucous stem rising from the ground several cm before branching; branches generally few and rigid leaves small (center leaflet 18—34 mm long, average 25 mm) and glabrous; stipules small (4—11(—22) mm long); flowers borne on a usually single Kosnik Et AL., Hybridization in Baptisia (Fabaceae) 497 rigidly erect raceme; floral bracts early deciduous; pedicels 5—15 mm long; corollas large (27-36 mm long), blue-violet of variable intensity; fruit usu- ally much longer than wide, 30-60 mm long, 20-30 mm wide, with a dis- tinct persistent beak that is noticeably widened at base, black at maturity, glabrous. Clay soil, prairies, pastures; Fannin and Grayson cos. in Red River drainage, also Collin Co., also Dallas Co. (Mahler 1988); otherwise only known in Texas from the Panhandle and Titus Co. to the east. Flowering Apr. Se n Campanula isophylla Moretti cv. Stella White 5716 Cultivated 16! Clermontia kakeana Meyen 8128 O’ahu: Kamaile‘unu 14¢ idge, 77 Cyanea hirtella (H. Mann) Hillebrand Wood 0903 (PTBG) Kaua'i: Kalalau 14° Rim, 1190 m Hippobroma longiflora (L.) G. Don 8166 Kaua'i: 3 mi W 144 of Hanalei, 20 m Lobelta erinus L. cv. Blue Moon 8693 Cultivated 28 Lobelia erinus L. cv. Blue Stone 8092 Cultivated elke Lobelia erinus L. cv. Cambridge Blue 86091 Cultivated 248 Lobelia erinus L. cv. Paper Moon 8694 cultivated 2i¢ Lobelia erinus L. cv. Rose Fountain 8584 Cultivated 144 Trachelinm a L. 8713 Cultivated Wis Collection numbers without collector name or herbarium acronym were collected by Lammers and Cultivated” plants were grown from Sata, obtained seeds in northeastern Illinois; the Remaining localities are in the Hawaiian Islanc First report for the species d Confirms a previous report oo the species. The counts reported for the lobelioids Clermontia kakeana and Hippobroma longiflora confirm previously reported counts compiled by Lammers (1993). Similarly, the counts reported for the campanuloids Campanula isophylla and Trachelinm coeruleum confirm previous reports of the same numbers. The former has been counted several times, most recently by Damboldt (1965); all reports show » = 16. The latter likewise has been counted sev- eral times, most recently by Contandriopoulos et al. (1984). Most of these reports show # = 17, though Contandriopoulos et al. (1984) found some plants with » = 16 in addition to 17-paired plants. ACKNOWLEDGMENTS We thank Kenneth R. Wood, National Tropical Botanical Garden, for providing the fixed and vouchered material of Cyanea hirtella. REFERENCES CONTANDRIOPOULOS, J., C. FAVARGER, and N. GALLAND. 1984. _ a l’étude onomique des Campanulaceae du Maroc. Bull. Inst. Sci. 8:101—114. Dampotpt, J. 1965. ae anaes revision der isophyllen een in Europa. Bot. Jahrb. he 84:302-358 DE VILMORIN, R. and M. ees 1927. Nombre des chromosomes dans les genres Lobelia, Linum et chez quelques autres espéces végétales. Comp.-Rend. Hebd. Séances Mém Soc. Biol. 96:166—-168. Lammers, T.G. 1993. Chromosome numbers of Campanulaceae. II]. Review and integra- tion of data for subfamily Lobelioideae. Amer. J. Bot. 80:660—675. 522 Stipa 17(2) sand D.H. Lorence. 1993. A new species of Cyanea ea aaa Lobelioideae from Kaua'i, and the resurrection of C. remy. Novon 3:4 436 THULIN, M. 1983. Some tropical African Lobeli comments on taxonomy and nomenclature. Nord. J. Bot. 3:371—382. —_— iaceae. Chromosome sotee new taxa and 523 NOTES CAPRARIA MEXICANA (SCROPHULARIACEAE), AN ENDANGERED ADDITION TO THE UNITED STATES FLORA While surveying proposed drilling sites in Falcon State Recreation Area (Starr County, Texas, USA) on 7 February 1993, I found a shrub I could not identify. Other south Texas botanists consulted did not know it. Billie L. Turner identified it as Capraria mexicana Moric. ex Benth. (Scrophulariaceae), a species heretofore not reported from the United States. A minimum of 66 mature Capraria mexicana, plus a number of seed- lings, grew in three small stands in a reservoir-influenced habitat in a state park at the western edge of the lower Rio Grande Valley. The site lies near the inlet (or east) side of the low point between hilltops (one with screened shelters, the other with a recreation hall) making up a peninsula with Fal- con Reservoir flooding the lower slopes. Elevation is about 305 feet. Ex- tremely high reservoir levels may flood the site. Large, solid stands of spiny aster (Erigeron ortegae Blake) alternating with coastal bermuda (Cynodon dactylon (L.) Pers.), separate the C. mexicana known. Among the C. mexicana grows a carpet of coast germander (Tewcrinm cubense Jacq.), with a few granjeno (Celtis pallida Torr.) saplings and twinevines (Sarcostemma cynanchoides Dene.). A few white-leafed Indian mallows (Abutilon hypoleucum Gray) and tree to- baccos (Nicotiana glauca Grah.) grow in the coastal bermuda. Capraria mexicana grows along streams widely in Tamaulipas north to the San Fernando area. Billie L. Turner notes (pers. comm., 2 April 1993) that it “gets to about the upper 2/3 of Tamaulipas state.” Bentham (1846) described the C. mexicana collected by Jean Louis Berlandier of Matamoros at Tampico, Tamaulipas and Greenman (1904) cited collection localities. Sprague (1921) revised the genus. No subsequent citations were found. Capraria mexicana tanges from Starr County, Texas at least to Tantoyuca, Veracruz (Tantoyuca is 105 km SW of Tampico). Intermediate localities (Sprague 1921) include Sota la Marina and Tampico, Tamaulipas. A provi- sional map provided by B.L. Turner (Fig. 1) shows the range of the species, based upon records at TEX and LL. Previous to this report, the northern- most record is Harde Le Sueuer’s 1939 collection from the “shores of La- guna Anda la Piédra,” near San Fernando, Tamaulipas. The tallest of the much-branched shrubs bearing numerous flowers reach 2 m in height. Leaves linear-lanceolate, nearly sessile, acute leaves, serrate distally on mature shrubs, while remaining entire on seedlings; specimen leaves 2—5.5 cm long and 0.4—1 cm wide; older leaves shed, perhaps with increasing aridity; pedicels linear, axillary, ranging from | to 1.5 cm, shorter Sioa 17(2): 523. 1996 Fic. 1. Documented distribution of Capraria mexicana, based upon records at TEX and LL. than subtending leaves; corolla white, greatly exceeding sepals; dry corol- las measure about 8 mm, including lobes measuring 6 mm; corolla lobes lanceolate, acute, and twice as long as corolla tube; style exceeds stamen length, but less than corolla length, with one enlarged stigma; anther of each of five stamens bilobed basally; fully grown (green) capsules nearly twice the length of the (4)5 sepals (with calyx lobes shriveling as capsules mature and brown), 5-8 mm long and 3—4 mm thick, cylindrical, the two capsular lobes separated by a longitudinal septum and appearing to open near the distal (upper) ends and releasing numerous minute seeds. On 7 February, stems were leafed to the ground; on 28 February, many of the arger lower leaves had withered; on 14 April, the lower trunks were leafless. Flowers were observed from 7 February (numerous) to 14 April (few). In the similar C. bzflora, corolla lobes are oblong, obtuse, and as long as or longer than the tube (Sprague’s 1921 key). Justin Williams adds (pers. comm., 17 July 1995) that C. mexicana is glabrous and C. biflora is pubes- cent. [Description (other than corolla from key) stems from specimens ex- amined and field observations in 1993 by the author. } — Stipa 17(2): 524. 1996 NOTES 525 Despite the extensive medicinal literature on the allied C. biflora (Brenneker 1961; Cruz 1965; Diaz 1976; Edwin 1971; Eldridge 1975; Garcia Rivas 1982; Hurtado 1894; Lavadores 1969; LeCointe 1947; Martinez 1969; Morton 1975; Morton 1977; Morton 1981; Nunez- Melendez 1964; Roig y Mesa 1945; Souza-Novelo 1943; Standley 1930; Stehle & Stehle 1962; Wong 1976), I found no evidence that C. mexicana has been examined yet for medicinal components. Capraria biflora is extensively used as a medicinal plant for a wide variety of ailments (references above), as an herbal tea (Morton 1981; Tanaka 1976), and in place of China tea (references in Sprague, 1921) from the Bahamas and south Florida across the West Indies into South America, north through Central America into northeastern Mexico. Its many English common names usually include a geographic name plus Tea and its Spanish, French and Portugese names frequently are direct translations. Since the species has been most commonly collected in the southern half of the Tamaulipan B1- otic Province and since its congener is widely used as both medicinal and herbal tea, I propose using the common name of Tamaulipan Tea for C. mexicand. The Texas Organization for Endangered Species listed Capraria mexicana as Endangered in Texas in 1993 (TOES 1993) and the Native Plant Project listed it as Endangered in the lower Rio Grande Valley of Texas in 1993 (NPP 1993). The species is not considered Endangered in Mexico. With the extremely low water levels in Falcon Reservoir and low rainfall in 1995 and early 1996, the plants suffer greatly from drought. A wind storm on 7 April 1993 snapped some Capraria branches and uprooted some Tree To- baccos. In 1993 the park population escaped becoming part of a proposed drilling pad (Ideker 1993a,b). I gratefully thank R.I. Lonard, M. Johnston, B.L. Turner, J. Poole, G.C. Janssen, J. Williams, M.D. Reed, and B. Lipscomb for discussion, identification, and/or review. Research and initial collection were made persuant to State Park Scientific Study Permit #7-93 (and subsequent re- newals) granted by D. Riskind. Initial collection sent to TEX.— Joe Ideker, Native Plant Project, P.O. Box 1433, Edinburg, TX 78540-1433, U.S.A. REFERENCES BentHam, G. 1846. Capraria. In: De Candolle, A.P., Prodromus 10:429. BRENNEKER, P. 1961. Jerba-Kruiden van Curacao en hun gebruik. Boekhandel St. Augustinus, Curacao, N.A Cruz, G.L. 1965. Livro verde das plantes medicinais e industriais do Brasil. Vols I and I. Velloso, S.A., Belo Horizonte, Brasil. Stipa 17(2): 525. 1996 526 Sipa 17(2) Diaz, J.L. 1976. Indice y sinonimia de las plantas medicinales de México. Instituto Mexicano para el Estudio de Is Plantas Medicinales, A.C. Monograficas cientificas 1 Epwin, G. 1971. Flora of Peru. Field Mus. Nat. Hist. Publ. 13(513): 461-717. Evpripce, J. 1975. Bush medicine in the Exumas and Long Island, Bahamas; a field study. Econ, Bot. 29:307—332. SPECIES COMMITTEE, NATIVE PLANT Project (NPP). 1993. Endangered, threat- el ace. wateh list, and extinct plane species of the lower Rio Grande Valley of Texas, ed. III. Sabal 10(6):3-10 Garcia Rivas, H. 1982. Enciclopedia de plantas medicinalas mexicanas. Editoria Posada, Mexico. 655 pp. GREENMAN. 1904. {Ticle unseen.] Proc. Amer. Acad. Arts. 40:31. Hurtapo, C. 1894. Compendio de le Botanica elemental. Flora de las Isla de Curazao— 1891-1892. pppendice: Plantas Medicinales, pp. 366-422. Imprenta de la Libreria de A. Bethencourt e Hijos. Iprker, J. 1993a. Hearing testimony given at the hearing on additional petroleum explo- ration in Falcon State Recreation Area. The Sabal 10(1):2-4 IpDEKER, J. 1993b. a State Recreation Area drilling cedars Sabal 10(2):9. Lavapores, V., Dr. G. 1969. Estudio de las 119 tae medicinales mas conocidas en Yucatan, México. Anes Mérida, Yucatan, Méxi LeCointe, P. 1947. Arvores e plantas uteis ene e aclimadas). 2d. ed. Brasiliana Ser. 5., Vol. 251. Companhia Editora Nacional, Sio Paulo, Brasil. Martinez, M. 1969. Las plantas medicinales de México, quinta edicién. Ediciones Botas, od. México. Morton, J.B. 1975. Current folk remedies of northern Venezuela. Quart. J. Crude Drug Res. 13:97-122. —_—______. 1977. Medicinal and other plants used by Hees on North Caicos (Turks and Caicos Islands, West Indies). Quart. J. Crude Drug Res. —_______.. 1981. Atlas of medicinal plants of Middle eae Charles C. Thompson, ue lee Illinois. Nunez-MELENDeEZ, E. 1964. Plantas medicinales de Puerto Rico. Bol. 176. Univ. Puerto Rico Est. Exper. Agricola, Rfo Piedras. Roic y Mesa, J.T. 1945. Plantas medicinales, aromaticas, 0 venenosas de Cuba. Cultural, S.A., Havana. Souza-Nove to, N. 1943. Plantas medicinales que viven en Yucatan. Inst. Técnico Agrico Henequenero. Chaczikin, Yucatan. [Mimeo SPRAGUE, T.A. 1921. A revision of the genus Capraria. Kew Bull. 5(24):205—212. STANDLEY, P.C. 1930. Flora of Yucatan. Field Mus. Nat. Hist., Bot. Ser. 3:159—492. STEHLE, H. and M. Stents. 1962. Flore medicinale ilustree (Flore Agr. Antil. Francaises, ee: a vol. IX). Imprimerie Parisienne, Pointe-a-Pitre, Guadaloupe. TANAKA, T. 1976. Tanaka's cyclopedia of edible plants of the world. Keigaku Publ. Co., Tokyo. TEXAS ORGANIZATION FOR ENDANGERED Species (TOES). 1993. Endangered, threatened and watch lists of Texas plants. TOES Publ. 9 3d. rev (and 1993 addenda). Wonc, W. 1976. Some folk medicinal plants from Trinidad. Econ. Bot. 30:103—142. Sipa 17(2): 526. 1996 PERESKIA ACULEATA (CACTACEAE), IN THE LOWER RIO GRANDE VALLEY OF TEXAS Ona 11 May 1996 field trip, a landowner told me he had cactus climbing into trees on a tract of land which had never been cleared. The site is 3.3 km northwest of Sebastian, Willacy County, Texas, U.S.A. The owner stated that the wooded tract and his residence had been in the family for many years and the cactus had not been planted. Nor did it grow around the residence. The tract is now surrounded by fields of Upland Cotton, Gossypium hirsutum L., and Grain Sorghum, Sorghum bicolor L. var. durra Hubbard & Rehd. I found the hanging Pisonia-like leaves with little trouble. That 1s, the small spines found me several times as I searched until I found the wick- edly-spined stems and areoles largely hidden in guinea grass (Panicum maxi- mum Jacq.). The stems climb into trees, branch, and spread like vines. The leafy, distal ends with hooked spines hang down within two meters of the ground. William MacWhorter identified it as Pereskia aculeata Maller, lemonvine. Benson (1982) gives the range only as “Carribean Tropical For- est” and cites localities where the species has escaped cultivation in Florida. Britton and Rose (1963) give the range as the West Indies and Atlantic and Carribean Coasts of South America. Bravo Hollis (1937) indicates it is encountered in the wild in the littoral regions of the Gulf of Mexico.) I assume these plants are naturalized from some former site now under the plow. The habitat is typical of riparian woodland dominated by sugar hackberry (Celtis laevigata Willd.) with a ground cover of guinea grass. This habitat may be a relict reflecting an ancient channel of the Rio Grande. The soil is Willacy fine sandy loam, 0 to 1% slopes (Turner et al. 1982). Where undisturbed, the natural ground cover is mostly subshrubs such as pigeonberry (Rivina humilis L.) and chaff-flower (Achyranthes aspera L.), and scattered clumps of rusty-seeded paspalum (Paspalum langei (Fourn.) Nash), manzanita (Malpighia glabra L.), and tropical sage (Salvia coccinea Buchoz ex Etlinger). A list of all species found on the tract appears in Ideker (1996). Voucher specimens: Pereskia aculeata: TEXAS. Willacy Co.: 3.3 km NW of Sebastian, | 1 May 1996, Ideker ABRI1. Achyranthes aspera: TEXAS. Willacy Co.: 3.3 km NW of Sebastian, 11 May 1996, Ideker ABROS (BRIT). —Joe Ideker, Native Plant Project, P.O. Box 1433, Edinburg, TX 78540-1433, REFERENCES BENSON, L. 1982. The cacti of the United States and Canada, Stanford Univ. Press, Stanford. Brirton, N.L. and J.N. Rose. 1963. The Cactaceae. Dover, New York. Sina 17(2): 527. 1996 $28 Sipa 17(2) Bravo Ho.tis, H. 1937. Las cactaceas de México. Univ. Nac. México, Cd., México. IDEKER, J. 1996. Vegetation of the ABR tract, Willacy County, Texas. Sabal 13(6):7—8. Turner, A.J., H.H. Hype, and J.L. Jacons. 1982. Soil survey of Wil U.S. Gove. Printing Off., Washington, D.C. acy County, Texas. Sipa 17(2): 528. 1996 Book REVIEW 529 BOOK REVIEW Witttam A. Weper and RONALD C. WitTMANN. 1996. Colorado Flora: Eastern Slope (Revised Edition). (ISBN 0-87081-387-0, pbk). University Press of Colorado, Niwot. $29.95, xl + 524 pp. AND: WittiaM A. WEBER and RONALD C. Witrman. 1996. Colorado Flora: Western Slope (Revised Edition). (ISBN 0-87081-388-9, pbk). University Press of Colorado, Niwot. $29.95, xxxvil + 496 pp. William Weber is a veritable Roger Tory Peterson of Colorado Botany, given his fifty year output of user-friendly identification manuals and his concomitant influence in popu- larizing the identification of a regional biota. The nearly inevitable results of such guides are an increased awareness, appreciation of, and concern for the continued viability of local species. These recently published volumes are revisions of earlier works written by Weber alone (Weber 1987, 1990) and are of considerable merit. Given Ree striking similarity in aces format it seemed appropriate to review both simultaneous volumes are remarkably similar to their predecesso seer 1987, 1990) and fol- ae a sequence of Preface, Introduction, Key to Families, ae and Fern Allies, Gymno- sperms, Angiosperms, Figures, Glossary, Index to Common Names, and Index to Scientific Names. The hydrologic Continental Divide separates the geographical ranges covered by these volumes. The Eastern Slope flora treats approximately 2300 species, whereas the Western Slope flora covers some 2100 s pri Infraspecific taxa, not included in their floristic summaries, are included in the The Preface in both manuals is nearly aeseel. Dr. Weber notes with satisfaction his own influence on the increased appreciation and study of botany in Colorado. His contri- butions are substantial, but the volumes lack acknowledgments and literature citations, and the section on eponymy by itself seems somewhat insufficient as a tribute to the work of earlier botanists. I am happy to see the authors flying a warning flag regarding the deleterious effects of invasives on native vegetation, an increasing worldwide problem and recent symposium topic in California. One major conduit for the spread of invasives not mentioned by the authors is the turning of the soil during the creation of fire lines in fire fighting efforts (Snow 1992-1994: 62), a factor that land management agencies may wish to add, however slightly, to their overall considerations of fire policy. ntroductions are long. However, for those new to the joys of wildflowers and plant identification the Introductions provide a good summary of a wide body of information. Both volumes begin with “Scope of the Book” and “The Eastern for Western} Slope”, wherein physiognomic provinces are defined. For example, the Eastern Slope describes: gh Plains, North and South; The Mountain Front; The Intermountain “Parks”; North Park: South Park; The San Luis Valley; The Sanraceh Range and Upper Arkansas Valley, and The Wet Mountain Valley and Spanish Peaks. These categories are followed by discussions of Floristic Zones, Plant Geography, Endangered Habitats, Rarity, Revegeta- tion, Alien Plants, Some Floristic Statistics, How to Learn to Recognize the Big Families, Fariilies with a unique character, Common Names, Pronunciation, Miscellaneous Notes, How to Collect and Preserve Botanical Specimens, The — of Voucher Saarinen. Collecting Ethics, Plant Identification, How to Use the Keys, and “Eponymy. Botanists honored in Colorado Plant Names”. The authors’ attention to user-friendliness is evident in “How to Learn to Recognize the Big Families” and “Families with a Unique Character”. Sipa 17(2): 529. 1996 530 Sipa 17(2) One oversight in “How to Collect and Preserve Botanical Specimens” was their a to mention the now increasingly common use of Global Positioning Systems (GPS tems to pinpoint collection localities on ale and the importance such data will ne for use with eae aphical Information Systems (GIS) Within each major category (Ferns and Fern Allies, Gymnosperms, etc.) the taxa are arranged pe here by family and by genera within families. The family and generic descriptions may include interesting information such as: hints for identification; geo- graphical ranges; medicinal, ornamental, and folk uses; and, among other things, invita- tions to botanize!, like their suggesting where the rar ryptogramma stelleri might be discovered (Eastern Slope volume). The species descriptions are brief and emphasize infor- mation relevant for diagnostic purposes. Taxonomic authorities, common names, and geo- graphic ranges are included. I have not used the present volumes in the field, but based on personal experience during a ten year residence in the Rocky Mountain region, I found the Key to the Families of earlier editions easy to use Interspersed in the Eastern Slope are two sets of 32 color photographs, whereas the Western Slope has four sets of 16 photographs. The sets have no apparent taxonomic or other logical arrangement such as flower color (popular in some guides), but in most in- stances the photos are crisp. The Eastern slope text is followed by 103 black and white plates prepared by Ann Papageorge, Carolyn Ensle, and Harold Roberts, nearly all of which illustrate more than one species. The companion volume of the Western Slope has 107 plates (sixteen of which are the fourth set of color plates mentioned above). e only obvious printing problems I spotted were large columnar gaps in the Index to Scientific Names (pp. 488, 492, 494) and fuzzy text (p. 471) in the Western Slope manual. The biggest concern with these volumes, like their predecessors, is the extensive use of neric names unrecognizable to even well-trained students of the North American flora. The following examples probably ring few bells of Soe a to most American plant systematists: Sabina monosperma, Bolophyta alpina, Breea arvensis, Psychrophila leptosepala, Cc ba opyrnm ¢ ita wcum, Nuttallia decapetala, or Seriphidium tr ae hese are reco zed by botanists as Juniperus monosperma, Parthenium alpinum, Cirsium arvensis, Caltha yal Auli cylindricum, daa os eniias and Artemisia tridentata. As an example il jab) at the family level, the authors segregate Alsinaceae from a larger and more inclusive te which does not accord with other soles floristic (Larson 1986; Hartman 1993) or integrative family treatments (Cronquist 1981; Thorne 1992). These are but few of the many nomenclatural departures when compared with other North American treat- ments. To their credit, however, the authors indicate the more commonly used nomencla- ture in the text The lengthy exposition in an earlier edition regarding Weber's generic concepts has been removed from these volumes (Weber 1987: 9-13), but the effects of that philosophy are the primary basis for the large number of unfamiliar generic (and even family) names. Weber (1987: 12) once argued that “it is not necessary ... co produce detailed justification for adopting a particular taxonomic point of view (a name)”. This statement suggests that the process of classification relies on (or did at one time) significant measures of experience and intuition. Today, classifications based even partly on intuition are much less tenable, and the creation of new names in the absence of new data has been criticized deserv edly (Brown et al. 1993). It is true Weber often has merely raised existing infrageneric taxon names up a notch ot the generic level. However, a phylogenetic analysis is now seen by many (most?) practicing Sas a necessary antecedent of revising classifications and ene name ae. new or aenebwise: Floras should largely reflect recent system- Sipa 17(2): 530. 1996 Book REVIEW >31 atic revisions that are based squarely on sound data (see also Stevens 1990). This is a view that some taxonomists will still find uncomfortable, but it reflects what now appears to be a majority view. Unfortunately, the use of many of the generic names in these volumes has not been justified by recent monographic and/or phylogenetic work. On the other hand, I predict that at least some of the genera recognized by the authors ultimately will be up- held by phylogenetic work. If we cast concerns about nomenclature aside and look at the larger picture, however, we must be grateful for what we have: Modern and knowledgeable accounts of the higher vascular plants of Colorado, written by the person who knows them best. Residents of some other Western states are less fortunate, given the lack of up-to- date state Floras written by an expert with years of experience. I intend to buy my own copies, and suspect these volumes will be purchased and used by many others as well. LITERATURE CITED Brown, G.K., H.E. Luter, and W. Joun Kress. 1993. Comments on the responsibilities of taxonomists. J. Bromeliad Soc. 43:154—156. Cronaguist, A. 1981. An integrated system of classification of flowering plants. Columbia Un Bees Press, New Hartman, R.L. 1993. Caryophyllaceae. In: J.C. Hickman, ed. The Jepson Manual: Higher Plants of ae aen University of California Press, Berkeley. Pp. 475-497. Larson, G.E. 1986. Caryophyllaceae. In: Great Plains Flora Association, Flora of the Great Plains. University Press of Kansas. Pp. 192-2 Snow, N. 1992-1994. The vascular flora of southeastern Yellowstone National Park and the headwaters —. of the Yellowstone River, Wyoming. The Wasmann Journal of Biology 50(1-2):52 THORNE, R.F. 1992. An canes phylogenetic classification of the flowering plants. Aliso 13:365-389. Weber, W.A. 1987. Colorado flora: Western Slope. Colorado Associated University Press, Boulder . 1990. Colorado flora: Eastern Slope. Colorado Associated University Press, Niwot. —Neil Snow, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missourt, 63166. Current address: Queensland Herbarium, Meiers Road, Indooroopilly, QLD 4068, Australia. Spa 17(2): 531. 1996 532 Sipa 17(2) CORRECTION Figure 4, p. 26, Sida 17(1), Sep 1996. The caption in Figure 4, Bacon and Spellenberg, “Hybridization in Qvercus” [Sida 17(1): 26] was inadvertently reversed in attempting to make a correction in final proof. The top figure, labelled “A” in the right margin is indeed Q. eduardii, not 4C as stated in the caption. The bottom figure, labelled “C” in the right margin is Q. conzatti1, not 4A as stated in the caption. Sida and the authors apologize for this unfortunate error. Noteworthy Carex, Cyperus, Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia, Louisiana, Mississippi, North Carolina, Tennessee, and ‘Texas Charles 1. Bryson, fob Rk. MacDonald, Richard Carter, and Stanley D. Jones 501 Documented chromosome numbers 19964. Chromosome numbers of Campanulaceae. LV. Miscellaneous counts Thomas G. Lammers aud Nancy Hensold 519 Notes Capraria mexicana (Scrophulariaceae), an endangered addition to the United States flora 523 Pereskia aculeata (Cactaceae), in the Lower Rio Grande Valley of Texas 527 Book notices and reviews 320, 306, 412. 416, 422, 436, 529 Correction 53 V NQ ISSN QO36- 1458 CONTENTS Taxonomy of North American species of Zizania (Poaceae) Edward E. Terrell, Paul ML. Peterson, James L. Reveal, and Melvin R. Duvall 533 Passiflora cerasina. a new species of Passiflora subgenus Passiflora (Passifloraceae) from French Guiana Hilaire Annonay and Christian Feuillet 55] Calcaratolobelia (Campanulaceae): A new genus of spurred lobelioids from Mexico and Central America Robert L. Wilbur 555 Sideroxylon alachuense, 1 new name for Bumelia anomala (Sapotaceae Loran C. Anderson 505 Crataegus nananixonii (Rosaceae, Ser. Intricatae) a new species of hawthorn from Eastern Texas J.B. Phipps and RJ. O’Kennon 569 Palynological evidence supporting the identity of two taxa of Berberis (Berberidaceae) from Tibet Arti Garg, Tariq Husain, and R.R. Rao yD Nomenclatural notes and « svnopsis of the genus Myrsine (Myrsinaceae) in Mesoamerica Jon M. Ricketson and John J. Pipoly Ill 579 Nomenclatural notes and a synopsis of Mesoamerican Stylogyne (Myrsinaceae) Jon M. Ricketson and Jobn J. Pipoly Il 591 Floristic composition of the San José del Cabo oasis. Baja California Sur, Mexico José Luts Leon de la Luz, Raymundo Dom iniguez Cadena, Miguel Dominguez Leon, and Jose Judn Perez Navarro 599 Contributions to the Hora aad ecology of tie northern Long)cat Pine Belt in a oe Rankin County, Mississippi CONTRIBUTIONS Michael Wayne Morris TO BOTANY Oe) VOLUME 17 NUMBER 5 SEPTEMBER, 1997 CONTRIBUTIONS TO BOTANY FOUNDED BY LLOYD H. SHINNERS 1962 Wm. F. Mahler Publisher 1971-1992 Director Emeritus HH. Sohmer Director Barney L. Lipscomb Editor Botanical Research Institute of Texas, Inc. 509 Pecan Street Fort Worth, Texas 76102-4060, USA 817 332-4441 / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http: www. brit.org/sida John W. Thieret Prof. Dr. Felix Llamas Associate Editor Contributing Spanish Editor Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia Northern Kentucky University Universidad de Leon Highland Heights, Kentucky 41076, USA H-24071 Leon, Spain lidelines for contributors are available upon request a on oon inside back cover of the last issue of each volume. Subscription per year: $25. Individual, $35. USA eta $45. Outside USA: numbers issued twice a year Om © SIDA, CONTRIBUTIONS TO BOTANY, Volume 17, Number 3, pages 533-6 Copyright 1997 Botanical Research Institute of Texas, Ine. Printed in the United States of America ISSN OO30- 1488 TAXONOMY OF NORTH AMERICAN SPECIES OF ZIZANIA (POACEAE) EDWARD E. TERRELL Department of Plant Biology University of Maryland College Park, MD 20742-5815, U.S.A. PAUL M. PETERSON Department of Botany National Museum of Natural History Smithsonian Institution Washington, DC 20560, U.S.A. JAMES L. REVEAL Department of Plant Biology University of Maryland College Park, MD 20742-5815, U.S.A. MELVIN R. DUVALL Department of Biology and Microbiology South Dakota State University Brookings, SD 57007-0595, U.S.A, ABSTRACT The grass genus Zizania (Poaceae: Oryzeae) is defined to include four species, three of temperate North America and one of eastern Asia. Keys, descriptions, distributions, and synonymies are provided for the North American entities: Z. aquatica, Z. palustris and Z texana. We discuss recent literature and the possible origin of Z. palustris var. interior. RESUMEN El género de gramineas Zizania (Poaceae: Oryzeae) se define como un género con cuatro especies: tres de la parte templada de Norteamérica y una de Asia oriental. Se incluyen claves, ie distribuciones y sinonomias para las entidades norteamericanas: Z. aquatica, Z. palustris y Z. texana. Examinamos la bibliograffa reciente y el origen posible de Z. palustris var. interior. The genus Zizania (Poaceae: Oryzeae: Zizaniinae) is defined here to in- clude four species. Three are North American (Z. aquatica L., Z. palustris L., and Z. texana Hitchc.) anda fourth (Z. latifolia (Griseb.) Turcz. ex Stapf) is native to eastern Asia. Zizania aquatica and Z. palustris are important constituents of aquatic plant communities. The caryopses (grains) of 7 — Sipa 17(3): 533-549. 1997 534 Stipa 17(3) palustris (northern wild-rice) are an important food being harvested in the wild and from cultivated paddies in Minnesota, California, Ontario and Saskatchewan. Zizania aquatica (southern wild-rice), native to the Atlantic and Gulf Coastal Plains, is not cultivated, but has been grown experimen- tally. Zizania texana, a local endemic 1n southcentral Texas, treated by Terrell et al. (1978), was included in experimental studies by Duvall (1987) and Duvall and Biesboer (1988a, 1989). We further understand that technical studies are being carried on by Francis Horne (pers. comm.) in San Marcos, Texas. Basal parts of the culms and rhizomes of the Asian species, Z/zania latifolia, are edible when infected with the fungus Usti/ago esculenta Hennings (Terrell & Batra 1982). It is important to note that the presence of the fungus prevents flowering. Living material of Z. latifolia should be banned from importation into North America because infection of our native spe- cies might have serious ecological and economic consequences. Hitchcock (1951; revised by Chase) referred to Zizaniopsis miltacea (Michx.) Doell & Aschers. as “southern wild-rice,” implying close relation- ship to Zizania; however, Zizaniopsis and its related genus, Lwzsola (includ- ing Hydrochloa), are quite distinct in important reproductive characters from Zizamia and were later placed in a separate subtribe (Terrell & Robinson 1974; Duvall et al. 1993). This paper cae keys, descriptions, distributions and synonymies for the North American species of wild-rice. Representative specimens cited are primarily those in the US herbarium. Recent publications concerning the taxonomy of Z/zania are discussed, with emphasis on Zizania palustris var. 7vferior. SYSTEMATIC TREATMENT sat i ,op. El. 2: 2 1753 {& Gen. Pl. ed. 5. 427. 1754]. Hydropyrim Hortus Berol. 1: 252. 1827, nom. illeg. Ceratochaete Lunell, Amer. Midl. Nacu- pee. 4.1915, nom. illeg. by lectotypification. Lecrorype: Zizania aquatica L. Revel ; ee 72:6. 1992]. = Melinum Link, Handbuch 1:96. 1829, nom. illeg. (non Medikus, Philos. Bot. 2:40. 1791). Type: Melinum palustre (L.) Link Me palustris LJ. Monoecious aquatic annuals or perennials. Culms scout or slender, erect or decumbent. Leaves with sheaths open, rounded dorsally; ligules mem- branous or scarious, glabrous; blades flat, linear to broadly lanceolate, api- ces acuminate to caudate. Panicles terminal, in American species the stami- nate spikelets on lower branches, pistillate on upper branches, in the Asian species Z. latifolia both types on same branches. Spikelets 1-flowered, dis- articulating from the expanded cupulate apices of pedicels, glumes absent, lemmas as long as spikelets, paleas subequal to lemmas, narrower. Stami- TERRELL ET AL., Taxonomy of North American Zizania pay) nate spikelets pendulous, caducous, lemmas and paleas membranous, loosely covering the 6 stamens, lemmas usually 5-veined, paleas usually 3-veined; anthers 3-10 mm long in our species, linear, yellow. Pistillate spikelets cylindrical, caducous, lemmas and paleas chartaceous or coriaceous, pli- cately clasped along their margins, lemmas 5-veined, terminally awned, paleas 3-veined, awnless; stigmas 1—-2.5 mm long in our species, short- plumose, at anthesis laterally exserted. Caryopses cylindrical, hilums lin- ear, embryos linear. From zézanion, an old Greek name for a weed growing in grain. KEY TO NORTH AMERICAN ZIZANIA SPECIES 1. Plants perennial; culms geniculate, stoloniferous, completely immersed or only upper parts of culms emergent; upper ligules acuminate or ee restricted to San Marcos River, Hays Co. iia _ 2. texana 1. Plants annual; culms erect, not scoloniierons: rarely completely eas upper ligules often truncate or erose, occasionally acuminate; not along San p) Marcus River, les Co., Teas 2. F rtile (bea 1 | ) piscillate inflorescences 10— B10) cm wide, with ee colts peanehies (immature inflorescences or those bear- ing only aborted spikelets may have appressed branches); lemmas of pis- tillate spikelets chartaceous (papery) and flexible, dull or sublustrous, bearing scattered short prickle hairs not or only slightly denser at lemma apices, lemmas and paleas sometimes partly separating along their mar- ).4—1 mm wide, often thread- gins at caacUltey, aborted aa spikelets 0 like; plants to 4 or 5 m tall; leaf blades (3—)10-72 mm or more wide 1. Z. aquatica Fertile pistillate inflorescences 1—10(—50 in var. snterzor) cm wide, with t of the branches upwardly appressed or in var. sterzor some- what spreading ; lemmas of pistillate spikelets coriaceous (firm and leath- ery) or indurate (hardened), lustrous, sometimes w whitish, glabrous or with hairs in lines, the hairs often densely grouped at lemma apices, lemmas and paleas tightly clasped along their margins; aborted pistil- late spikelets 0.6-2.6 mm wide; plants 1—2(—3 in var. ivterror) m tall; leaf blades 3—20(—40 in var. ivfervor) mm wide 2. Z. palustris No 1. Zizania aquatica L., Sp. Pl. 2:991. 1753. Annuals with culms 0.2 to ca. 5 m tall, erect or decumbent, often stout, glabrous, most culms emergent or rarely immersed. Leaves with sheaths glabrous or scabridulous, margins ciliate or glabrous; ligules 5-30 mm long, upper ones ovate or triangular, truncate to acuminate, often erose or irregularly lobed; blades to 1.5 m or more long, (3—)10—55(—ca. 72) mm wide, scabrous or glabrate above and beneath, margins scabrous or glabrate. Panicles 20-120 cm long, (S—)10—50 cm wide; mature staminate branches ascending to widely spreading or reflexed, pedicel apices 0.2—0.6 mm wide; mature pistillate branches widely spreading or divaricate (sometimes appressed if immature or bearing only aborted spikelets), pedicel apices 536 Sipa 173 wa 0.5—1.2 mm wide. Staminate spikelets 5—12.5 mm long excluding awns, lemmas 1—2.5 mm wide, lanceolate, acuminate or with awns to 3 mm long, paleas with awns 0—2 mm long. Mature pistillate spikelets 5-24 mm long excluding awns, 1—2.5 mm wide, lemmas lanceolate or oblong, chartaceous, flexible, dull or somewhat lustrous, with numerous scattered short prickle hairs not or scarcely denser at lemma apices, mature lemmas and paleas often partly separating along margins, awns to ca 9 cm long, scabridulous; aborted pistillate spikelets 0.4—1 mm wide, linear, shriveled, often filiform. Caryopses 6-22 mm long, 0.8—2 mm wide. 27=30. KEY TO THE VARIETIES OF ZIZANIA AQUATICA 1. Plants to 5 m tall; blades (S—)10-—72 or more mm wide; pistillate spikelets 7-24 mm long; lemma awns to 90 mm long a. var, aquatica 1. Plants 0.2—1 m tall; blades 3—12(—20) mm wide; pistillate spikelets 5— mm long; lemma awns 1—8 mm long Lb. var. brevis la. Zizania aquatica L. var. aquatica. Ceratochaete aquatica (L.) Lunell, Amer. a Naturalist 4:214. 1915. Type: without collector or data (LEcrorypE: LINN 19.1!; [Reveal, Phytologia 72:6. 1992]) Zizania clavulosa Michx., Fl. Bor. Amer. 1:75. 1803. Type: North America: A. Michaux. Hydropyrum esculentum Link, Hore. Berol. 1:252. 1827. Stipa angulata \.. ex Steud., omencl. Bot. ed. 2. 2:642. 1841, as syn. of Hydropyrum esculentum Link. Type: North America. Lizania a var. svbbrevis B. Boivin, Naturaliste Canad. 94:528. 1967. Type: CANADA. Ontario: Russell Co.: Casselman, 21 Aug 1884, J.-M. Macoun 28938 eee CAN). Common name.—Southern wild-rice Distribution.—In fresh or somewhat brackish marshes, swamps, rivers, streams, occasionally lakes. Coastal plain and locally in piedmont from New England and New Brunswick to Florida and west to southern Louisiana; from New England westward across southern Quebec and southern Ontario through the southern part of the Great Lakes region to central Ohio, northern Indiana and Illinois, southern 3/4 of Wisconsin, and rare in southern Min- nesota; rare or absent in northeastern Iowa, western Kentucky and Mis- sourt (Fig. 3). Occasionally introduced elsewhere for waterfowl food. A variant population in the Wading River area of southern New Jersey has narrow leaves and plants mostly immersed (Ferren & Good 1977; observed and collected in 1977 by Terrell and Ferren, and in 1983 by Duvall). Nar- row-leaved populations occur locally in the New England states and near Ottawa, Ontario, and individual narrow-leaved plants occasionally occur elsewhere. Figs. 1, 3 Selected representative specimens examined (US). CANADA. Onrario: pana & Lennox Cos.: Napanee, along the Napanee River, 24 Jul 1952, Dore 13840. Carlton Co. Marvelville, tributary of Castor River, 14 Sep 1973, Dore & Carteret 21157. ene on TERRELL ET AL., Taxonomy of North American Zizania Fic. 1. Zizanta aquatica var. aquatica. A. Culm leaf (Terrell 4467). Inflorescence (Terrell 4911). C. Staminate spikelet (Terre// 4459). D. Pistillate spikelet, ventral view (Terre// 4467, D-H). E. Pistillate spikelet, dorsal view. F. Aborted spikelet. G. Caryopsis showing embryo at base. H. Caryopsis showing hilum. 538 Stipa 17(3) 2 mi S$ of Marionville, Castor River, 14 Sep 1972, Dore & Carteret 25159. Essex Co.: Hillman, 19 Aug 1949, Dore 10340. Frontenac Co.: Verona, Hardwood Creek, 17 Aug L951, Dore & Mulligan 13549. Iberville Co.: Henryville, South River, 12 Aug 1952, Bassett & Hamel 2454. Leeds Co.: 10 mi SW of Brockville, Jones Creek, 25 Sep 1950, Dore 12895, Ontario Co.: 3 mi E of Port Perry, Lake Scugog, 26 Aug 1949, Dore, Senn, & Frankton 10310, Prescott Co.: E of Alfred Station, Alfred Peat Bog, 12 Jul 1955, Dore 16000. QueBEC: Missisquoi Co.: 2.5 mi N of Clarenceville, South River, 13 Aug 1952, Bassett & Hamel 2505. UNITED STATES. ALaBama. Mobile Co.: Mobile River, MoAr s.n. DeELAwArE: New Castle Co.: Wilmington, | Sep 1897, cone s.m. Sussex Co.: Nanticoke River at Wood- land, 15 Sep 1976, Terrell & Jones 4559. Froripa: Duval Co.: Freshwater creeks, Oct, Curtiss 3364. Marion Co.: Dunnellen, Withlacoochee River, 9 Sep 1898, Cambo 906. Or- ange Co.: Cold Spring Creek, 23 Sep 1898, Cambo & Baker 1139. Georgia: Bryan Co.: Ogeechee River at bridge fees Route 17, 7 Sep 1973, Terrell 4467. Camden Co.: White Oak Creek along Route 17, 8 Sep 1973, Terrel/ 4470. Glynn Co.: Alramaha River near Route 17, 8 Sep 1973, PTs McIntosh Co.: Champney River along Route 17, 8 Sep 1973, Terrell 4408, Kentucky: Reelfoot Lake, 6 Oct 1915, Alexander 308. LOUISIANA: Saint” Tammany Par.: vicinity of Covington, Sep 1919, Arsene 11596. Maine: Barnstable : Harwich, Allen’s Harbor Creek, 2 Sep 1918, Fernald & Long 16183. MARYLAND: rere Arundel Co.: marsh along Patuxent River, 26 Aug 1973, Terrell & Terrell 4460. Calvert Co.: Chesapeake Beach, 18 Aug 1912, Chase 61326. Caroline Co.: Denton, 6 Oct L973, Terrell & Terrell 4476. Dorchester Co.: 1 mi NW of Sharptown, 15 Sep 1976, Terrell & Jones 4565. Queen Anne’s Co.: Millington, 30 Sep 1973, Terrell & Terrell 4475. Massacuusetrs: Middlesex Co.: Cambridge, 26 Aug 1911, Know/ton s.n. MICHIGAN: Berrien Co.: New Buffalo, Margin of River, 20 Jul 1911, Lansing 3279. Monroe Co. shores of Lake Erie, 27 Aug 1924, Farwell hen Saint Gan Co.: Port Huron, 12 ee 1910, Dodge 12. New Jersey: Atlantic Co.: mi S of Green Bakes ilong Route 563, 24 1977, Terrell 4911. Burlington Co.: Sain River just above Route 542, 21 Jul 1974, Ferren & Good s.n. NoRTH CAROLINA: Brunswick Co.: 2 mi W of Wilmington, Brunswick River, 5 Sep 1973, ae — Chowan Co.: Edenton, 20 Jul 1938, Godfrey . Jones es W of Pollocksville, Miry Hole Branch, 9 Sep 1966, Bradley, Sears, & Bini 3407. PENNSYLVANIA: ee Co.: West Chester, Darlington s.n. SouTH CARO- LINA: Colleton Co Combahee River at bridge on Route 17, 7 Sep 1973, Terre// 4465. Jasper Co.: 2 mi NE of Coosawhatchie, Tullipinny River, 7 Sep 1973, Terrell 44606. Georgetown Co o.: Cat Island, 16 Aug 1915, Alexander 100. Ruove IsLAND: Providence o.: Diamond Hill, 26 us 1942, Palmer 46579. VirGinia: Charles City Co.: Graves Landing, oe River, 10 Sep 1941, Fernald & Long 13537. Fairfax Co.: Hunt- ing Creek, 20 Jul 1915, McAtee 2377. Staff be Co.: Potomac River at Aquia Creek, 28 Aug 1938, Hermann 9744. WASHINGTON D.C.: vicinis Washington D.C., 12 Sep 1880, Ward s.n. Roosevelt Island, 10 Oct 1958, McC/vre s.n. WISCONSIN: ara Lake Co.: Fox River, 2 mi W of Princeton, 23 Aug 1981, Terre// 5016. LaCrosse Co.: 4 mi E of LaCrosse, 22 Aug 1974, Terrell 4487. 2.7 Ib. Zizania aquatica var. brevis Fassett, Rhodora 26:157. 1924. Z. aquatica subsp. brevis (Fassett) S. L. Chen, Proc. Int. Symp. Bot. Gard., oe China 600. 1990. Type: CANADA. Quebec: rocky tidal flats of St. Lawrence River, Levis, 9 Aug 1923, H.K. Svenson G N.C. Fassett 853 (HoLoryre: GH; fragment US 2797822!: photo Dore 1969: 18) Estuarine wild-rice. Common name. TERRELL ET AL., Taxonomy of North American Zizania Be! Distribution.—Occurs only in tidal mud flats along the upper St. Lawrence River in Quebec about 80 km upstream and downstream from Quebec City. Fig. 3. Representative specimens examined (US). CANADA. Quesec. Tidal flats, er 23 Aug 1947, Swallen 9703; just west of Pierre Laport Bridge, Anse Grigras, 1 L983; Darbyshire 2030. Levis Co.: § of ferry dock along St. Lawrence, 13 Aug oe olsers 2031. Lotbiniere :U er tidal flat of St. Lawrence estuary at Ste. Antoine des Fonds, 31 Aug 1947, a 971. Portneuf Co.: estuarine marsh of St. Lawrence, Portneuf, 9 Sep 1976, Dore 25522. Quebec Co.: upper tidal beach of estuary of St. Lawrence, Beauport Flats, 27 Aug 1975, Dore 25471. 2. Zizania palustris L., Mant. Pl. 2:295. 1771. Annuals with culms to 2(—3 in var. /vterior) m tall, erect, glabrous, part or most of culms immersed. Leaf sheaths glabrous or with scattered hairs, margins glabrous; ligules 3-16 mm long, lanceolate to triangular, truncate or erose; blades 20—60 cm or more long, 3—20(—40 in var. interior) mm wide, glabrous, margins glabrate or scabrous. Panicles 24—60 cm long, to 20(—40 in var. interior) cm wide; mature staminate branches of panicle 3— 20 cm long, branches ascending or spreading, pedicel apices 0.2—0.4 mm wide; mature pistillate branches of panicle 1—8(—15) (—50 in var. znterzor) cm long, appressed or ascending or 1—few branches spreading (to widely spreading in var. znterior), pedicel apices 0.7—1.2 mm wide. Staminate spike- lets 6-17 mm long excluding awns, lemmas 1—2.3 mm wide, lanceolate, acuminate or with awns to 2 mm long, paleas with awns 0—2 mm long. Mature pistillate spikelets 8-22(—33) mm long excluding awns, 1—2.6 mm wide, lemmas lanceolate or oblong, coriaceous or indurate, lustrous, some- times whitish, glabrous or with short prickle hairs in lines, these hairs often more densely aggregated at lemma apices, mature lemmas and paleas remaining firmly clasped along their margins, awns to ca 9.3 cm long, hirsute at bases to scabrid distally; aborted pistillate spikelets 0.6-2.6 mm wide. Caryopses 6—30 mm long, 0.6—2 mm wide. 27=30. Fig. 2. KEY TO THE VARIETIES OF ZIZANIA PALUSTRIS —_ . Leaf blades 3—15(—20) mm wide; lower pistillate branches with 2-8 spikelets; pistillate inflorescences usually 1-10 cm wide, with appressed or ascending branches; plants to 2 m tall 2a. var. palustris Leaf blades (10—)20—40 or more mm wide; lower pistillate branches with 9-30 spikelets; pistillate inflorescences (S—)10—50 cm wide, with branches ascending to widely spreading; plants 1-3 or more m tall ................. 2 2b. var. interior _ 2a.Zizania palustris L. var. earns Mant. Pl. 2: 295. 1771. Melinum palustre eee Handbuch 1:96. 1829. Type: garden plant grown at Uppsala (LEc- ToTYPE: LINN 1119.2!) [Reveal, Seeaes 72:7. 1992). Z. aquatica var. angustifolia Hitchc., Rhodora 8:210. 1906. Z. aquatica subsp. angustifolia 540 Sipa 17(3) Fic. 2. Zizania palustris var. palustris. A. Calm base (Terrell 4496). B. Inflorescence (Terrell 4515, B-G). C. Pistillate spikelet, ventral view. D. Pistillace spikelet, dorsal view. E. Aborted spikelet. F. Caryopsis showing hilum. G. Caryopsis, side view. TERRELL ET AL., Taxonomy of North American Zizania 541 s n. Fic. 3. casei aecahacen: of Zizania. aquatica var. aquatica (A) and Z. aquatica vat. brevis (B). (Hitchc.) Tzvelev, Novosti Sist. Vyssh. Ras 1971. Type: U.S.A. MAINE: es Aug 1895, FE Lamson- ae s.n. nes oTyPe: US 80339!) Zz: onli forma elas Dore in McNeill & Dore, Naturaliste Canad. 103:565. 1976. E: CAN ARIO: Russell Co.: in Castor River 2 mi E of Embrun, 21 Sep ee WG. Dore e RY. Moore 10587 (HOLOTYPE: DAQ). Common name.—Northern wild-rice Distribution.—In shallow water of lakes and streams, often forming ex- tensive stands in northern lakes; Nova Scotia (introduced), New Brunswick, southern Quebec, and New England states westward across the Great Lakes states, central and southern Ontario, to central Alberta (range extended westward by planting fide Aiken et al. 1988), the Dakotas, and Nebraska. Occasionally planted elsewhere for waterfowl food, known to have been introduced but probably not — cin in British Columbia, Ari- zona, West Virginia, and Idaho. F Selected representative specimens examined (US). CANADA. MANnirosa: near Winnipeg, 54? Sipa 17(3) 30 Jul 1933, Chambliss 391. New Brunswick: Westmoreland Co.: Aulac, Chignecto Isthmus Marsh, 21 Aug 1945, Dore & Ingalls 45-1068. ONrario: Hemlock Island, Geor- gian Bay Islands, 25 Jul 1942, McDonald 257. Roundtable Lake, Hardwick Township, 20 Aug 1951, Garton 1627. Addington, Co.: 3 mi SW of Napanee, Napanee River, 16 Aug L951, Dore G hee Sel 13540. Carleton Co.: Jock River, 11 Sep 1950, Dore oe Frontenac i 8 mi SW of Arden, Salmon River, 14 Aug 1951, Dore & Mulligan 13 Leeds Co.: 10 mi SW of Brockville, St. Lawrence se 25 Sep 1950, Dore 1 ee eres Co.: te Bonnechere River, 18 Aug 1952, Cody & Parmelee 6598. Victoria Co.: 2 m E of Sebright, Head River, 20 Aug 1952, C a & Parmelee 0082. QUEBEC: W of ne Lacolle Creek, 6 Aug 1952, Dore & Cody 13918. Noyan, Richelieu River, 6 Aug 1 Dore & Cody 13919, Chambly Co.: Longueuil, 21 Sep 1 1924, Victorin 18258. Labelle Co.: 2 Petite River, 3-5 Aug 1941, Marie-Victorin, Rolland-Germain ie oon . Nicolet : W of Becancour, St. Lawrence ee 983, Darbyshir U ne DS TATES. INDIANA: LaGrange Co.: 2 mi E of On Wtario, Pigeon a 30 Aug 1914, Deam 15045. Steuben Co.: Lime Lake, 19 aes 916, Deam 20913. White Co.: 3 mi S of Buffalo, 24 Aug 1919, Deam 29542. me Joliet, 22 es 1904, Skeels 462. lowa: Dickinson Co.: Lower Gar Lake, 15 Aug 1 , Thorne 18599, Fayette Co.: 18 Jul 1894, Fink 180. Maine: Franklin Co.: Farmington, “al 1899, Knowlton 314. Hancock Co.: Long Island, Aug 1872, Young s.n. : — Co.: Keg Stream, 28 Aug 1934, Walker 2032. Sagadahoc Co.: Bowdoinham, 14 1916, Fernald & Long 12580. aren Co.: Princeton, 2 Sep es C sinner a re HuSETTS: Concord, Sudbury River, 21 Aug 1904, Worthen s.n. Mict : Allegan Co.: Kalamazoo River above Allegan, 12 Sep 1902, Wight 124. Emmet on rete river at Devil’s Elbow, 9 Aug 1932, Gates i POC. Huron Co.: near Caseville, Saginaw Bay, 9 Sep 1910, Dodge 14. Kalamazoo Co.: SW of Vicksburg, Barron Lake, 21 Jul 1938, Rapp 2178. Saint Clair Co.: Lake Saint C - 12 Aug 1910, Dodge 13. MINNESOTA: Cass Co.: S of Deer River, Mississippi River, 31 Aug 1975, Edie )7. Clearwater Co.: Lake fue, 6 Aug 1929, Grant 3016. Mille Lacs Cox Mille Lacs Lake, 3 Sep 1975, Terrell 4515, Otter Tail Co.: East Battle Lake, Aug oe Gide sm. NEBRASKA: Bone Creek, 29 Aug 1893, Clements 2921; near Halsey, 6 Aug 1934, Martin 439; Rat Lake, 14 Aug 1915, Thomson 51. New York: Clinton Co.: Plattsburg, Lake Champlain, 22 Aug 1917, Hitchcock 16018. Jefferson Co.: Woodville, along Lake Ontario, 23 Jun 1921, Howse 8217. Oswego Co.: Selkirk, Salmon River, 4 Aug, Clausen & Hinkey 4289. St. Lawrence Co.: Waddington, along the St. Lawrence, 22 hes 1935, House 20744. Nortu Dakora: Hep Co.: Tn ee James river, 22 » L913, Lunell s.n. SourH Dakota: Beadle : Huron, edge of James River, 23 Aug 196, . Griffis 118. Brookings Co.: aa Tal 1892, Williams s.n. /VeKsion West 18 Aug 1913, Kirk 1042. Wisconsin: Sawyer Co.: Couderay Lake, 4 Aug 1930, cone 08. oe} 2b. Zizania palustris var. interior (Fassett) Dore, Publ. Dep. Agric. Canada 1393:20. 1969. Z. aquatica var. interior oa Rhodora 26:158. 1924. Z. inte- rior (Fassett) Rydb., eninge 1:82. L931. Z. palustris subsp. tnterior (Fassett) S. L. Chen, Pro. Int. Symp. . Gard., Nanjing, China 600. 1990. Type: U.S.A. lowa: Emmet Co.: eee 7 Aug 1897, L. H. es R. 1, Cratty 764 (HOLOTYPE: GH; tsorype: US 328087!; photo Dore 1969: | Common name.—Interior wild-rice Distribution.—Sporadically in eastern Canada and westward across the Great Lakes region to southern Manitoba and central Alberta (range ex- tended westward by planting; see Aiken et al. 1988), Nebraska, Kansas (?), TERRELL ET AL., Taxonomy of North American Zizania 543 Minnesota, Iowa, Wisconsin, Missouri, and southward (by planting?) in Illinois, Indiana, and Ohio). Fig. 4, 5. Selected representative specimens examined (US). CANADA. Maniropa: Starbuck, 20 mi W of Winnipeg, La Salle River, 16 Aug 1950, Dore 12734. ONTARIO: Northumberland Co.: 3 mi W of Hastings, Rice Lake, 3 Sep 1952, Dore & Hammond 13981. Russell Co.: 3 mi W of Cumberland, Ottawa River, 18 Aug 1953, Jenkins & Dore 3532. QUEBEC: Chateaguay Co.: San River, Aug 1941, Boivin 4425. UNITED STATES. Illinois: Henderson Co.: near Oquawka, 1871, Patterson s.n, Madi- n Co.: Margins te 8 Sep 1891, Esuer 280. Ree ee Co.: N of Spring Bay, 12 Sep oe Chase 6181. INDIANA: Fulton 1 mi SE of Grass Creek, Grass Creek, 9 Aug 1928, Deam 46049. Iowa: Cook je Des ae River, 10 Aug 1901, Chase ona Hamilton Co.: 3 mi S of Jewell, Little Wall Lake, 16 Aug 1936, Hayden 10325. Linn Co Cedar Rapids, 20 Jul 1895, B. Shimek 68. M1nNEsoTa: Fort Snelling, Minnesota River, 28 Aug a ee hee Chippewa Co.: epee Chippewa River, 28 Jul 1895, Moyer 25. Itase a Co: nd of Bass Lake, 2 mi NNW of Cohasset, 31 Aug 1975, Terre// 4510 ee Co.: 3 mi . of Pine City, 24 Aug 1974, a 4491, Wabasha Co.: river along rt N of Weaver, 25 Aug 1981, Terre// 5020. NEBRASKA: Grant Co.: 15 mi S of Whitman, : ye 1893, Tulen 1630. Wisconsin: Buffalo Co.: slough in backwaters of Mississippi River, 1/2 mi SSW of Nelson, 4 Sep 1975, Terre// 4516. Trempealeau Co.: delta of Trempealeau River, Perrot State Park, 24 Aug 1981, Terrell 5017 3. Zizania texana Hitchc., J. Wash. Acad. Sci. 23:454. 1933. Type: U.S.A, Texas. Hays Co.: San Marcos River near San Marcos, Apr 1932, W. A. Silveus 518 (HOLOTYPE: US-1537174!; isotypes: K! US-1720531!). Perennial with culms 1—2(—5) m long, decumbent, geniculate, rooting at lower nodes, stoloniferous, long-streaming in river a or the upper parts of culms emergent. Leaf sheaths glabrous; ligules 4-12 mm long, caudate or acuminate; blades to ca 1 m long, to 13(-25) mm wide, glabrous. Panicles 16-31 cm long, 1-10 cm wide; mature staminate branches as- cending or somewhat spreading, pedicel apices ca 0.3 mm wide; mature pistillate branches appressed or ascending, pedicel apices 0.5—0.9 mm wide. Staminate spikelets 6.5—11 mm long excluding awns, lemmas |.2—2 mm wide, ovate or oblong, acute to acuminate, paleas usually awnless. Mature pistillate spikelets 9-12.5 mm long excluding awns, 1.2—1.8 mm wide, lemmas lanceolate, somewhat coriaceous, somewhat lustrous, with scat- tered short prickle hairs, apices scabrous or hispidulous, awns 9-35 mm long, with scattered hairs; aborted pistillate spikelets 0.7—1.5 mm wide. Caryopses grown in cultivation 4.3-7.6 mm long, 1—-1.5 mm wide, 1/2—3/ 4 as long as their lemmas. 2”7=30. See Fig. 814 in Hitchcock (1951). Common name.—Texas wild-rice Distribution.—This species is restricted to a 2.4 km length of the head- waters along the San Marcos River, within the city limits of San Marcos, Hays Co., in southcentral Texas (Terrell et al. 1978). The neutral or slightly alkaline (pH 6.9-7.8) water temperature varies only 5° (20.4—25.5) C an- nually. Officially listed as an endangered species. 544 Sipa 17(3) q fh | il ) | i Teh poe i iN vat SSRs a | aa LARA rt yee aceon gill : ‘i \ A Di Mt oS Me Pc / { a sul gy : ie { pet y | , oan Pref v Fic. 4. Geographic distribution of Zizania palustris var. palustris (A), Z. palustris var. inte- rior (B), and Z. texana (C). Central region shows overlap of A and B. ~ Representative specimens examined. UNITED STATES. Texas: Hays Co.: | mi below Spring Lake, San Marcos River, 14 Sep 1973, Terrell, Emery & Beaty 4472 (US); San Marcos, Aug 1892, Nealley 91 (MO, US); San Marcos, 6 Nov 1897, Tre/ease s.n. (BM, MO); “Marcos River,” 31 May 1938, Sz/vews 2731 (BM); San Marcos River, 14 Jan 1936, Hotchkiss s.n. (BM, MO). ExcLUDED NAMES Zrzania effusa Munro, J. Linn. Soc., Bot. 6:52. 1862, nom. nud. Munro alluded to a speci- men of Phragmites australis (Cav.) Trin. ex Steudel (LINN, 1119.3!), annotated by J. E. Smith with this name. Other excluded Z7zania names were listed by Fassett (1924). DISCUSSION In the past Zizania palustris (or Z. aquatica var. angustifolia) was gener- ally included in Z. aquatica , the older name, in most publications (e.g., TERRELL ET AL., Taxonomy of North American Zizania 545 Fic. 5. Zizania palustris var. interior. A. Culm leaf (Terre// 4516, A-D ). B. Inflorescence. C. Pistillate spikelet, ventral view. D. Pistillate spikelec, dorsal view. 546 Stipa 17(3) Fassett 1924). However, Wiegand and Eames (1925) listed both Linnaean species in their flora of the Cayuga Basin. Dore (1969) recognized the two species in his monograph of the Canadian wild-rices. The two distinct spe- cies are borne out by morphological data: They differ in size of the plants, in leaf width, in size and branching of the inflorescence, and the lemmas and paleas differ in texture and thickness, with prickle hairs differing in their arrangement (see key to species). The caryopses differ slightly in length and thickness, but measurements overlap (see photographs of caryopses 1n Dore (1969), Dore & McNeill (1980), and Aiken et al. (1988)). The two species also differ in ligule size and shape, but these differences can over- lap. Terrell and Wergin (1981) found differences in the shapes of the silica bodies in the lemmas: Z. aquatica usually has C-shaped silica bodies, whereas Z. palustris has suborbicular silica bodies with 1—3 shallow sinuses. Present data on distribution show that the two species have distinct but overlapping ranges (Figs. 3, 4) with Zizania palustris the wild-rice espe- cially of northern lakes and Z. aquatica centered in the southeastern Atlan- tic and Gulf Coastal Plains. The marked differences in the spikelets of Z/zania aquatica and Z. palustris were well-known to Fassett (1924) and undoubtedly to others (see the early literature cited by Aiken et al. 1988). Duvall and Biesboer (1988a), found an anatomical basis for the textural differences in the lemmas and paleas, there being a single layer of thin-walled subepidermal fibers in 2. aquatica, whereas Z. palustris (with its coriaceous texture) has at least two layers of thick-walled fibers. Warwick and Aiken (1986), in an electrophoretic study, concluded that genetic identity values between Zizania aquatica and Z. palustris were on levels expected for congeneric species. They did not find any isozyme evi- dence for gene flow between the two species. However, Duvall and Biesboer (1988b), in a study of artificial hybridization, found that hybrids were pro- duced (at a low level) only when Z. aquatica was the female parent. These authors concluded that (1) the “near-unilateral crossability barrier” sup- ported the taxonomic distinctness of the two species, and (2) the successful artificial hybridization (although limited) shows the potential for success- ful natural hybridization. Zizania aquatica var. brevis, native to a portion of the St. Lawrence River, has been studied in detail by Darbyshire and Aiken (1986). Warwick and Aiken (1986) found no isozyme differences with Z. aquatica vat. aquatica. Similarly, Duvall and Biesboer (1989) found no distiguishing seed storage protein differences between var. aquatica and var. brevis. These two varieties were also found to be fully interferctile under greenhouse conditions (Duvall & Biesboer 1988b), supporting their recognition as varieties rather than as species. TERRELL ET AL., Taxonomy of North American Zizania 547 In 1924, Fassett described Zizania aquatica var. interior, which resembles Z. aquatica in leaf width and inflorescence characters, but Z. palustris in the characters of the pistillate spikelets. Dore (1969) transferred var. interior to Z. palustris, and in this we follow Dore. Like Z. palustris, plants of var. interior have been purposely introduced to wildlife refuges and private wild- life preserves; consequently, it is sometimes difficult to determine whether a population of Interior wild-rice occurred naturally or introduced. Aiken et al. (1988) maintained var. terior as distinct, but noted that it is some- times difficult to separate var. palustris from var. interior, and on their map of Z. palustris in Canada they did not attempt to separate them. Zizania palustris var. interior occurs mainly where the ranges of Z7zania aquatica and Z. palustris overlap (Figs. 3, 4); consequently it has been hy- pothesized that var. znterior includes hybrids (Dore and McNeill 1980). Herbarium specimens of var. ivterior are mostly concentrated in the central United States. Field studies by the senior author in 1974, 1975, and 1981 in the Mississippi Valley of western Wisconsin and southeastern Minne- sota produced several collections of var. swterior. Fassett (1927) saw it “grow- ing abundantly along the Mississippi River from Lake Pepin to Fountain City”; certain of these specimens exist at WIS. One of the Terrell collec- tions (4516 at US) included a range of morphological expressions varying from aquatica-like plants with broader leaves and more spreading pistillate branches to palustris-like plants with narrow leaves and compact appressed inflorescences (Fig. 5). These plants were like Z. palustris in spikelet mor- phology, thus agreeing with Fassett’s (1924) original description of var. interior. Cultivated strains of Z7zania palustris have been selected by Minnesota growers for retention of mature spikelets to facilitate harvesting. In 1975, Terrell collected samples from several cultivars in Minnesota (vouchers at US). These were generally similar to Z. palustris, except in having some individuals bearing pistillate inflorescences with one or a few spreading or divergent branches, a condition called “crowfoot” by growers. Such plants are somewhat similar to Z. aguatica, which has spreading pistillate inflo- rescences. The electrophoretic study of Warwick and Aiken (1986) sug- gests that “Cultivated varieties, previously considered to be improved strains of Z. palustris var. palustris, were found to be more closely related to Z. ” palustris var. interior ... If var. zzterior is of hybrid origin we point out that the combination Z. x interior (Fassett) Rydberg (pro sp.) (Arc. 50, Greuter et al. 1994; see also the present synonymy) is available for use. We also note that given not only the long expanse of evolutionary history but also the Pleistocene glaciations in particular it seems possible that var. ‘nterior could be the result of past introgressive hybridizations. 548 Stipa 17(3) We emphasize, however, that some other evidence concerning var. /fe- rioy argues against its origin as an interspecific hybrid. As noted above, Warwick and Aiken (1986) found no isozyme evidence of introgression between the annual species. A preliminary chloroplast DNA restriction sites study also failed to indicate that var. ivterior could be of hybrid origin (Duvall 1987). Even though an artificial hybridization study indicated that hybrids can be produced between the annual species, this study also showed virtually no difference in crossability between the two varieties of Z. palustris with those of Z. aquatica (Duvall & Biesboer 1988b). If var. interior were, in fact, of hybrid origin, one would expect it to (back) cross more readily with Z. aquatica, than does var. palustris. These data, coupled with the observa- tion that all of the artificial F; hybrids that were produced in the hybridiza- tion study displayed the diagnostic spikelet characters of the Z. aguatica parent, is compelling evidence against the hypothesis of hybrid origin. These data suggested that vegetative similarities between Z. palustris var. interior — and Z. aquatica vat. aquatica are possibly an overlap of a morphological gradient of plastic and highly variable characters. Unlike other North American Zizania species, Zizania texana behaved as a perennial in greenhouse experiments by Duvall (1987). A close rela- yonsye between Z. aquatica and Z. texana is supported by storage protein data (Duvall & Biesboer 1989) and distribution. However, crosses of Z. texana (ovulate parent) with three other taxa: Z. aquatica var. aquatica, Z. palustris var. palustris, and Z. palustris var. interior, resulted in hybrid prog- eny in only one case, that of Z. texana x Z. palustris var. interior (Duvall 1987). The pistillate spikelets of these hybrids were somewhat intermedi- ate, unlike those of any other interspecific hybrid. Thus, full resolution of the relationships between annual and perennial species of North American wild-rice awaits further study. ACKNOWLEDGMENTS We wish to acknowledge the help of Mary Barkworth for advice on taxo- nomic descriptions. Alice Tangerini prepared the illustrations. Robert Soreng contributed helpful reviews of the manuscript. Work on this project is supported in part by Project F-12 of the Maryland Agricultural Experi- ment Station and Cooperative Extension Service. REFERENCES AIKEN, S.G. 1986. The distinct morphology and germination of the grains of two species of wild rice (Zizania, Poaceae). Canad. Field-Naturalist 100:237-240. P.F Ler, D. Punrer, and J.M. Srewarr. 1988. Wild rice in Canada, Agricul- ture Canada Publ. 1830. New Canada Publ., Toronto. 130 pp. DARBYSHIRE, S.J. and S.G. AIKEN. 1986. Zizania aquatica var. brevis (Poaceae): A 1983 TERRELL ET AL., Taxonomy of North American Zizania 549 distribution survey and a scanning elec ron microscope study of epidermal features. Naturaliste Can. (Rev. Ecol. Syst.) 113:355—360. Dore, W.G. 1969. Wild rice. ee Dept. ‘Agric Publ. 1393. Information Canada, Ottawa. and J. McNeill. 1980. Grasses of Ontario. Biosystematics Research Insti- tute, Research Branch, pee Canada Monograph 26 Duva.t, M.R. 1987. A systematic evaluation of the genus Z/zania (Poaceae). Dissertation, University of Minnesota, St. Paul. and D.D. Biesporr. 1988a. Anatomical distinctions eee the pistillate spikelets of the species of wild-rice neem Poaceae). Amer. J. Bot. 75:157-159. —______ and D.D. Birsporr. 1988b. Nonreciprocal byberdizabicn ae in crosses between mee wild-rice species (Zizania palustris x Z. aquatica: Poaceae). Syst. Bot. 13:22 34, nd D.D. Biespoer. 1989. Comparisons of el ECHRODHOESUE $8 seed pole - files among North American populations of Zizania. Biochem. Syst. 1 —_______, P.M. Perrerson, E.E. TERRELL, and A.H. CHRISTENSEN, 1993, Seen of North American ery: grasses as construed from maps of plastid DNA restriction 30; sites. Am Fassett, N. C. 1924. A ae of the genus Zizania. Rhodora 26:153-160 1927. Notes from University of Wisconsin—1. Rhodora 29: 228-229, FERREN, W.R., Jr. and R.E. Goon, 1977. Habitat, morphology, and phenology of southern wild rice (Zizania aquatica L.) from the Wading River in New Jersey. Bull. Torrey Bot. Club 104:392—396. Greuter, W., ER. BARRE, H.M. Burpbet, W.G. CHaLoner, V. DEMOULIN, D.L. HawkswortnH, M. JorGENSEN, D.H. Nicotson, P.C. Sitva, P. TREHANE, and J. MCNEILL (eds.). 1994. International code of botanical nomenclature. Regnum Veg. 131. Hircucock, A. S. (2nd ed., revised by A. Chase). 1951. Manual of the grasses of the United States. U.S. Dept. Agric. Misc. Publ. 200. Washington, D.C TERRELL, E.E. and H. Rosinson. 1974. Luziolinae, a new subcribe of oryzoid grasses. Bull. Torrey Bot. Club 101:235—245 oe es P. EMery and H. E. Beaty. 1978. ot ee on Zizania texana (Texas wildrice), an endunwenda species. Bull. Torrey Bot. Club 105:50—-57. and WP. Wercin. 1981. Sure features ne ree ‘oa in lem- mas and awns of Zizania (Gramineae). Amer. J. Bot. 68: eau L.R. Batra. 1982. Zizania latifolia: and Usti ulenta, a grass-fungus and association, Te Bot. 36:274—285. Warwick, S.I. and $.G. AIKEN. 1986. Electrophoretic evidence for be eae of two species in annual he rice eee Poaceae). Syst. Bot. 11:464— WIEGAND, K.M A.J. Eames. 1925. The flora of the ee ia Bak New York. Cornell ore a Exp. on Mem. 92. Sipa 17(3) BOOK REVIEW CHEATHAM, SCOOTER and MARSHALL C. JOHNSTON with LYNN MARSHALL. 1995. The Useful Wild Plants of Texas, the Southeastern United States, the Southern Plains, and Northern Mexico. Volume I. Abronia thru Arwndo. ISBN 1- 887292-01-2, hbk). Useful Wild Plants, Inc., 2612 Sweeney Lane, Austin, TX 78723, U.S.A. $125.00. xxi+ 568 pp. 304 color photos, 267 distribution maps. This first of twelve volumes provides encyclopedic treatment of 78 genera and 267 species of native and wild growing plants of Texas and of botanically, read ecologically, related adjacent areas. Useful applies to a plants utility as food, natural products, medi- cine, fiber, landscape or aesthetic value, oil, folk use, etc. Each genus is treated alphabeti- cally and includes a description of the ‘group’ and each taxon, botanical and common nomenclature, plant identification, habitat, a detailed rangewide distribution map, EX- CEL color photograph(s), a broad category of uses as appropriate for the taxon, in- cluding ethnobiology, author experience by category of use, literature cited, index and more. The treatments are well written and easy to read and understand. Marginal notes with keywords are provided for ease of location of subjects of particular interest. Unlike most i encyclopedias, the overall design is aesthetically appealing and wholly supports the user friendliness of the volume. Useful Wild Plants is the firse fruic of a labor of love that spans at least two decades. According to the authors and publisher, six additional volumes are ready for press; but, with financial constraints only one or, hopefully two!, will be published per year. I, for one, am confident this daunting task will be accomplished. Anyone interested in Texas’ natural history, botany, ethnobiology, folk use of plants, conservation of Texas’ natural heritage, natural plant products, traditional medicine, eco- nomic botany, plant propagation, or native plant landscaping will want to own this vol- ume and those to come. This is a must have volume for every university and botanical pias ae Sea every major city or county libra Useful Wild Plants is authoritative, copetinsne and thoroughly researched by an army of Penne cooperators and volunteers. The distribution maps are, in my opin- ion, the bese available for the treated species. The ai pin raphs ARE the best. In support of the production of the Useful Wild Plants of Texas encyclopedia, the au- thors have established a not-for profit corporation, Useful Wild Plants, Inc. Membership supports publication of the remaining volumes, includes the Useful Wild Plant Newslet- ter usually issued ‘seasonally,’ and entitles members to discounts on volumes to be pub- ished, and supports “Botany for the real World.” Further information can be obtained ae the publisher or from uwp@jumpnet.com. (See also Sida 16(3):609. 1995). David Rik punt i Program, Texas Parks and Wildlife, 4200 Smith School Rd. Aus- tin, TX 7 mee Sipa 17(3): 550. 1997 PASSIFLORA CERASINA, A NEW SPECIES OF PASSTFLORA SUBGENUS PASSIFLORA (PASSIFLORACEAE) FROM FRENCH GUIANA HILAIRE ANNONAY “Le Ramses,” 5 Avenue Desambrots 06000 Nice, FRANCE CHRISTIAN FEUILLET Department of Botany, MRC-166 Smithsonian Institution Washington, DC 20560-01606, U.S.A. ABSTRACT Passiflora cerasina, sp. nov., 1s newly described and placed in Passiflora subgenus Passiflora. This species has been found in northeastern French Guiana on Montagne de Kaw, and in Surinam on Mt. Brownsberg. Passiflora cerasina belongs in the P. laurifolia group, more precisely Cac he few species that have the first row of corona filaments shorter than the second row. RESUME Passiflora cerasina, esp. nouv., est décrite et placée dans Passiflora sous-genre Passiflora. Cette espéce a été trouvée dans le nord-est de la sais frangaise sur la Montagne de Kaw et au Surinam sur le Mont Brownsberg. Passiflora cerasina appartient au groupe de P. /aurifolia, . oti elle est proche des espéces dont la courone a les filaments du rang externe plus courts que ceux du second ran A new species Passiflora cerasina has been collected by the senior author on Montagne de Kaw (type) in the NE region of French Guiana. Montagne de Kaw (Kaw Mountain), French Guiana, is a narrow table mountain, with a lateritic crust on top. It is about 35 miles long and is a locality rich in species of Passiflora (Feuillet 1989, 1994), now with 19 known species. The new species has been discovered, though not collected, in Surinam on Mont Brownsberg at an elevation of about 500 m (30 Dec 1996). Passiflora cerasina H. Annonay & C. Feuillet, sp. nov. (Fig. 1). TYPE: FRENCH GUIANA. Spe de Kaw, road from Roura to Kaw, PK 6.3, 30 km SE of Cayenne, 4°60'N, O'W, 75-100 m, 21 Apr 1997, H. Annonay ; (HOLO- type: US). cies haec in subgen. Passiflora pertinens; folia integerrima; stipulae linearia; bracteae Pe enemies ibrae 1 integrae foliaceae rubrae : glan a dulosae; sepala et petala rubra; mesocarpium crassum, endocarpium rubrum, placentae candidae. Sipa 17(3): 551-554. 1997 A era ho Sipa 17(3) | 0.3.cm Fic. | — Passiflora cerasina Annonay & Feuillet, sp. nov.: a. Leaves. b. Petiole. c. Flower. d. Fruit. e. Longitudinal cut of the fruit. f. Seed. Scale: ae. | cm. f. 0.3 cm (after the type). ANNONAY AND FEUILLET, Passiflora cerasina, a new species from French Guiana 553 Semiwoody climber with tendrils, glabrous throughout. Leaves simple, alternate; stipules linear, 3-5 mm long, pale green, early deciduous; peti- ole 2.0—2.2 cm long, canaliculate, bearing one pair of sessile glands near the middle, glands hemispherical-elongate, 2 x 2 x 2 mm, green; lamina elliptic, 13-15 x 8.0-8.5 cm, upper surface dark green and shiny, margin entire, darker or somewhat reddish, rounded to cuneate at base, apex acumi- nate, acumen slightly curved, venation brachidodrome (looped), with 6—8 pairs of secondary veins, these prominent above and beneath, at lamina base the Ist pair marginal, the 2nd pair 3 mm from the margin, in the basal half of the lamina the minor lateral veins are perpendicular to the midrib. Flowers hanging, one per leaf axil, fragrant; peduncle ca. 2.2 cm long (elongating to about 5 cm in fruit), bearing three bracts; bracts ovate, verticillate, sessile, with marginal, yellow, sessile glands, 4.2-4.8 x 2.2— 2.9 cm, apex rounded in flower, emarginate in fruit, cherry-red in flower, dark brownish violet speckled with dark green and with medium green margin in fruit; floral tube wider than long; sepals cherry-red, ca. 4 x 0.8— 1.2 cm, bearing a curved, subterminal, yellowish awn, 0.7 cm long; petals cherry-red, ca. 4 x 0.4-0.6 cm wide; corona in several rows, the 2 outer rows of thick filaments, outermost row of filaments mostly white, with violet banding in the basal two thirds, curved upward, 2.0—2.5 cm long, the second outer row of filaments longer than the outermost row, 4.7—5.2 cm long, mostly violet with white banding in the basal half, curved down- ward forming a bell-shaped shade around the androgynophore, inner rows much shorter; stamen filaments flat, perpendicular to the androgynophore, yellowish white with light purple at apex, anthers dorsifixed, perpendicu- lar to the filament and to the androgynophore, when the anther opens the whitish pollen is facing the throat of the floral tube (up on a hanging flower), ovary yellowish white, with 3 clavate apical styles, styles yellowish striped with light purple, stigmas whitish. Fruit obovoid, looking like a small quince, 4.5-5.0 x 3.9-4.2 cm, green with yellowish markings when un- ripe, dark purple densely freckled with pale green when ripe, soft due toa spongy mesocarp about 0.5—1 cm thick, endocarp cherry-red, the 3 placentar zones white and bearing red ovules; seeds elliptic, coarsely reticulate, ca. 6 x 5mm, nearly flat, 1.5 mm thick the middle, canaliculate at margin, each half crenate at margin, brown, in a white, juicy aril. Distribution.—Passiflora cerasina is docamented only by the type collected on Montagne de Kaw, in the slope forest at an elevation of 75-100 m. It has also been observed by the senior author on top of Mt. Brownsberg in Surinam at about 500 m elevation. Passiflora cerasina is the fifth species of the P. /aurifolia group to be found on Montagne de Kaw. The other species from Montagne de Kaw are P. /aurifolia L. and P. nitida Kunth with distri- bution areas much broader than the Guianas, and P. crenata Feuillet & 554 Sipa 17(3) Cremers and P. rufostipulata Feuillet found only in NE French Guiana. In addition, P. acuminata DC., of the same group, has been seen between Cay- enne and Montagne de Kaw by the junior author. With six out of 21 spe- cies of the P. /aurifolia group being known from NE French Guiana, three of them only known from the area, the coastal table mountains of NE French Guiana are the probable center of diversification for this species group. Systematics. —Passiflora cerasina belongs to Passiflora subg. Passiflora (lec- totype: P. incarnata L.) and is part of what Killip (1938) called “ser Laurifoliae” (nom. nud.: without type and without Latin description). Ic can be inserted into Killip’s key between P. capparidifolia Killip (sp. # 209) and P. laurifolia (#213), among species with the outermost row of corona filaments shorter than the second row. In this group, Passiflora cerasina is caracterized by the combination of its floral color pattern, fruit shape, and endocarp color. Within this group it differs also from P. capparidifolia (# 209) and P. popenovii Killip (# 210) by its broadly elliptic leaves, from P. nigradenia Rusby (# 211) . its shorter peduncules and larger bracts and owers, from P. ambigua Hemsl. (# 212) by the marginal glands on its bracts, from P. /aurifolia (# 213) by its larger leaves with very different venation, and from P. rvfostipulata (described recently) by its colored, con- cave bracts and by the shape and color of the stipules. As is common in the P. lauvifolia group, the end of lateral branches tend to be hanging and look like inflorescences with reduced leaves and shortened internodes. Etymology.—The epithet “cerasina” (= cherry red) refers to the color of the bracts, sepals, petals, endocarp, and ovules of the new species. The cherry red floral parts contrasting with the corona filaments banded white and purple-violet are reminiscent of the Peruvian P. phoenicia Lindley in the closely related P. guadrangularis grou — S) REFERENCES FEUILLET, C. 1989. ela and distribution of Guianan Passifloraceae. In: L.B. Hotm- , LC. Nigtsen and H. Bats ev, eds. — forests, botanical dynamics, specia- 18 Qo tion ae diversity. ee ieee Press. Pp. 311-3 Pruner, C. 1994, Two new species of Passfora (Passifloraceae) from French Guiana. Novon 4:23 36. 341. Kitup, E. P. 1938. is American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1-613. CALCARATOLOBELIA (CAMPANULACEAE): A NEW GENUS OF SPURRED LOBELIOIDS FROM MEXICO AND CENTRAL AMERICA ROBERT L. WILBUR Department of Botany Duke University Durham, NC 27708, U.S.A. ABSTRACT The case is presented for the recognition of two genera of spurred lobelioids, Heterotoma and aa es, both presumably derived from the genus Lobelia. Hosmer Pe scien ao era were included in the genus Heterotoma Zucc. by most authors. Ayers (1990) c l, however, that fe spurred, blue- or pink-flowered lobelioids nee ee to be behnded within Lobelia section Hemipogon Benth. & Hook. A key and descriptions ae the two spurred genera derived from the genus Lobelia, 1.e. Heterotoma Zuc Calcaratolobelia Wilbur gen. nov., are provided together with the needed ern oe with the synonymy of the twelve species and one variety that form the genus Calcaratolobelia: C. aurita (Brandegee) W Wilbur, C. cordifolia (Hook. & Arn.) Wilbur, C. flexuosa (C. Presl) Wilbur, C. flexuosa var. intermedia (Hemsley) Wilbur, C. gzbbosa (S. Watson) Wilbur, C. goldmanii (Fern.) Wilbur, C. knoblochis (Ayers) Wilbur, C. macrocentron (Benth.) Wilbur, C. mevaughii (Ayers) Wilbur, C. margarita (F. Wimmer) Wilbur, C. pringle: (B.L. Robinson) Wilbur, C. tenella (Turcz.) Wilbur, and C. ene (Ayers) Wilbur. The synonymy of each taxa forming the genus Heterotoma is also provided > RESUMEN Se pretenden reconocer dos géneros lobeioides con flores espolonadas, Heterotoma y anes ambos pees derivados del o Lobelia. Anteriormente la mayoria de los autores inclufan ambos géneros en oe Zuee Ayers (1 990) cone sin embargo, que las lobeioides, de flores espolonadas azules 0 rosadas deberian incluirs Lobelia seccion edi ie Benth. & Hook. Se eneeen una clave ee ee 8 para Feet los dos géneros de flores espolonadas derivados del género Lobelia: Heterotoma Zucc. Calera Wilbur gen. nov., junto con las unertnaee necesarias con ot sinonimiuas }. IF pecies y una variedad que forman el gén C, aurita (Brandegee) = ee G. ash (Hook. & Arn.) Wilbur, é ee (C. Presl) Wilbur, flexuosa var. intermedia Hem ey) Wilbur, C. gébbosa (S. Watson) Lie C. goldmantii pe ' Wilbur, C. &noblochti ee nae C. macrocentron (Benth.) Wilbur, C. mcvaughit (Ayers) Wilbur, C. margarita (F. Wimmer) Wilbur, C. pringlei (B.L. — Wilbur, C. tenella (Turcz.) Wilbur, y C. villaregalis (Ayers) Wilbur. También se ofrece la sinonimia de los taxa que forman el género Heterotoma. My introduction to the lobelioid genus Heterotoma (Campanulaceae: Lobelioideae) was in the early spring of 1948. It was in the old National Herbarium of Mexico then located in Chapultepec Park in Mexico City. Sipa 17(3): 555-564. 1997 556 Sipa 17(@3) Rogers McVaugh, whose assistant I had the great good fortune to be on his first trip to Mexico, was paying a courtesy call on its then Director, Maximiliano Martinez. Dr. Martinez, whose English was said to be excel- lent, kept that fact from us. Dr. McVaugh spent a difficult afternoon of one day and the morning of another struggling to express himself in Spanish with very little help and no evidence of sympathy from Dr. Martinez. | soon escaped the linguistic ordeal by wandering off to browse in the her- barium. Afterwards I commented on the extreme heterogeneity of Heterotoma as clearly demonstrated by the collections. Dr. McVaugh indicated that the genus was not very natural, in that one species, H. /obelioides, seemed to be derived from the genus Lobelia in the vicinity of Lobelia laxiflora, and the small-, and blue-flowered species of Heterotoma more closely resembled the numerous blue-flowered Lobelias. The hypanchial spur was the principal feature binding together the disparate species composing Heterotoma. McVaugh commented that this was not an ideal example of a natural genus but perhaps it was the best that could be done with what was in all prob- ability merely “a genus of convenience.” This appraisal was later reaffirmed by him (1965), noting that taxonomy would not “be well served” by the addition to Hererotoma, of a very different spurred species of Lobelia from Colombia. Pragmatic treatments such as this may have served us well in mid-century and for a few decades thereafter, but their chances for contin- ued unchallenged acceptance in the age of cladism is slight. McVaugh’s (1940) study of the problems of generic recognition within the lobelioids led him to the following generalizations: “It appears, then, that almost any attempt at a natural classification of the Lobelioideae must strike a balance among the following courses: (1) Define a considerable number of small genera, each fairly uniform in char- acter, but set off from closely related groups by purely arbitrary characters. This course involves the creation of a number of new names. (2) Make generic limits ample enough to allow for the inclusion of most of the anoma- lous species. This will tend to make the genera larger and fewer, and will also necessitate the creation of many new names. (3) Make the genera as small and homogeneous as is compatible with logic, at the same time rec- ognizing the weight of convention as it bears on the subject of generic limits. It seems to me [i.e., McVaugh} that best results may be obtained from this last course, if at the same time it be remembered that the ‘genus’, as ordinarily defined, is a conventional concept; it is less a natural unit than the species and more to be thought of as a means of classification. Conve- nience, therefore must be taken into account as well as apparent kinship between species or groups.” Ayers (1990), having investigated the species comprised in the tradi- tional genus Heterotoma and its presumed closest relatives, concluded that Witpur, Calcaratolobelia, a new genus from Mexico and Central America 557 her admittedly “abbreviated study ... supports the hypothesis of parallel evolution of nectar spurs in at least two separate lineages within the Mexi- can and Central American members of Lobelia” and “makes the recognition of the polyphyletic genus Heterotoma untenable ... .” Some may recall the “good old days” when Simpson’s definition (1961, p.124) that monophyly “is the derivation of a taxon through one or more lineages ... from one immediately ancestral taxon of the same or lower rank,” was thought adequate. Under sucha concept of monophyly, it would seem that even Heferotoma in the traditional sense would be considered mono- phyletic, because the red and yellow, large-flowered Heterotoma lobelioides is suspected of having been derived from Lobelia section Homochilus A.DC., while the pink, white or bluish, small-flowered species may well have evolved from Lobelia section Hemipogon Benth. & Hook. f. In this event, the genus Heterotoma in its entirety would be thought to have evolved from the genus Lobelia, an ancestral taxon of the same rank. This of course is contrary to Hennigian principles since the “stem” Hererotoma is by inference a Lobelia. Although not convinced that Hererotoma in the sense of Mc Vaugh (1943) and Wimmer (1953) and other pre-Hennigian authors has been demon- strated to be unacceptably “polyphyletic” as claimed by Ayers, at least as the term was used in the first half of the century, it must be conceded that times have changed and that a stricter and narrower interpretation is now the prevailing practice in delimiting genera. If Heterotoma, in the restric- tive sense of the original publication and of Ayers’ (1990) conclusion, is deemed worthy of generic status, at least until further studies are com- pleted, it would seem that the spurred, small-flowered derivatives of Lobe- fia are equally deserving of generic recognition. Ayers concluded emphatically that the generic independence of Heterotoma s./, Gncluding both the red-yellow flowered species (H. /obeltoides) and the blue-flowered species) was unacceptable. Ayers stated that there were three acceptable alternatives to the scheme she favored. 1) Place all species of Heterotoma in the appropriate sections of the Lode- lia from which they apparently evolved i.e. place the blue or pink flowered species in Lobelia sect. Hemipogon and the red & yellow, long- spurred species in Lobelia section Homochilus. 2) Retain Heterotoma lobelioides as a separate monotypic genus and recog- nize the two groups of small-flowered species as members of the paraphyletic genus Lobelia. 3) Recognize a monotypic Heterotoma lobelioides with its two varieties and establish a new genus for the two small-flowered species groups, Macrocentron and Cordifolia. 4) Transfer Lobelia margarita to a newly erected genus containing the members of Hererotoma, excluding H. lobeltoides. on 558 Sipa 17(3) Ayers considered the fourth alternative acceptable only if evidence in addi- tion to the presence of floral spurs could be found relating Heterotoma pringlei and the very similar, pouched but non-spurred Lobelia margarita to the small- spurred taxa. It would seem to me that if Ayers’ investigation demonstrated anything convincingly it was that margarita and pring/lei are closely related. The establishment of a new genus for the small-spurred, blue-flowered species, although admittedly theoretically justifiable, was not favored by Ayers, fearing the proliferation of small, even though monophyletic, genera. Ayers favored the second of the listed options even though her solution continued recognition of a paraphyletic Lobe/za. Because she could not demonstrate that Lobelia laxiflora is a sister “group” to the small, blue-flowered herba- ceous species of Lobelia, Ayers concluded that it would be premature to trans- fer Heterotoma lobelivides into Lobelia section Homochilus since such a change might prove superfluous after the “final” analysis of Lobelia is completed. My sense of logical order within the spurred lobelioids leads me to favor the fourth alternative listed by Ayers: namely the recognition of Heterotoma as one genus probably derived from Lode/ia in the vicinity of section ene A.DC. and the establishment of a new genus for the bluish or pink-flowered spurred (or at least pouched) species apparently also derived from Lobelia but from section Hemipogon Benth. & Hook. f.. Cladists will condemn both Ayers’ arrangement and mine. The genus Lode/ia is admit- tedly rendered paraphyletic in Ayers treatment; cladistically my arrange- ment is even less tolerable since in all likelihood both Heteropogon and Calcaratolobelia are derived from different sections of Lobelia and hence Lo- belia is in all likelihood twice paraphyletic. (Perhaps one should leave well enough alone, but surely more trouble is in the offing for Lobelia and the stability of classification in the Lobeloideae. There would seem to be no possibility of maintaining Diastatea Scheidw. as a genus distinct from Lobe- fia if the same criteria are applied to it as have been to the small-spurred species here referred to Calcaratolobelia.) McVaugh (1945, p. 15-17) presented a series of eight recommendations to serve as a formal set of generic criteria for the Campanulinae, with which he was then primarily concerned, but which have proved helpful in other families. Two of his recommendations perhaps offer guidance (and sup- port) for the course of action taken here: “Recommendation 3 ... Genera of the nature of satellites, if also com- prising two or more series of species, invite suspicion especially if united by a single character which is also the only character separating them from a more inclusive genus.” This recommendation would argue against the recognition of Heterotoma s./, since it comprised two or more series of species, the red and yellow, large flowered H. /obelioides and the blue and white or pink, small-flowered Witsur, Calcaratolobelia, a new genus from Mexico and Central America 559 species as both series are united by a single character (i.e. the nectariferous spur) which 1s “also the only character separating them from a more inclu- sive genus” [1.e., Lobelia}. Recommendation 6. “Any segregate genus should be sharply delimited; that is, any species which 1s cvtermediate in one or more respects towards a more inclusive genus should be relegated to the latter. The retention of the anomalous species in the more inclusive genus will change its limits, if at all, but very slightly, and only in this way can the segregate genus be pre- cisely defined.” This recommendation would support Wimmer’s original placement of Heterotoma pringlei (= Lobelia gypsophila Ayers) in the genus Hererotoma rather than in Lobelia, where Ayers placed it, but to say this is not fair to Ayers for she placed all of the bluish or pinkish to white-flowered species in the genus Lobelia. It was her firm conviction that the spurred Herterotoma pringlei is not only a Lobef/za but that it is more closely related to the pouched but non-spurred Lobelia margarita than it is to any of the other bluish or pink- ish, small-flowered, spurred species as shown by its gypsophilous soil pref- erence, pedicellary movements during fruit maturation, similar and unique pollen exine, and similar, somewhat smaller, unusually colored seeds. Ac- knowledging the close relationship between Heterotoma pringlei (= Lobelia gypsophila) and Lobelia margarita as demonstrated by the shared features listed above, it would seem best to place the two species in one genus. Ayers concluded “that despite the difference in hypanthial morphology, they should not be placed in separate genera” and the genus to which both were assigned by her was Lobe/ia. In contrast I place both of these species (i.e. the species that Ayers refers to as Lobelia margarita FE. Wimmer and Lobelia gypsophila Ayers (= Heterotoma pringler B.L. Robinson) together with both groups of spurred “lobelioids” that Ayers designated as the “cordifolia species groups” and the “macrocentron species group’ into the same new genus here designated as Cal/caratolobelia. These three species groups can be separated by the following key. 1. Decumbent or procumbent perennial herbs spreading from underground rhizomes; pedicels erect and stationary during fruit maturation; corolla pinkish purple A) the Macrocentron species group. Erect annual herbs or tap-rooted perennials; capsule reoriented during fruit maturation by movement of the pedicel; corollas blue, pale lavender or white — or rarely pink-purple. 2. Tap-rooted perennials; pedicels divergent at 90° angles, in fruit deflexing abruptly at the hypanthium and reorienting the She: with spur or pouchside down and nearest the stem axis .......... B) tl 1e Gypsophila : species group. . Fibrous-rooted or tap-rooted annuals; ae arcuate at 45° angles fruit reflexing abruptly at the hypanthuim and reorienting the c ie with the spur-side nearest the stem axis... C) the Cordifolia species group. bo 560 Sipa 17(3) This is Ayers’ fourth alternative treament of the possible relationships of the components of Heterotoma s./, and their presumed ancestral groups within Lobelia. She did not favor this alternative until additional evidence besides the presence of floral spurs was found relating “Heterotoma pringlei (and thus L. margarita) to the other small-spurred taxa.” It is disconcerting that one is now placed in such a defensive position by recognizing segregate genera such as Hererotoma and Calcaratolobelia. Sach segregates Clearly render the very large genus Lobe/za paraphytetic and there- fore are unacceptable to a great many cladistically indoctrinated system- atists. Brummitt (1996), in a most interesting defense of the unavoidabil- ity of paraphyletic taxa, urges us “not to be ashamed of paraphyletic taxa” as they are and have been “an unavoidable and essential feature of our tax- onomy.” Clearly, if the orthodox cladist are correct, we systematists have barely begun our tasks for an alarming percentage of genera and families currently recognized are paraphyletic (Funk 1985; Judd et al. 1994). If one’s taxonomy admits that the evolutionary process has produced the in- credible diversity which we attempt to classify, then it seems inevitable that a great many of our taxa will be demonstrably ancestor-descendantly related. Such groups conflict with the cladists concept of monophyly since the recognized group does not include all of its descendants i.e. in this case the genus Lobelia does not include two of its probable descendants, the segregate genera Heterotoma and Calcaratolobelia. Perhaps brief comment is in order concerning the species described by McVaugh (1965) from the northern Andes of Colombia. McVaugh went to some pains to point out that, despite the gibbosity of the hypanthium, his new Columbian species clearly had nothing to do with the Mexican and Central American spurred /obeliads formerly placed in the genus Heterotoma; morphology suggested to McVaugh that the Andean species apparently was derived from Lobelia subg. Tupa. KEY TO THE NORTH AMERICAN SPURRED OR POUCHED GENERA OF THE LOBELIOIDEAE 1. Floral spurs crescent shaped, inflated laterally, che tube narrowing distally and with an abruptly dilated base; corolla red and yellow; calyx lobes attached to the distal end of spur, appressed to corolla tissue Heterotoma 1. Floral spurs conical or cylindrical, not inflated laterally, the base not dilated; corolla white, lavender-blue, or pink-purple in spurred species; calyx lobes variously atrached to the hypanthium, when discal at che sides of a cylindri cal spur, then oriented at a 90° or greater angle from the spur axis... Calcaratolobelia HETEROTOMA Zucc. Heterotoma Zucc., Flora 15(2) (Beibl.): 100. 1832. Tyre specirs: H. lobelioides Zucc. WitBur, Calcaratolobelia, a new genus from Mexico and Central America 561 Myopsia C. Presl, Prodr. Monogr. Lobel. 8. 1836. Type species: Myopsia mexicana C. Presl = H. lobeltoides Zucc.). Perennial, suffrutescent herbs to 1.5 m tall. Leaves petiolate, ovate, pubes- cent, with acuminate apices, the margin with yellow or purple, gland-tipped teeth. Flowers resupinate by twisting of the ebracteolate pedicel. Hy- panthium asymmetrical, the calyx and corolla elongated on the lower side into an arcuate, inflated spur with a bulbous base. Corolla unilabiate, the spur with all 5 lobes on the apparent abaxial side and pointing downward, the tube slit dorsally to base; tube and spur yellow-orange to burgundy; lobes subulate, yellow. Staminal column ca. 2 cm long, exserted vertically from dorsal slit in corolla; ventral filaments adnate to hypanthial rim and continuous with it to base of spur; ventral anthers with many, stiff, subu- late trichomes at apex. Corolla, stamens, and style persistent on the fruit. Fruit capsular, dehiscent by apical valves. Seeds numerous (60—100/cap- sule), ellipsoid, 0.55—0.6 mm long. 7=7. la. rusia a Onelioides Zucc. var. lobelioides, Flora 15(2)(Beibl.): 101. 1832. Type: MEXICO. Oaxaca(?): La Cumbre de San Antonio, 8000 ft, no date, Karwinski s.n. ie ee M; tsotypes: JE, M, ae mexicana C. Presl, Prodr. Monogr. Lobel. 8. 1836. Type: MEXICO, Oaxaca (?): a Cumbre de San Antonio, 8000 ft, no date, Karwinski s.n. (NEOTYPE: ?). a Bertoloni, Fl. Guatimal. in Novi. Comment. Acad. Sci. Inst. Bononiensis 4:409. 18 40. Type: GUATEMALA: Pinula, 4200 ft, J. Donnell Smith 1925 (NEOTYPE: NY, designated by Ayers 1990). eo ene Ortgies, in Regel, Gartenflora 12:50. 1863. Type: illustration in van te, Fl. Serres 14: pl. 1454. 1863 (Lectotype, designated by Ayers 1990). H. lobelioides var. tonelii (Ortgies) F. Wy amie Ann. Naas Mus. Wien. 56:371. 1948. Ib. Heterotoma lobelioides Zucc. var. glabra Ayers, Syst. Bot. 15:311. 990. Tyre: MEXICO. San Luis Porosi: in mountains near Santa Maria del Rio, Aug 1876, Schaffner 736 (HOLOTYPE: GH; Isotype: US) CALCARATOLOBELIA Calcaratolobelia gen. nov. Tyee: Calcaratolobelia cordifolia (Hook. & Arn.) Wilbur (=Lobelia cordifolia Hook. & Arn.) sea annuae vel perennes. Pedicelli graciles, saepe ebracteolati vel raro infimo cum bracteolis minutis et filiformibus. Hypanthia infirme ad valde asymmetrica, partim in ee corollae distenta, irregularia. Corolla tubulosa, in uno latere longitudinaliter fissa, basi in calcari plus minusve longum extensa vel solum gbbosa; limbus corollae oes ee quingquepartitus. Filamenta a corolla libera, sed duo longiora et in apicem calca decurrentia. Antherae 2 minores apice piloso-penicellatae. Stylus filiformis inclusus. oe sula semi-infera Annual or perennial herbs (1—)10—70(—-150) cm tall. Leaves petiolate or the cauline sometimes sessile. Inflorescences often secund and appearing pedunculate due to the naked upper part of the stem. Pedicels spreading- 562 Sipa 17(3) ascendent, slender, often ebracteolate or in two species with minute, fili- form bracteoles especially on the lowermost pedicels. Flowers inverted in anthesis by the twisting of the pedicel. Hypanthium weakly to strongly asymmetrical, the calyx and corolla extending on the abaxial side into a nectariferous knob or spur, the spur straight, cylindrical or tapering or at least not possessing a bulbous base. Corollas bilabiate with the two upper and three lower lobes, respectively, pointing in opposite directions, the tube slit dorsally to base or at least to within 1 mm of base. Staminal column less than 8 mm long and slightly exserted through the dorsal slit; the 2 shorter anthers with a stubble of numerous, short, apical bristles. n=7 or 14 when known. A small genus of twelve species known only from Mexico and Central America. Ayers’ (1990) key and descriptions adequately summarize the differences between the species and it seems unlikely that they can be improved upon until considerably more material has been collected and evaluated. The revised nomenclature is presented below with the species arranged in the same order as in Ayers’ publication except for the insertion of L. margarita as species #12 1. Calcaratolobelia macrocentron (Benth.) Wilbur, comb. nov. Hererotoma macrocentron Benth., Hooker's ae sas 2: 68, pl. 1177. 1876. Lobelia macrocentron oes 1.) Ayers, Brittonia 39:418 . TYPE: co. {DURANGO or Nayarit? }: ra Madre, NW of Mexico, eels 1849-50, Seemann 2049 (HOLOTYPE: K; ISOTYPE: BM). o 2. Calcaratolobelia villaregalis (Ayers) Wilbur, comb. nov. Lobelia villaregalis Ayers, Brittonia 39:419, 1987. Typi: MEXICO. Jazisco: cafion humido, Sierra de La Venta, 24 Nov. 1968, Villareal de Puga 2463 (HOLOTYPE: MICH; tsorype: IBUG). 3. Calcaratolobelia eigen (Ayers) Wilbur, comb. nov. Lobelia knoblochii yers, Brittonia 39:42 87. Type: MEXICO. Cuinuanua: shadec Moyarachic, 25 May ee Pie 5097 (HOLOTYPE: F; ISOTYPE: MSC), rf Ayers, Britconia 39:421. 1987. Type: MEX . DuRANGO: 34 road mi Salto, along ney from nanan to ee 2: noe L951, McVaugh 11528 ae ryPE: MICH; tsoryrees: MICH, NY, TEX). 4. Calcaratolobelia eon (Ayers) le comb. nov. Lobelia ae W © 5. Calcaratolobelia cordifolia (Hook. & Arn.) Wilbur, comb. nov. Lobelia cordifolia Hook. & Arn., Bot. Beechey Voy. 301. 1838. Heterotoma a (Hook. & Arn.) McVaugh, Bull. Torrey Bot. Club 67:143. 1940. Type: MEX . Nayarit: vicinity of Tepic, Jan-Feb 1827, Beechey s.n. ore E; ISOTYPE: ~~ Heterotoma salvadorensis F. W/immer, Ann. Naturhist. Mus. Wien 56:371. 1948. Type: EL SALVADOR. San Salvador, no date, Bernou!7 & Cario 8 (HOLOTYPE: GOET; IsOTYPES: GOET, W) Witsur, Calcaratolobelia, a new genus from Mexico and Central America 563 6. Calcaratolobelia Siar (Turcz.) Wilbur, comb. nov. Heterotoma tenella Turcz., jull. Moskovsk. Obsc. Isp. Prir., Otd. Biol. 25(3):175. 1852. Lobelia volcanica ie Syst. Bot. 15:317. 1990. Type: MEXICO. VERACRUZ: Xalapa, 600 m, Dec 184C ieee 7029 aut: K; tsoType: BR). Heterotoma cordifolia var. — es JE Wimmer, Pflanzenr. IV. 276b:717. 1953 Lobelia tenella L., Mantissa Pl. 10 177i. vue. Prodr. Fl. Cap. 40. 1794, non ae. Sicul. Pl. Cent. 1:53.t.2. 1806. 7. Calcaratolobelia gibbosa (S. Watson) Wilbur, comb. nov. Heterotoma gibbosa Watson, Proc. Amer. Acad. Arts 23:280. 1888. Type: ME . CHIHUAHUA: Oritz, no date, Pringle 1478 (HOLOTYPE: GH), non Lobelia gibbosa Labill., Nov. Holl. pl. 1:50. 1805 aig endlichii F. Wimmer, Feddes Repert. a Nov. Regni Veg. 26:1 pn Lobelia endlichii (F. Wimmer) Ayers, Syst .15:319. 1990. Typr: _ . eee bei Huejotitan in Tal des Rio a 1720 m, 5 Apr 1906, Endlich 1172 (Lectotype: W, designated by Ayers 1990). 8a. Calcaratolobelia flexuosa (C. Pres!) Wilbur var. flexuosa, comb. nov Rapuntinm flexuosum C. Presl, Prodr. Monogr. Lobel. 23.1836. Type: MEXICO. pre- sumably collected between Acapulco and Mexico City, no date, Haenke s.n. (HOLO- TYPE: =~ photo: NY, US). non Rapuntinm flexuosum C. Presl, Monogr. Lobel. 16.1836. Type: Cape of Good Hope. mee ntinm flexuosum C. Presl (p. 23) has priority over R. yeoae C. Presl (p. 16), Arc. 53. 6. ICBN. 1993.] Lobelia flexuosa (C. Presl) Cz dr. 7:378. 1839, ee flexuosa (C. Presl) McVaugh, Bull. Torrey Club 67:143. 1940. Lobelia arabidoides Hook. & Arn echey Voy. . pl. 66. 1838. Heterotoma avabidoides (Hook. & Arn.) oe ie Icon. Pl. 12:68. i Type: MEXICO. Nayarit: coos of Tepic, Jan—Feb 1827, Beechey 5.n. eee K). Heterotoma flexuosa var. liecbmanniana F. Wimmer, Pflanzenr. 1V. 276b:718. 1953. Type: MEXICO. [Oaxaca?}: “San Miguel de Pueto, Guatala,” no date, Liebmann 7801 (LEC- TOTYPE, designated by Ayers (1990), who selected from among three syntypes: C). 8b. Calcaratolobelia flexuosa (C. Presl) Wilbur var. pie orig ene OL ur, comb. nov. Heterotoma intermedia Hemsl., Biol. r. Amer. Bot 2:269. a eis len aad var. intermedia (Hemsley) FE. oa Pflanzenr. IV. 276b:7 1953. Type: MEXICO. [possibly collected between the cities of Durango eee Hesie.a 1850, Seemann 2051 (HOLOTYPE: K; IsoTyPE, BM). 9. i a etic (Fern.) Wilbur, comb.nov. Plies , Pre 36:504.1901. Lobelia goldmanii (Fern.) Ayers, Syst. ee 15: ih ae [as oe Type: MEXICO. SINALOA: road ae Las Flechas to La Rostra, 22 Feb 1899, Goldman en (HOLOTYPE: GH). The spelling employed by Fernald, H. goldmanii, was altered by Wimmer to H. eae - oe IV 276b:715- o 1953), and this spelling was sometimes adopted ers. The original collector's name was Edward A. Goldman but Wiramerm cited the collector's name as Goldmann. Asuicle 60 applies; the original spelling must be retained. } = 10. Calcaratolobelia aurita (Brandegee) Wilbur, comb.nov. Heterotoma aurita Brandegee, Proc. Calif. Acad. Sci. Il. 3:149. 1891. Lobelia anurita (Brandegee) Ayers, 564 Sipa 17(3) Syst. Bot. 15:324. 1990. Type: MEXICO. Baja CALIFORNIA DEL Sur: Sierra la La- guna, 21 Jun 1890, Brandegee 353 (LucroryPE: UC, ISOLECTOTYPE: G Lobelia cotensis M.E. Jones, Contr. W. Bot. 15:152. 1929. TYPE: MEXICO. Baja Cati- (ORNIA DEL SUR: Cota Ranch, Laguna Mts, 14 mi E of Todos Santos, 21 Feb 1928, 4147 (HOLOTYPE: POM, ee at US; isorypes: FE; MICH, MO, NY, POM). ms eae M.E. Jones, Contr. W. Bot. 18:68. 1933. Type: MEXICO. Baja CALI- FORNIA DEL SUR: The Laguna, Laguna Mes, 22 Sep 1930, Jones 27428 (HOLOTYPE: POM; IsoTyPes: BM, MO, NY, US). > = 11. Calcaratolobelia pringlei (B.L. Robinson) Wilbur, comb. nov. Heterotoma pringlei B.L. Robinson, Proc. Amer. Acad. Arts 44:615. 1909. Type: MEXICO. Nugvo LEON: daly mountains near Doctor Arroyo, 7 Nov 1904, Pringle 13274 (HOLOTYPE: GH; tsorypes: F, MIC ). 4 Lobelta gypsophila Ayers, Sida 13: .1988, based on Heterotoma pringle: B.L. Robinson, not Lobelia pringle: S. Watson, Amer. Acad, Arts. 25:157:. 1890. 12. Calcaratolobelia margarita (F. Wimmer) Wilbur, comb. nov. si margarita F. Wimmer, Ann. Naturhist. Mus. Wien. 36:355. 1948. Type: ME Nuevo Leon: Mpio. Galeana, Haciendo Pablillo, 14 Aug 1936, M. Taylor 167 aM LOTYPE: F; 1sorypr: TEX) Nothing in this paper should be taken as criticism of Dr. Ayer’s admi- rable contribution which is clearly based on her very careful study with its attention to field work especially in the mountains of Nuevo Leén. Quite obviously and admittedly my proposed solution is largely one that she listed as an “acceptable” alternative to the one that she prefers. REFERENCES Ayers, T.J. 1990. Systematics of Heterotoma (Campi — and the evolution of nectar spurs in the New World Lobelioideae. Syst. Bot. 15:296—-32 Brummitt, R.K. 1996. In defense of paraphyletic taxa ; eds. The biodiversity of African plants. Kluwer Aci sdemaic Publishers. FuNK, V.A. 1985. Cladistics and g n the Compositae. Taxon 34:72—80. 1985. sa W.S., R.W. SANDERS, and M. 1. Dende E. 1994. Angiosperm family pairs: prelimi- van der Maesen et al., = S = ON y phylogenetic analysis. Harvard Pap. Bot. 5:1—5 NeeVaiou, R. 1940. A revision of “Lawrentia” and al iiied genera in North America. Bull. Torrey Bot. Club 67:778-798. McVaucu, R. 1943. Campanulaceae, Lobelioideae. N. Amer. Fl. 32 A (pt 1):1—134 McVaucu, R. 1945. The genus Triodanis Rafinesque, and its een to SG heciiavik — 13-52. [Recommendations on generic criteria pp. 15 et McVauau, 1965. South famenican. Lobelioideae new to science. ae . Missouri Bo Gard. oe 409. [see page 404-406 for discussion of a spurred Lobelia that shou ; not be included i in Heterotoma or C Sa a Simpson, G.G. 1961. Principles of animal taxonomy. Columbia U. Pre WIMMER, EE. 1953. Campanulaceae - Lobelioideae. 2. Pflanzenreich rv 276b:261-814 {Heterotoma pp. 713—720.] STIDEROXYLON ALACHUENSE, A NEW NAME FOR BUMELIA ANOMALA (SAPOTACEAE) LORAN C. ANDERSON Department of Biological Science Florida State University Tallahassee, FL 32306-2043, U.S.A. ABSTRACT Bumelia anomala is transferred to the genus S/deroxylon, where the species must be given ecause the former specific epithet is preoccupied in the latter genus. Addi- tional (new) morphological data are added to the species description, and the species’ sa- lient features and rarity are evaluated. RESUMEN Bumelia anomala se transfiere al género Sideroxylon, donde la especie debe recibir un nombre nuevo porque el epiteto especifico esta ocupado en este género. Se afiaden nuevos datos morfolégicos a la descripcié6n de la especie, y se evaluan las caracterfsticas mds sobresalientes y raras. Key worpbs: Sapotaceae, Sideroxylon, Bumelia, silver buckthorn, Florida, rare plants, endemics. The silver buckthorn, Bumelia anomala (Sarg.) Clark, has a varied taxo- nomic history. It was originally described as a variety of B. /anuginosa (Michx.) Persoon (Sargent 1921) and was elevated to species rank by Clark (1942). Cronquist regarded it as “an uncommon but widely distributed form” of B. tenax (L.) Willd. (Cronquist 1945). Godfrey (1988) treated it as a species, whereas Wunderlin (pers. comm.) includes it within B. tenax with no infraspecific taxonomic status. Pennington (1990, 1991) has made a good case for transferring all Bumelia into Sideroxylon, but he did not mention B. anomala at any level. Our studies indicate that Bumelia anomala is a distinctive, rare endemic of central Florida, and that it merits recognition as a species. Unfortu- nately, the epithet anomala is preoccupied within Sideroxy/on, soa new name is proposed here: Sideroxylon alachuense L.C. Anderson, nom. nov. BasionyM: Bumelia lanuginosa var. anomala Sarg., J. Arnold Arbor. 2:168. 1921. Bumelia anomala (Sarg.) Clark, nn. Missouri Bot. Gard. 29:169. 1942. a ee forma anomala (Sarg. )Crong, : J. see Arbor. 26:456. 1945. Type: FLORIDA: [ALacHua Co.:} Gainesville, Jun 1917, T.G. Harbison 47 (A-2 sheets). one (1921) erroneously listed re ee from Hancock County (which does not exist in Florida), 2on Sideroxylon anomalum (Urban) T.D. Pennington, Fl. Neotropica Monogr. 52:123. 1990 Stipa 17(3): 565-567. 1997 566 Sipa 17(3) The species is named for its occurrence at Alachua Sink (type locality), part of Paynes Prairie State Preserve near Gainesville in Alachua County, Florida. Additional collections: Alachua Co.: Paynes Prairie State Preserve near Gainesville, 25 Jul 1918, 7 ne oo (A), in Harbison 64 (A oO Ju) 1919, T.G. Harbison 97 (A); 30 Nov 1934, L.E. Arnold & E. West s.n. (FLAS); 7 1980, W.S. Judd 2824 (FLAS); 25 Jul 1982, res anti 932 aoe < May nn re Godfrey 81292 (FSU), 22 Jun 1984, R.K. Godfrey 81389 (FSU); 12 Jun 1996 (three trees, Alachua Sink progeny on R.W. Simons’ Gainesville property), L.C. Anderson 16579 (FSU), L.C. Anderson 16580 (FSU), L.C. Anderson 16581 (FSU). Marion Co.: Silver River State Park, May 1985, R.W. Simons 2. (FSU). Orange Co.: near Orlando, 11 Nov 1917, 7.G. Harbison 51 (A). Small trees up to 9.2 m tall and 1.3 dm DBH; juveniles occasionally growing as low shrubs, spreading by horizontal rhizomes; long shoots (elon- gation or leader) bright green and with scattered pale hairs when young, often with sharp green thorns to 20 mm long, at nodes; some thorns be- coming short spur-shoots (cf. Godfrey 1988 for illustration). Mature shoots glabrous, gray-white, becoming thick and rigid. Leaves (3—)4—6(—9) cm long, (11-)16-30(-39) mm wide, petioles up to 5 mm long; blades on long shoots ovate-elliptic to rhombic or broadly obovate, tips obtuse to emarginate, those on spur-shoots smaller, usually oblanceolate to elliptic. All blades lustrous, dark green above with finely bony-cartilaginous re- ticulate veins, metallic-silver below with compacted dolabrate, sericeous hairs that hide the surface. Flowers S—6-merous, 10-20 in umbellate clus- ters on spur-shoots, tawny pubescent pedicels 4.5-5 mm long. Calyx 3— 4.5(-6) mm long, outer two sepals ore a orbicular, longer inner sepals narrower and often emarginate; all scruffy pubescent with mostly silver hairs plus small patches of light golden-brown hairs. Corollas white, 4.2-5(—6) mm long, petals 3-pronged, central prong longer and slightly spreading. Staminodes narrowly deltoid, obtuse, rarely minutely erose, nearly as long as and alternate the petals. Stamens opposite the petals, filaments somewhat dilated proximally, 1.7—1.9 mm long, anthers saggitate, 1.4— 1.5 mm long. Pistils velutinous, ovaries 1.5—1.8 mm, styles 1.4—1.5 mm long at anthesis. Single-seeded berries shiny black, oblong or rarely ovate, glabrescent with patches of hairs distally, 11-13 mm long and 10-11 mm wide excluding persistent 1—-1.5 mm style. Seeds light brown, smooth, 9— 10 mm long and 6 mm wide. This species has often been considered conspecific with S. tenax. How- ever, the two entities may be distinguished as follows: Twigs oe colored, glabrous or nearly so at maturity. Leaves (3—)4—6(—9) cm ong, (1 l-) 30(— 39) mm wide, dark green above, s hiny white below. Flowe ers 5—6-merous, in small clusters on spur-shoots, on pedicels 4.5—5 mm long, calyces 3—4.5(—6) mm long, anthers 1.4—1.5 mm long, styles 1-1.5 mm long. ae mid-late July. S. alachuense ANDERSON, Sideroxylon alachuense, a new name for Bumelia anomala 567 Twigs dark brown to oa ey or rusty, densely aia Leaves (2.5—) 4-5 m long, (6—) )mm wide, dull green above and brown- ish or rusty ie Se Z i merous, in larger sae on pedicels 4—9(— 12) mm long, calyces 2—2.8 mm long, anthers + | mm long, styles l.2 mm long. Flowering early— a -June in central Florida. S. tenax The silvery dolabrate hairs of S. a/achuense are strongly appressed and unidirectionally aligned, whereas the generally scruffy brown hairs of S. tenax are more interwoven, some twisted and ascending when observed through scanning electron microscopy. Distinctive stomatal patterns on the leaves provide additional characters diagnostic among Sideroxylon taxa (Anderson 1996). Stomatal density per unit of leaf surface is lower in S. alachuense than in S. tenax. The prominent peristomatal rims that nearly hide the guard cells are generally narrower and longer in S. a/achuense than in S. tenax Sideroxylon alachuense (as Bumelia anomala) is listed as endangered in Florida (Coile 1993). Known occurrences are few; perhaps only 20 plants exist in the wild. About 20 plants are in cultivation in Gainesville through the efforts of R. W. Simons. It is the rarest and perhaps the most beautiful buckthorn in the United States. ACKNOWLEDGMENTS Bob Simons is responsible for most of our distributional knowledge about the silver buckthorn. He is thanked for allowing me to collect from his plants, taking me into the field to observe buckthorns, and providing me with flowers and fruits (which are quite tasty). Ted M. Barkley made help- ful suggestions on the manuscript. REFERENCES ANDERSON, L.C. 1996. New es hical and morphological data for Sideroxylon thorner (Sapotaceae). Sida 17:343— Ciark, R.B. 1942. A revision of ‘ae genus Bumelia in the United States. Ann. Missouri Bot. Gard. 29:155-18 Coie, N.C. 1993. aa s endangered and threatened plants. Fla. Dept. Agriculture and Consumer Serv., Division of Plant Industry-Botany Section Contr. 29, Gainesville Cronauist, A. 1945. Studies in the Sapotaceae, III. Dipholis and Bumelia. J. Arnold Arbor. 26:435—- 471. Goprrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. Univ. Georgia Press, Athe PENNINGTON, T.D. 1990. Sapotaceae. FI. Neila Monogr. 522] cea _«d'91. The genera of Sapotaceae. Royal Botanic Gardens SARGENT, cs. 1921. Notes on North American trees. VII. J. Arnold rea ir 264-174. 568 Sipa 17() BOOK NOTICES Ho ianp, V.L. and Davip J. Krit. 1995. California Vegetation. (ISBN 0- 7872-0733-0, pbk). Kendall/Hunt Publishing Company, 4050 Westmark Drive, Dubuque, IA 52004-1840. $42.95. 516 pp. This book, originally done as a text but now revised for wider use, tells all that an average visitor or resident might want to know about California vegetation. The first 120 pages cover topography, geology, and soils; climate; vegetational history; community classification; and plant names. Most of the rest of the book is devoted to Lucid discussions of California’s various types of vegetation from marine habitats to alpine areas, including general descriptions, basic environmental factors (e.g., climate, soils, and human impact), and lists of important plants. Concluding each type’s section is a list—often extensive—of references for further reading, a most welcome addition. The index is detailed. Liberal use is made of black-and-white eaten ice of them helpful, a few disappointing. Use- ful, too, are the maps showing location the various vegetation ty pes and the ranges of selected California plant species. When you, as a visitor, go to California—or, as a resident, take this book with you. You'll be glad you did.—John W. Thieret. explore your state Seymour, RANDY. 1997. Wildflowers of Mammoth Cave National Park. 1997. ISBN 0-8131-0898-5, pbk). The University Press of Kentucky, 663 South Limestone Street, Lexington, KY 40508-4008. $17.95 + $3.50 shipping. 254 pp. This book includes color . hotographs (mostly excellent), earmarks, habitat data, flowering times, and uses/folklore of 400 species found in Mammoth Cave National Park’s 53,000 diverse acres. The flowers are arranged, first, by flowering dates and, second, by color. The book tells the user where in the park—t.e., in what areas or along what trails—to find the various flowers, a welcome bonus. This work, a labor of love for its author, is beautifully done. Even if you do not visit the park, W/dflowers is a pleasure to look at. If you do visit, you will find the guide useful. | recommend tt.—John W. Thieret KorMONDY, Epbwarb J. 1996. Concepts of Ecology. Fourth Edition. ISBN Q-13-478116-3, pbk). Prentice-Hall, Inc., Simon & Schuster/A Viacom Company, Upper Saddle River, NJ 074 458, U.S.A. Price: not ziven, 322 pp. From the cover: “In this new edition, Edward J. Kormondy continues to focus on eco- system ecology with primary emphasis on energy flow, biogeochemical cycling, popula- tion ecology, and community ecology. A final section on human ecology integrates these four major areas. Designed for sophomore and junior students with minimal background in biology and chemistry, terminology is Sica and elementary mathematics is used for quantitative aspects. Stipa 17Q): 568. 1997 CRATAEGUS NANANIXONI (ROSACEAE, SER. INTRICATAE) A NEW SPECIES OF HAWTHORN FROM EASTERN TEXAS J. PEEPS The University of Western Ontario Department of Plant Sctences London, Ontario, CANADA NOA 5B7 R.J. OKENNON Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-40060, U.S.A. ABSTRACT A new species of Crataegus, C. nananixonii, is described from Texas and illustrated. The new taxon is a small plant endemic to sand hills near Nacogdoches. RESUMEN Se describe y se ilustra una nueva especie de Crataegus, C. nananixonii, procedente de . El nuevo taxon es una pequefa planta endémica de las colinas arenosas cerca de Nacogdoches. In preparation for the forthcoming Flora of Texas, this paper describes a new species of Crataegus which has been known for a number of years from sand plains near Nacogdoches, Texas. This species cannot be matched with anything in lists of Texas plants (e.g. Hatch et al. 1990, Johnston 1990, Mahler 1988, Nixon 1985, Simpson 1988, Vines 1977) or Louisiana (MacRoberts 1988). In Correll and Johnston’s (1970) Manual of the Vascu- lar Plants of Texas it would key down unambiguously to C. iflora Muenchh., if che ripe fruit color were known. However, it is very different from that species, as discussed below. Crataegus nananixonu J.B. Phipps & R.J. O’Kennon, sp. nov. ae 1). Type: TEXAS: NacoGpocues Co.: | mi N of jet. of FM1087 and Co. Rd. 153, on Co. Rd. 153, 14 Apr 1989, E.S. Nixon 17430 (HoLoryPE: ASTC; 1soTyPE: UWO). Frutex, 1—2 m altus, intricate ramosus; a 1-3 cm longae, tenues, rectae vel plus usve recurvatae, sordide atrocinereae. Folia decidua, petiolata; laminae 1.5—3.5 cm longae, rhombo-ovatae, vadosissime lobatae vel non-lobatae, mes sinious nonnihil irregulariter seratis, in juventice sparsim scabro- caus scentes adaxiale, glabrae abaxial Daxlale, in maturitate g pubescentia adaxiale in venas, glanduloso-maculatae in allis dentes, praecipue proxime, cae ca. 3 venarum secondariarum, petioli 5-15 mm Stipa 17(3): 369-574. 1997 570 SIDA 17(3) $8 Fic. 1. Crataegus nananixonit J.B. Phipps & R.J. O’Kennon. Fruiting specimen, fruit and parts of whole leaf from J.B. Phipps 6277 (UWO). Flowering specimen and flower parts from E.S. Nixon 17340 (UWO). S. Laurie-Bourque de/. Puipps AND O’KENNON, A new species of Crataegus from Eastern Texas 71 ‘ longi, glanduloso-maculati. Inflorescentiae 3—5 florae; pedicellis patento-pubescentibus, r bracteolis caducis linearis glandulo-marginibus conspicuis in anthesem praecocem; anthe- sis in Aprili. Flores 12-15 mm diam.; hypanthium glabrum; lobi calycis 4-5 mm longi, glandulo-marginati, glabri adaxiale; petala + orbiculares, plus minusve unguiculata; stamina ca. 10, antheris rose-purpureis; carpelli et styla 5. Infructescentiae 1—4 fructae, pedicellis tenuiter patento-pubescentibus. Fructus ca. 1 cm diam., orbiculares, cupreo-rubri, in maturitate glabri, reliquiis calycis adsentibus vel non in ore leviter elevato; pyrena 5. 5 Shrub, 1—2.5 m tall, quite intricately branched; thorns 1—3 cm long, fine, straight to slightly recurved, dark dull grey. Leaves deciduous, peti- olate; leaf-blades 1.5—3.5 cm long, rhomb-ovate, extremely shallowly lobed or unlobed, margins somewhat irregularly serrate, thinly scabrous-hairy adaxially when young, glabrous abaxially, at maturity almost entirely gla- brous except for some pubescence on the veins abaxially, and with gland- dots in some of the teeth especially proximally, with about 3 pairs of sec- ondary veins; petioles 5-15 mm long, gland-dotted. Inflorescence 3—5 flowered, branches with spreading pubescence, caducous linear gland-mar- gined bracteoles conspicuous in early anthesis; anthesis April. Flowers 12— 15 mm diam; hypanthium glabrous; calyx lobes 4-5 mm long, gland-mar- gined, glabrous abaxially; petals + orbicular with slight claw; stamens about 10, anthers rose-purple in color; carpels and styles 3—5. Infructescences 1— 4 fruiced; pedicels with thin spreading pubescence. Fruits 1 cm diameter, + orbicular, coppery red when ripe, glabrous, calyx remnants present or not ona slightly elevated rim; pyrenes 3—5. Additional specimens examined: TEXAS: Nacogdoches Co.: ca | mi N of jet. of FM 1087 & Co. Rd. 153 on 153, occasional, regenerating on a clearcut area, 1.9 m tall, white flowers, anthers rose- elas 14 Apr 1989, E.S. Nixon 17334 (UWO); ca. 1 mi N of jet. of FM 1087 & Co. Rd. 1 153, regenerating on a clearcut area, eee 1.8 m tall, white flowers, anthers rose- ae 14 Apr 1989, E.S. Nixon 17336 (UWO); ca. 1 mi ie of jct. of FM 1087 & Co. Rd. 153 on 153, og aaa on a clearcut area, occasional, tall, white flowers, anthers rose-purple, 14 Apr 1989, E.S. Nixon 17339 (U Gas : i N of jet. of FM 1087 & Co. Rd. 153 on 153, regenerating on a clearcut area, occasional, 2 m call, white eae eens rose- pi 14 Apr 1989, E.S. Nixon 17340 (UWO); ca. 1 mi N of jct. of F & Co. Rd. n 153, regenerating ona clearcut area, occasional, 1 m tall, a howe een rose- mle 14 Apr 1989, E.S. Nixon 17342 QO); ca. 1 mi N of jet. of FM 1087 & Co. Rd. 153 on 153, regenerating on a clearcut area, occasional, 1.5 m tall, white et anchersr rose- purl, 14 Apr 1989, E.S. Nixon 17343 (UWO); ca. 1 mi N of jet. of FM 1087 & Co. Rd. 153 on 153, regenerating on a clearcut area, occa- sional, 1.2 m tall, white Done anthers rose-purple, = Apr 1989, E.S. Nixon 17344 (UWO),; 1.3 mi W of jet. of FM 1087 & Co. Rd. 153 on 1087, dry upland, post oak, shortleaf pine & sandjack -— 1.8 m tall, white flowers anthers rose-purple, 19 Apr 1989, E.S. Nixon 17321 (UWO); 1.3 mi W of jcc. of FM 1087 & Co. Rd. 153 on 1087, dry upland, post oak, shortleaf pine & sandjack oe 2.1 m tall, white flowers, anthers rose- Hee 19 Apr 1989, E.S. Nixon 17322 (UWO); 1.3 mi W of jct. of FM 1087 & Co. Rd. on 1087, dry ieee post oak, shortleaf pine & sandjack oak, 2.1 m tall, white flow- ers, cae: rose-purple, 19 Apr 1989, E.S. Nixon 17323 (UWO),; 1.3 mi W of jct. of FM )87 & Co. Rd. 153 on “1087, dry upland, post oak, shortleaf pine & sandjack oak, 2.4 m 572 Sipa 17(3) tall, white flowers, anthers rose-purple, 19 Apr 1989, E.S. Nexon 17324 (UWO); 1.3 mi W ena of FM 1087 & Co. Rd. 153 on 1087, dry upland, post oak, shortleaf pine & sand- fee ee a m tall, white flowers, anthers rose- ie 19 Apr 1989, E.S. Nixon 17325 (U 3 mi W of j ~FM 1087 & Co. Rd. 153 on 1087, dry upland, we ci short- ne pine & sete oi 71 m tall, white flowers, nee rose-purple, 19 Apr 1989, E.S. Nixon 17326 (UWO); 1.3 mi W of jet. of FM 1087 & Co. Rd. 153 on 1087, i: up post oak, shortleaf pine & sence oak, 2.4 m tall, white flowers, anthers rose-purple, | Apr 1989, E.S. Nixon 17327 (UWO),; 1.3 mi W of jet. of FM 1087 & Co. Rd. 153 on > and, Cc 1087, dry upland, post oak, shortleaf pine & sandjack oak, 2.3 uite flowers, anthers rose-purple, 19 en 1989, E.S. Nexon 17328 (UWO); tight-1 yand side (northwards) of Co. d. 153, several miles W of Garrison on FM 1087, just N of scout camp, 31°52'N 94°37'W, 500! a.s.1., extreme sandy soil, sparse scrub of dwarf oak, 1 m bush with clusters of ruddy fruit, 18 Oct 1988, J.B. Phipps 6277 (UWO); Hwy 1087, 3.7 mi E of Hwy 250, N of Nacogdoches, sandy forest, 13 Apr 1993, R.J. O'Kennon 11239 (BRIT, UWO). This distinctive and attractive species is restricted to open sandy scru- bland of oak, in Nacogdoches Co., Texas, where it is locally common and we are naming it for Elray S. Nixon, formerly of Stephen EF. Austin State University, Nacogdoches, who provided J.B. Phipps with a quantity of flower- ing material and directed him to its site in September 1988. The first part of the root “nana” recognizes the dwarf stature of the plant. At first sight, Crataegus nananixonii resembles C. uniflora Muenchh. of series Parvifoliae Loud.) Rehder, but this is primarily due to its small habic, leaves, flowers, and fruits and few-flowered inflorescences. Detailed charac- teristics however, are more similar to those of series [#tricatae (Loud.) Rehder with which C, zananixoni shares a thorny nature, rhomb-elliptic leaf shape, glandular leaf-bases and petioles, abundant caducous glandular bracteoles and ruddy fruit with a small collar. Also, the unspecialised calyx lobe of C. nananixoni excludes this plant from series Parvifoliae. There is, however, a form of Crataegus uniflora superficially similar to C. nananixonii with rhom- bic-obtrullate rather than elliptic-obtuse leaves more typical of the species that is widely scattered across the southern states. This has been recorded for Texas on sandhills near Tyler in Smith Co. and also has several flowers to the inflorescence, like C. nananixonii, a feature however, not rare in true C, uniflora. The possibility that Crataegus nananixonii was in reality a highly dwarfed form of a generally larger species of series Itricatae therefore needed to be addressed in some detail and for this purpose we considered those species listed in Vines (1960), i.e. all members of the series occurring west of the Mississippi. Crataegus intricata, C. neobushii, C. buckley1, C. rubella, C. padifolia and C. pagensis may be rejected immediately on account of their glabrous inflorescences and young foliage since this is a generally reliable character at the species level in North American Crataegus taxonomy. Among the species with pubescent inflorescence and foliage all are much too large but Puipps AND O’KENNON, A new species of Crataegus from Eastern Texas 13 need to be compared, nevertheless. Of those with some substantive simi- larity to C. nananixonii, C. harveyana and C. ouachitensis have deep-pink, rose purple or rose-colored (‘red’ in Vines’ key) anthers but 20 stamens while C. bi/tmoreana has 10 stamens but cream-colored anthers. Stamen number and anther color are, of course, two generally good taxonomic char- acters at the species level in Crataegus and have been considered so by Sargent, Palmer, Kruschke and J.B.Phipps, among others. Indeed, Kruschke (1965) took an absolutist stance on the stamen number issue declaring the impos- sibility of 10 and 20-stamen entities belonging to the same species. In examining section Dowglasii (black-fruited hawthorns) Dickinson et al. (1996) have shown that 20-stamen forms are self-incompatible, often dip- loid (and presumably always sexual) while 10-stamen forms are self-com- patible, often not diploid (and may be apomictic), thus providing an expla- nation for Kruschke’s viewpoint. My own (JBP) opinion, however, is that the distinction is not necessarily that rigid and that this could be in part due to the stage of evolutionary divergence reached at a particular point in time. The point to note, however, is that stamen number and anther colour are strong characters and that in the case of C. nananixoni they support its specific status as a rare local edaphic Texan (for the time being) endemic rather than as a dwarfed form of some other species in series Intricatae. ACKNOWLEDGMENTS We wish to thank the National Sciences and Engineering Research Coun- cil of Canada for funding this work, Susan Laurie-Bourque of Hull, Quebec for preparing the illustration and Antony Littewood of the Department of Classical Studies, The University of Western Ontario for checking the Latin diagnosis. REFERENCES Corre, D.V. and M.C. a de Manual of the vascular plants of Texas. Texas esearch Foundetion, Renner DICKINSON, T.A., S. BELAOUSSOFF, R. M. Love, and M. MuniyamMa. 1996. North American black-fruited eee Cpineus sect. ” Douglas Loud.): 1. Variation of floral con struction, eas — and their possible evolutionary significance. Folia Cab Phytotax. 31:3 Hatcu, S.L., KN. on and L.E. Brown. 1990. Checklist of the vascular plants of Xds. JOHNSTON, M.C. 1990. The vascular plants of Texas, 2nd ed. published by Author, Austin, exXas. Kruscuke, E.P. ee Contributions to the taxonomy of Crataegus. Milwaukee Public. us. Public. Bot. MacRoberts, D.T. 1988. A documented checklist and atlas of the vascular flora of Louisiana. Louisiana State Univ., Shreveport. Sipa 17(3) WN | Maner, Wm. F 1988. Shinners’ manual of the north central Texas flora. Sida, Bot. Misc. 3. Nixon, E.S. 1985. a shrubs and woody vines of east Texas. Bruce Lyndon Cunningham, Nacogdoches, Simpson, B.J. | es t eld guide to Texas trees. Texas Monthly Press, Austin, TX. Vines, R.A. 1960. Crataegus In: Trees, aneubs and woody vines of the southwest. Univer- sity of Texas Press, Auscin, TX. Pp. 329-387 Vines, R.A. 1977. Trees of east Texas. Univ. of Texas Press, Austin. TX. PALYNOLOGICAL EVIDENCE SUPPORTING THE IDENTITY OF TWO TAXA OF BERBERIS (BERBERIDACEAE) FROM TIBET ARTI GARG, TARIQ HUSAIN, and R.R. RAO Taxonomy and Biodiversity Division National Botanical Research Institute Rana Pratap Marg, Lucknow - 226001 U.P, INDIA ABSTRACT The identity of Berberis replicata var. dispar and B. griffithiana var. pallida is supported on the basis of pollen morphology. Phylogenetically B. rep/icata was found to be a primitive, and B. griffithiana an advanced taxa. RESUMEN La identidad de Berberis replicata var. dispar y B. griffithiana var. pallida se mantiene en base a la morfologia polinica. Filogenéticamente B. replicata es un taxon primitivo, y B. griffithiana uno derivado. INTRODUCTION In continuity of Husain et al. (1994) paper on The identity of two taxe of Berberis ( Berberidaceae) from Tibet, the identity of Berberis replicata W.W/. Sm. var. dispar Ahrendt and B. griffithiana Schneid. var. pallida (Hk.f. & Th.) Chamberlain & Hu—is discussed in the light of pollen morphological char- acters using light and scanning electron microscope. MATERIAL AND METHODS Polliniferous material for the present study was obtained in the form of dried flower buds from Herbarium sheets. The material was soaked in 70% alcohol for 48 hours and pollen was subsequently acetolysed according to the method of Erdtman (1952). One part of the acetolysed pollen was mounted on slides in glycerine jelly and observed under compound micro- scope while the other part was mounted on brass stub, gold coated, ob- served and photographed under scanning electron microscope. RESULTS AND DISCUSSION The morphological characters of the three taxa (distributed in Eastern Himalaya and China) namely B. replicata var. dispar (India: Arunachal Sipa 17(3): 575-578. 1997 576 Stipa 17(3) Pradesh: T. S. Rana & Party 210335, LWG), B. griffithiana var. griffithiana (India: Arunachal Pradesh: T) S. Rana & Party 210330, LWG) and B griffithiana var. pallida (China: Ludlow, Sheriff & Taylor 3697, BM) are tabu- lated and a key for the seggregation of B. replicata and B. griffithiana are provided in Husain et al. (/. c.) communication. The pollen morphology of these taxa is given in Table | and explained as follows: Berberis replicata var. dispar (Figs. 1, 2) Grains inaperturate, 2—3 colpoid streaks occur in association with ir- regular fissures on the exine surface. Exine 1.7 jim thick; ectoexine equal to endoexine, columella indistinct; surface psilate, ill-defined with irregular folds. Grains spheroidal; diam 45 ppm (44—47 jam); tectum ill-defined. Berberis griffithiana (Figs. 3—G) Grains spiraperturate, spiral twines irregulary dileneating the surface into elongated and rounded areas; aperture narrow, margin smooth. Exine 3 pm thick, ectoexine thicker than the endoexine, columella indistinct; surface coarse, organised into irregular raised lumps of piloid elements or warted. Grains spheroidal, diam 34/40 pm (32—42 pm). Tectum irregular. KEY TO VARIETIES 1. Grains spheroidal; diam 34 jm; surface with closely packed, homogenous piloid elements var. griffithiana 7 x 40 pm; surface warted var. pallida 1. Grains sub-spheroidal; 3 The pollen grains of B. griffithiana var. griffithiana and B. griffithiana vat. pallida were found to be similar to each other but revealed considerable differences with those of B. replicata var. dispar in primary (apertural), sec- ondary (exine ornamentation) and tertiary (exine strata, shape and size) characters (see Table 1). This supports the view of Husain et al. (/ ¢.) that B. replicata var. dispar is a distinct taxonomic entity and being closer to B. replicata in macro-morphological characters, should be kept as a variety under B, replicata and not be merged under B. griffithiana var. pallida. Walker and Doyle (1975) phylogenetically classified pollen with respect to the number of apertures as (1) Inaperturate (ii) mono-aperturate (iii) di- aperturate (iv) tri-aperturate or (v) poly-aperturate with 4,5,6 apertures and aperture shape as (i) elongate, furrow like (ii) round, pore like (iii) encircling ring or band like. The pollen grains of B. replicata var. dispar are basically inaperturate, the primitive type whereas those of B. griffithiana are spiraperturate (encircling the grain), the advanced type. Hence, replicata is considered primitive and B. griffithiana advanced. The two taxa represent the two extreme ends of the evolutionary status and thus strongly support the distinct identity of these two taxa. GarG ET AL., Palynological evidence for two taxa of Berberis a7 7 a a Fics. 1-6 Scanning electron micrographs of pollen. Fics. 1-2. Berberis replicata var. dispar, 1) surface with cracks and fissures. 2) surface en- ere showing irregular folds. Fics. 3-6. Berberis griffithiana, 3) pollen with semi-spiral. 5) surface with piloid elements. his var. pallida, 4) pollen with one free colpus. 6) warted surface. 578 Sipa 17(3) ACKNOWLEDGMENTS The authors are thankful to the director, National Botanical Research Institute, for use of facilities. REFERENCES Erptman, G. 1952. Pollen morphology and plant taxonomy. Angiosperms. Almqvist and Wiksell, Stockholm Husain, T., B. Darr, and R.R. Rao. 1994. On the identity of two taxa of Berberis (Berberidaceae) from Tibet. Sida 16:17—2 Wacker, J.W. and J.A. Doyie. 1975. The basis of angiosperm phylogeny: Palynology. Ann. Missouri Bot. Gard. 62:664—7 23 NOMENCLATURAL NOTES AND A SYNOPSIS OF THE GENUS MYRSINE (MYRSINACEAE) IN MESOAMERICA JON M. RICKETSON Missouri Botanical Garden . Box 299 St. Louts, MO 63166-0299, U.S.A. ricketso@ mobot.ore JOHN J. PIPOLY II Botanical Research Institute of Texas O09 Pecan Street Fort Worth, TX 76102-4060, U.S.A. ppipoly @ brit. org ABSTRACT A nomenclatural summary is provided for the species of the genus Myrsine occurring in Mesoamerica. An updated description of the genus and its taxonomic synonyms is pro- vided, along with a key to the species. Seven taxa are recognized, including the new com- r.ex Roem. et Schult. subsp. cane (Lundell) Ricketson bination Myrsine coriacea (Sw.) R. B ¢ synonyms provided, and & Pipoly. All taxa are enumerated, nomenclatural and taxonom two species, Myrsine juer. ties and M. pellucidopunctata, are ie oeeiaed. Sixteen bino- ys: mials are newly relegated to s RESUMEN Se ofrece un resumen de las especies mesoamericanas pertenecientes al género Myrsme. Se incluye una descripcidn actualizada del género y sus sindnimos taxonémicos, junto con . Se reconocen siete taxa, incluyendo la nueva Br. ex Roem. et Schult. subsp. #/grescens (Lundell) anto nomenclaturales una clave para identificar las especies Seba Myrsine coriacea (Sw.) R. Ricketson & Pipoly. Se enumeran los taxa, se ofrecen sinénimos tan ze) nee He y se lectotipifican dos especies, Myrsine guerguensenii and M. pellucidopunctata. Se relegan dieciséis bindmenes como nuevos sindnimos taxonomicos. INTRODUCTION The genus Myrszve R. Br. contains ca. 300 species of which nearly 1/4 remain undescribed. C. Chen and Pipoly (1996), Pipoly (1991, 1992a, 1992b, 1996) and Pipoly and C. Chen (1995) have provided summaries of evidence for broader circumscription of the genus, particularly to include Rapanea Aubl. The genus is pantropically distributed, occurring in diverse vegetation types, from mangroves to subalpine scrub, but always in moist, wet or pluvial habitats. The genus is defined by lateral (axillary), fascicu- Sipa 17(3): 579-589. 1997 580 Sipa 17(3) late or umbellate inflorescences, sessile or on short, perennating peduncles girdled by persistent floral bracts, thus forming “short shoots.” In prepara- tion for our treatment of the genus for Flora Mesoamericana, it became nec- essary to assemble complete synonymies and bibliographic references, and lectotypify several species. Because of the somewhat abbreviated format of that Flora, the present synopsis is intended to provide a complete nomen- clator for this often misunderstood and nomenclaturally complex genus. TAXONOMIC TREATMENT Myrsine L., Sp. Pl. 1:196. 1753, Gen. Pl. ed 5:90. 1754; Roem. & Schult., Syst. Veg. 503. 1819; A. DC. , Trans. Linn. Soc., London, Bot. 17:104. 1834, Ann. Sci. Nat. Bot. 9:292. 1834, Ann. Sci. Nat. Bot. 16:65—97, 129-196, tab. 1-3, 8-9, 1841; A. DC in DC, Prodr. 8:92. 1844. Mig. in Mart., Fl. Bras. 10:306. 1856. Hook. f. in Benth. & Hook., Gen. Pl. 2:642. 1876. Mez in Engl., Pflanzenr. 9(V. 236):338. 1902. E. Walker, Philipp. J. Sci. 73:1940, Bot. Mag. Tokyo 67:249. 1954, Bull. Ag. Home Ec. Univ. Ryuku 2:76. 1955, Quart. J. Taiwan Mus. 12:164. 1959. Stearn, Bull. Brit. Mus. (Nat. Hist.), Bot. 4:174. 1969. Fosberg & Sachet, Smithso- nian Contr. Bot. 21:3-11. 1975. Lundell, Phytologia 48:137. 1981 Se se S6:418. a ee Novon . =e 1991, ier 17:1. 1992, Novon 2:392. Pipoly & C. Chen, on 5:3 1995; Harvey & Pipoly, FI. Pics das ve mas 995:C. ce & ‘Pipoly, = China | ete 996; Pipoly, Sida 17:115-162. a 487.1 996. Type species (by monotypy): Myrsine africana I Rapanea Aubl., Hist. Pl Guiane aie t. 46. 1775. 245. 1783. Duhamelia Dombey ex Lam., Encycl. Samara Sw., Prodr. 1:120. on pro non LL. Manglilla A. Juss., Gen. Pl. _ 1789. Arthrophyllum Lour., Fl. Cochi 7 ae Shae Caballeria Ruiz & Pa , FL. Peruy. Prodr. 1:141. Roemeria Thunb. (non Alin Nov. Gen. Pl. . 130. aa Scleroxylum he : Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten aturk. 3:57. 1809 Pi A. — i FI, Nouv.-Zel. 349. pl. 38 Mertsta Banks & Sol. ex A. Cunn. in A. oe in o an 8:95. 1844. (axillary), um- — Shrubs or small trees. Leaves alternate. Inflorescences latera bellate or fasciculate, sessile or on short accrescent branchlets girdled by persistent floral bracts. F/owers 4—5(—6)-merous, bisexual or unisexual, the plants then bisexual, monoecious, dioecious, or polygamous; sepals nearly free or united to 1/2 their length, imbricate or valvate, usually ciliate, punc- tate, persistent; petals nearly free or rarely united to 1/2 their length, usu- ally ciliate, glandular-granulose at least along margin and often through- out within, punctate; stamens and staminodes similar, subequal to corolla length, the filaments free or connate basally to form a tube, the tube with or without sterile appendages alternating with the filaments, and all merely adnate to the corolla tube; or developmentally fused throughout, the RICKETSON AND PipoLy, Myrsine in Mesoamerica 581 anthers thus appearing epipetalous, the anthers and anctherodes similar ovate or reniform, elliptic or oblong, rarely sagittate, 2-celled, dehiscing by longitudinal slits, or rarely by subterminal pores opening later into wide longitudinal slits; pistil and pistillode similar; conic, obturbinate obnapiform, or variously subglobose; ovary globose, costate or not, gla- brous or glabrescent; ovules few, uniseriate, completely immersed in pla- centa or seated below apical pores in placenta or variously projecting; styl obsolete to present, tapering into stigma; stigma morchelliform, liguliform sinuate to lobate, prismatic and 3-lobed, or rarely conical. Fruit a globose, subglobose, ellipsoid, ovoid, or subovoid drupe, with somewhat fleshy exo- carp and crusty or leathery endocarp, 1-seeded; endosperm horny, rumi- nate; embryo cylindric, transverse. In Mesoamerica, six species are known, that may be distinguished by the following key. KEY TO MYRSINE OF MESOAMERICA . Branchlets, petioles and/or midrib of leaf blade ferrugineous to rufous to- Se . = early glabrescent, the trichomes uniseriate 2, i af bla ra e apeewiate es a ar or rarely widely oblong, 0. 6- We 3.5cm ae the gaterminal mucron; petioles up to - 4cm long 2. Leaf blades elliptic, haan oblanceolate or lanceolate, 2 ue midrib not forming at M. dependens 5-13 cm ae inal mucron; petioles 0.1—2 cm oar . Branchlets usually persis eee ferrugineous to rufous tomentose; rib of leaf blade usually glabrous above ........... a. M. coriacea cae. coriacea . Branchlets glabrous, usually ferrugineous only along margins of ter- minal buds and petioles; midrib of leaf blade ferrugineous tomentose above 2b. M. coriacea subsp. nigrescens ; jaa petioles and midrib of leaf blade glabrous or glandular-granulose leaf surface prominently punctate and punctate-lineate, pe large bydropores scattered over the abaxial surface, leaf margins flat t inrolle ssile stign 3.M. pelucidepunceat Lower leaf Sree conspicuously but not prominently punctate or pron nently punctate-lineate, bearing small hydropotes densely ose over the abaxial surface, leaf margin usually inrolled to revolute; stigma sessile or stylopodic. a 5. nee sHoots 5-13 mm long, 4-5 mm wide; flowers 3-5 mm long; fr 4—6 mm diameter . M. calcarata 5. ere . 1.5-4 mm long, 1.5—3 mm wide; flowers 2.2—3.2 mm long; fruits 3—4.5 mm diameter. 6. Branchlets guage! lenticellate; inflorescence umbellate; pedi- m long; stigma on a stylopodium; fruits globose to depressed eee plants of premontane, montane or cloud forests (500—3,000 m), Mexico to Panama 6. Branches without prominent lenticels; . Juergensenii inflorescence fasciculate; pedicels 0.5—1.5 mm long; stigma sessile on the ovary; fruits ellip- Stipa 173) soid; plants of lowland coastal forests and mangroves (0-300 m), along the coast of the Gulf of Mexico and the Caribbean, Mexico to 6. M. cubana Panama 1. Myrsine dependens (Ruiz & Pav.) Spies Syst. Veg. 1:664. Caballeria dependens Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 281. 1798. aera dependens (Ruiz & Pav.) Roem. & Schult., an Veg. 4:506. 1819. oe dependens Ruiz & Pav.) Mez in Engl., Pllanzenr. 9(1V. 236):377. 1902. Type: TUANUCO: Near Muna, H. Razz GJ. Pavon s.n. (AOLoTYPE: MA, IsoryPeEs: F, a Myrsine ciliata Kunth in H.B.K., Nov. Gen. Sp. 3:248. 1819. Rapanea ce (Ruiz & ing. var. Say ume uatrec., Rev. Acad. Colomb. Ci. Exact. 8:324. Pa TOD 1: Tyre: COLOMBIA. Without locality, A. von Humboldt & A. Bonpland 326 (HOLOTYPE, P). Samara myrtifolia Willd. ex Schult. & J.-H. Schult. in Roem. & Sone Mant. 3:220. Prodr. 8:103. 1844. Type: 1827. Myrsine myrtifolia (Schult. & J.H. Schult.) A. DC. COLOMBIA. Without locality, A. von Humboldt & A, Bonpland s.n. ee PE: P). According to TL-2, the third volume of Mavtissa was co- oe: i Julius Herman Schultes, son si taser August Schultes, in their revision of the t. Veg. chat Josef and Johann Jakob Roemer had written earlier. Therefore, we a ‘ch: inged the litera- e citation pees used for the basionym oe gly. Caballo myrtifolia Ruiz & Pav. ex A. DC. in DC, Prodr. 8:102. 1844. pro syn, nom. inval, TYPE: ee Wid ea ality, H. Rurz & _ Pavon sn. CAOLOTYPE: G-DEL; IsOTYPES: G-BOIS, ede myrtotdes ania eo Pl. t. 87. 1852. Rapanea myrtordes (Hook. f.) Mez in Enel., PHanzenr. ne 236): 377. coe Rapanea ciliata (Kunth) Cuatrec. var. myrtoides (Hook. f.) Cuatrec., Trab. Mus. Nat. Ci. Nat. Ser. Bot. 33:108. 1936. Teas dependens (Ruiz & Pav.) Mez in Engl. var. myrtoides (Hook. f.) Cuatr cad. Colomb. Ci. ixact. 8:325. LOS1L. Type: COLOMBIA. Narino: Volcan ¢ Q arade PEL K Rapanea pittiert Mez in Engl., Lundell, Phytologia 48:142. 1981. Type: COSTA RICA. Cartaco: Volcan Irazi, - 14111 (HoLoryre: B-destr.; LecrotyPe: by Pipoly 1992b, BR; isoLecroryPt , Rev. Aca € an He m, W. Jameson jon Pflanzenr. OPV. 236):378. 1902. Myrsine pittier’ (Mez) 3,000 m, H. Pittier Myryine peruviana (Lundell) US). Rapanea peruviana Lundell, Wrightia 6:117. 1980. SYN. Nov. Lundell, Phyctologia 48:142. 1981. Type: PERU. Cuzco: Tres Cunees. upper ae of urque Nacional de Manu, 1-13 km NW of Paucartambo- pees Road, 3,300 3,500 m, 29 Jun 1978, A. Gentry et al. 23478 (HoLotyprE: LL-TEX; tsorypes: MO, USM). — pittiert Mez var. chirripoensis Suessenguth, Bot. Jahrb. Syst. 72 . Type: COSTA RICA. San José: Chirrip6é Grande, 28 Apr 1932, W. Kupper 1140 oe LOTYPE: M; isorype: M.) Considerable leaf variation occurs in Myrsine dependens throughout its range reminiscent of that found among Asian species distributed over broad archipelagos. Populations corresponding to Myrsine ciliata are notable only hile in bud. Popula- 7 :281. 1942. Syn. for their longer glandular cilia along the leaf margins w tions corresponding to the types of Samara myrtifolia, Caballeria myrtifolia, and Myrsine myrtoides are notable only because the leaf blades are cartilaginous in texture, much smaller, and almost plicate. It is interesting to note that 5 RICKETSON AND Pipoty, Myrsine in Mesoamerica 583 exactly the same leaf variation is seen in taxa from New Guinea, where R ip A) P77 args yaIVAIY IA Mez 1S a vegetative match to the type of Myrsin dep ay) lens, while Rapanea velutina corresponds to Myrsine myrtoides (both Rapanea spe- cies will be transferred to Myrs/ne). In other words, quantitative foliar varia- tion was responsible for their segregation in each case. The type of Rapanea pittieri was distinguished by Mez solely because of the epunctate (vs. punc- tate) dorsal side of the anthers. Our study of interpopulational variation re- vealed that the difference is a matter of conspicuous punctations (Costa Rican populations) vs. prominently raised punctations (Colombian to Pe- ruvian populations). While carrying out fieldwork in Costa Rica and Co- lombia, I have found that this varies even within populations. Populations corresponding to the type of Rapanea peruviana Lundell have a denser and deeper red-colored tomentum than the average, and those of Rapanea pittiert var. chirripoensis are in all ways exact matches to the type of Myrsine dependens. Such a broad ranging, polymorphic species has been termed a “polymor- phic ochlospecies” by White (1962), Prance (1982), and Pipoly (1983). Variation in these species is not clinal or necessarily ecotypic; rather it appears to be a series of uncorrelated permutations in quantitative characteristics. 2. Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult. As here circumscribed, Myrsine coriacea 1s the most widespread neotropical species and perhaps the only “weedy” neotropical member of the Myrsinaceae. It is the most polymorphic of the ochlospecies found in any myrsinaceous genus. While vegetative plasticity is greater in this species than in any other in the family, the conic-morchelliform stigma is unique. Among the other distinguishing features of this species are the few-flowered, sessile, fasciculate inflorescences, subsagittate anther and antherode bases, the per- sistent ferrugineous tomentum of the vegetative organs and pedicels, and the small, globose fruits. The only consistent difference we can find be- tween populations corresponding to the type specimen of Myrszne coriacea and those corresponding to that of M. wigrescens (Lundell) Lundell, is that in populations of the latter, the persistent tomentum is restricted to the peti- oles and adaxial leaf midrib, and the populations are largely restricted to open areas of wind-swept elfin forests. Therefore, we consider M. nzgrescens a subspecies of M. coriacea, and make the following new combination. 2a. pitas coriacea (Sw.) R. Br. ex Roem. & Schult. subsp. coriacea, ae Me 511. 1819. eg as Sw., Prodr. 1:32. 1788. Rapanea coriaced (Sw.) Mez ea Iban Symb. Antill. 2:428. 1901. Type: JAMAICA. Without locality, oe date, O. Swartz s.n. oe BM). Caballeria ferruginea Ruiz & Pav., Syst. Veg. Fl. Peruy. Chil. 280. 1798. Manglilla ferruginea (Rutz & ee) Roem. & Schult., Syst. Veg. 4:506. 1819. Myrsine ferruginea (Rutz & Pav.) Spreng., Syst. Veg. eee 1825. Rapanea ferruginea (Ruiz & Pav.) Mez in 584 Sipa L7G) Urban, Symb. Antill. 2:429. 1901. Type: PERU. HuAnuco: Near. Mufia, Without date, H. Rurz et_J. Pavin s.n. (AOLOTYPE: MA; IsoryPE: G). Myrsine Laie Kunth in H.B.K., Nov. Gen. Sp. 3:249. 1819. Type: COLOM- BIA. ca: Near a es 1,000 m, Humboldt & Boapland 1908 (HOLOTYPE: P). Myrsine Nie Zahibr., K. K. Naturhist. Hofmus. 7:3. 1892. Rapanea jelskii (Zahlbr.) Mez in Engl., ae pe 36): 79 1902. Type: PERU. Cajamarca: Near Cutervo, von Jelski 15 (HOLOTYPE: W; ISOT Samara saligna Willd. & JH. Schult. in es & Scul, M ant. 3:220. 1827. Myrsine is (Schult. & : H. Schult.) A. DC., Prodr. 8:103. 18 Type: Herb. Willd. 1039 (B WILLD, Myrsine mpricoides Schledl., Linnaea 1833: 525. 1833. SYN. Nov. Rapanea myricoides — Wrightia 3:109. 1964. Type: MEXICO. Jatapa: H. Galeotti 521, 22, 526 (synvypEs BR, F, G, HAL). We defer lectotypification of this binomial until all materials cited in the es and their duplicates can be assembled. Myrtine tomentosa Presl, Reliq. nk. 2:63. 1835. Typr: PERU. HudAnuco: Wichout further locality, T: on SM. tawny PR). Myrsine viridis Rusby, n. Torrey Bot. Club 6:74. 1896. Type: BOLIVIA. La Paz: Mapiri, Jul-Aug | ie i“ Bang ae ate NY). Myrsine guatematensis Gand., Bull. Soc. Bot. France 65:57. 1918. Type: GUATEMALA. ALTA VERAPAZ: Coban, 1,350 m, Dec 1906, H. von Tirckheim 1001 (HoLoryeeE: P; ISOTYPES: FE, G, LL-TEX, MICH, MO). Rapanea rufa Lundell, te 5:298. 1976. SYN. Nov. Myrsine rufa (Lundell) Lundell, ea 48:142. 1981. Type: COSTA RICA. San José: Cordillera de Talamanca, Pacific Slope of the tee Massif, 2,700— 3,000 m, 6 Apr 1969, G. Davidse & R. Pohl 1635 aes LL-TEX; isotype: ). = am — Myrsine vestita Lundell, Wrightia 7:274. 1984. SYN. Nov. Rapanea vestita (Lundell) Lundell, Phycologia 58:490. 1985. Type: COSTA RICA. ee Cordillera de ee slopes between Cerro Echandi and Cerro Burt, 2,600—2,700 m, 24 Aug 1983, Davidse et al. 24018 (HoLorypE: LL-TEX; isorypes: CR, (0), Myrsine microcalyx Lundell, Phytologia 58:277. 1985. a NOV. Rapanea microcalyx (Lundell) Lundell, Phytologia 58:490. 1985. ae PANAMA. C ae Cerro Colo- rado, 24 mi on gravel road from bridge over Rio San Félix, 1,430 m, 22 Nov 1979, T. Antonio Soi (HOLOTYPE: LL-TEX; isorype: MO) As noted above, this highly variable species exhibits great variation in quantitative features of its vegetative parts, and also in such features as the degree of curvature of the leaf base (obtuse, rounded or acute), leaf shape, vestiture thickness and trichome length, number of flowers and internode length. All of these factors are responsible for the various segregates that have been recognized in the past, including the new ones synonymized above. We believe we have assembled material from throughout subsp. coriacea’s range for the first time since Mez’s work in 1902 and only for that reason have we been able to appreciate how broadly variation occurs within and among populations, that has convinced us to adopt a very broad spe- cies concept. One of the new synonyms, Myrsine microcalyx, is a bisexual growth form with consequent quantitative floral variation. 2b. Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult. subsp. nigrescens RICKETSON AND Pipoty, Myrsine in Mesoamerica 585 (Lundell) Ricketson & Pipoly, comb. et stat. nov. Myrsine nigrescens Lundell, Phytologia 56:418. 1984. Rapanea nigrescens (Lundell) Lundell, Phytologia 58:490. 1985. Type: PANAMA. Bocas pr. Toro: Cordillera de Talamanca, headwaters of the Rio Columbre, 6 airline km NW of the peak of Cerro Echandi on the Costa Rica-Panamanian international border, 2,450—2,600 m., 2—3 March 1984, G. Davidse ef al, 25239 (noLotyPE: LL-TEX; isorypr: LL-TEX, MO) We have chosen to relegate Myrs/ne nigrescens to synonymy under Myrsne coriacea because we have not found any unique features by which we may define it. However, a combination of characters, including the tomentum restricted to the margins of leaf buds and petioles, and tomentose midrib of the leaf, are correlated with its habitats in open areas of wind-swept elfin forests, can be used to distinguish a series of populations consistently. There- fore, we recognize those populations as at the subspecific level. Our sub- species concept follows Pipoly (1987). 3: es eee ee age Oerst., oP Meddel. Dansk Naturhist. jobenhavn. 133. Lé OL. Rapanea pellucido-punctata (Oerst.) Mez in Enegl., sas ee 236):393. ce Type: COSTA RICA. cARTAGO: “In monte Irasuense, Orsted legit, Jan 1847, A.S. Oersted. 30” (LecrorypE, here designated: C). Rapnaea panamensis Lundell, Wrightia 4:169. 1971. SYN. Nov. Myrsine panamensis (Lundell) Lundell, Phytologia 48:142. 1981. Type: PANAMA. Panama: San José Island, Perlas Archipelago, Gulf of Panama, ca. 35 mi SSE of Balboa, main road W of Matta Puerco, 9 Nov 1944, LM. Johnston 441 (HoLoTYPE: LL-TEX; tsorypes: GH, , Ud). Myrsine sytsmae Lundell, Phytologia 58:278. 1985. Syn. Nov. Rapanea sytsmae (Lundell) Lundell, Phytologia 58:490. 1985. Type: PANAMA. Los Santos: Road to El Cortezo, 100—200 m, 24 Jan 1981, W D'Arcy & K. Sytsma 14355 (HoLoryPe: LL-TEX; sorype: MQ). In the Botanical Museum, University of Copenhagen Herbarium (C), there are six different collections of A.S. Oersted 30 made from three differ- ent locations (Aguacate, Jaris and Irazt in Costa Rica), between January 1846 and November 1847. In Oersted’s original description he states “Fre- quenter occurrit in Costa Rica, sylvarum humidarum incola, alt. 2,000— 5,000. Specimina in montibus Jarvis, Aguacate et Irasu lecta reportavi,” without specifying localities further. Mez (1902) and subsequent authors have failed to designate a lectotype. The C sheet of A.S. Oersted 30 from Irazi, Costa Rica, of Jan. 1847, is here selected as the lectotype because it is by far the best of the six collections, and has the greatest number of dissections and notes by Oersted associated with it. The two Lundell species here synonymized represent populations differ- ing only by quantitative characters that are not constant over the range of the species. Myrsine panamensis is notable only by the early glabrescent branchlet apices with small trichomes slighly more reddish than other popu- lations of M. pellucidopunctata. Populations corresponding to the type of 586 Sipa 17(3) Myrsine sytsmae are notable only for its narrow leaves, and somewhat shorter petioles. None of these populational characteristics can be consistently used to define taxa and therefore, we relegate Myrsine panamensis and M. systsmae to synonymy. 4. Myrsine calcarata (Lundell) wate ae Phytologia 48:142 (1981). alae calcarata Lundell, Wrightia 5:2 6. Type: COSTA RICA. ALAJUELA: S side of Volcan Pods, upper slopes, on sae e a past weather station, 24 Mar 1972, D, Stone 3141 (HOLOTYPE: LL-TEX),. 5. Myrsine juergensenu (Mez) Lundell, oo 48:142 1981. lai juergensentt Mez in Engl., PHanzenreich 9 36):388. Type: MEXICO. Oaxaca: without specific locality, date, H. sae ee (LECOTYPE, here ce eerie ISOLECTOTYPES: G, ) Rapanea allenit Lundell, Wrightia 4:168. 1971. SyN. Nov. Myrsine allenii (Lundell) Lundell), Phytologia 48:142. 1981. Type: PANAMA. Cuirigui: N forested face of Cerro Copete, an E spur of Chiriqui Volcano, 2,438-2,591 m, 29 July 1947, PH. Allen 4869 (HoLoryPE: US, LL-TEX-fragment; tsorypes: G, MO). Myrsine chiapensis Lundell, Phytologia 58: 275. 1985. SYN. Nov. Seabees Lundell, Phyrologia 58:489. 1985. Type: MEXICO, Cuiapas: Municipio of La Trinitaria, E of Laguna Tzikaw, ene Bello ae ene Li 300 m, 13 May 1973 D. Breedlove 35246 (Hotorypr: LL-TEX; tsoryee: , MO). ee Lundell, Sy ee 275. 1985. NOV. ee (Lundell) “un Phytologia 58:489. 1985. Type: PANAMA. Cocte: El Valle, Mesa, 6-6.5 km ae Main Street, ca. 1,000 m, 16 Jan 1977, J. Folsom 1352 (HOLOTYPE: MO ISOTYPE: LL-TEX bee Ally Laneelh Phytologia 58:276. 1985. Syn. Nov. Rapanea gillyi (Lundell) ell, meee 58:489. sen Type: MEXICO. Nayarir: Mirador de ie a, Ca, 14 mi N of Tepic, 450-600 m, 21 Aug 1959, C. Fedema 840 (HoLorypE: LL-TEX). Muyrsine costaricensis Lundell, Phy i 58:276. 1985. SYN. Nov. ie COStAVICENSIS (Lunde re ll, Phy loen. oo 489. 1985. Type: COSTA RICA. San Jost: Along gravel road, 2 km N of Hwy. 12, ca. 10 km W of oe Hwy., 2,200 m, 14 Aug 1977, T. Croat 43380 hoe eae LL-TEX; isoryee: Nea nels Lundell, Phytologia 58:277. 1985. SYN. Nov. a yatiscensis (Lundell) Lundell), Phytologia 58:490. 1985. Tver: MEXICO. Jatisco: Sierra de Caule, SW of Talpa de Allende; SW of the prominent peak called Piedra Rajada, S- and W-facing slopes, 2,200 m, 19-21 Nov 1952, R. McVaweh 14336 (HoLtoryPE: LL-TEX; isoTyPe: MICH- n.v., MO). Rapanea mexicana Lundell, ps 5:296. 1976. SYN. Nov. Myrsine mexicana (Lundell) Lundell, Phytologia 48:142. 1981. Typr: MEXICO. Cuiapas: see near Escunitla, 25 Jun 1941, £. ae 5998 (aes LL-TEX; tsoryprs: A, EF, US). Rapanea reflexiflora Lundell, Wrightia 5:297. 1976. SYN. NOV. he rier a) (Lundell) Lundell, Phy jue 48:142. 1981. Type: PANAMA. Coc it: Woods adjacent to chicken farm, La Mesa, above El Valle, 3 Jan 1974 , J. Dwyer 11885 (HoLotype: LL-TEX; IsOTYPES: BRIT, F, ao. PMA). ’ Mez (1902) cited several specimens, including: /#rgensen 326 and 570 (both staminate), Ji/rgensen 338 (pistillate), all from Sierra San Pedro Nolasco; Cuming s.n. anno 1859 (staminate), Ghiesbreght s.n. anno 1842 (staminate), RICKETSON AND Pipoty, Myrsine in Mesoamerica 587 and Galeotti 530 (pistillate) and 53/7 (staminate), all from Oaxaca. The bulk of the material was scanty, except for Ga/eotts 530, that had the most com- plete set of duplicates in the best condition, all of which had been seen by Mez. The US sheet was in the best condition by far, and therefore is se- lected as the lectotype. Myrsine juergensenii has heretofore been considered a species of northern Mesoamerica, with a range from Mexico to Honduras. During preparation of the treatment for Flora de Nicaragua, it became obvious that many of the entities described from Panama were actually extreme forms of Myrsine juergensenit, either from lower altitudes, isolated mountain ranges, or other habitat extremes. Many of the variations encountered in those populations, as represented by the respective type specimens, also occured sporadically in populations from Costa Rica, where this species had never been reported before. A series of quantitative variations in the vegetative features of this species’ populations and other variations directly attributable to sex ex- pression have given rise to overdescription. For example, the types of Rapanea allenii and M. costaricensis are notable only for its slightly longer pedicels and calyx lobes slightly narrower than most (but not all) of the M. jwergenseni populations. The types of Myrsine chiapensis and M. gillyi are notable only for variation directly attributable to sex expression, such as thicker, shorter sepals, and stouter pedicels than most of M. jwergensenii’s populations. The yellow-greenish color of the dried leaves in M. chiapensis and M. gillyz are drying artifacts. Populations corresponding to the types of Myrszne galiscensis and Myrsine mexicana differ from those of the average population for the species only by having thinner sepals. Populations corresponding to the type of Myrsine coclensis has fewer flowers per fascicle and larger, thicker leaf blades, both features typical of premontane Myrsinaceae, especially those growing in low light regimes. The types of Myrszne coclensis and Myrsine reflexiflora, collected from the same locality, differ only because they repre- sent staminate and pistillate growth forms of the population, respectively. 6. Myrsine cubana A. DC., Ann. Sci. Nat. Bot. 16:86. 1941. Type: CUBA. Without locality, 1829, R. de la . 243 (HOLOTYPE: G-DC). 4 Sideroxylon punctatum Lam., Tabl. 2:42. 1794. Bumelia punctata (Lam.) Roem. & Schult. in L., Syst. Veg. 498. ma 7 yrsine floridana A. DC., Trans. Linn. Soc. Lon- don, Bot. 17:107. 1834, nom. superfl. Myrsine punctata (Lam. ‘Steen, Bull. British Mus. Nat. Hist.), Bot. 4:177. 1969, non (H. Lév.) Wilbur, Pacific Sci. 19:522. 1965. Rapanea punctata (Lam.) Lundell, Wrightia 4:121. 1969. Type: U.S.A. Froripa: A. Michaux s.”. (HOLOTYPE: G-DC). Myrsine veraecrucis Lundell, Phytologia 56:418. 1984. SYN. Nov. Rapanea veraecructs (Lundell) Lundell, eae 490. 1985. Type: MEXICO. Veracruz: Municipio Alvarado, Km 22, Veracruz-Alvarado Hwy., 20 m, 21 May 1977, J. Calzada 3200 (HOLOTYPE: F; ISOTYPE: ae n.v.). 588 Sipa 17(3) Stearn (1969) discovered that the earliest applicable name for this spe- cies was Sideroxylum punctatum Lam., so he made the combination Myrsne punctata (Lam.) Stearn. Unfortunately, that epithet was preoccupied by a combination published in 1966 by Wilbur. Therefore, the next available name 1s Myrszne floridana A. DC. However, A. de Candolle cited Sideroxylon punctatum Lam. asa homotypic synonym of Myrsine floridana. Myrsine floridana A. DC. is a nomum superfluum, and the correct name is M. cvbana A. DC., the oldest available name. In the past, Myrs/ne cubana has been confused with Myrsine guyanensis (Aubl.) Kuntze, a South American species from Brazil, the Guianas, and eastern Venezuela. To compound the misinterpretation, Mez (1902) and all subsequent authors have confused Myrsine guyanensis (Aubl.) Kuntze with M. /atifolia (Ruiz & Pav.) Spreng., the latter restricted to the contiguous Andean mountain chain. Pipoly (1992a, 1992b) dis- cussed the diagnostic features separating these three taxa. In addition, Y.B. Harvey & Pipoly (1995) have further elucidated the systematic position of Myrsine monticola Martius, a name formerly considered a taxonomic syn- onym of M. cubana. The type of Myrsne veraecrucis is both quantitatively and qualitatively a perfect match to Caribbean populations of Myrsine cubana in all aspects. We hypothesize that Lundell’s comparison of this species to Myrsine mexicana may be responsible for his mistaking the Vera Cruz populations as a new species. ACKNOWLEDGMENTS We thank the Missouri Botanical Garden and the Flora Mesoamericana Project, for funding that allowed J. Ricketson (MO) to travel to Fort Worth, where he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Herbarium (LL-TEX) housed at the University of Texas at Austin. With- out access to that critical collection, assembled by C.L. Lundell over a pe- riod of nearly 60 years, the present study would not have been possible. We — are grateful to those who have been so instrumental in assisting us in our work, including Gerrit and Jeany Davidse, Linda Oestry, and Catherine Mayo, (MO), Barney Lipscomb, Lindsay Woodruff and Stacy Miles (BRIT), Billie L. Turner, Tom Wendt, Carol Todzia, Beryl Simpson, and José Panero (TEX). We also thank William Burger, Gerrit Davidse, Charlotte Taylor and Barney Lipscomb for review and helpful comments on the manuscript. —— REPERENCES C. Cuen, C. and J. Prpory. 1996. Myrsinaceae. In: Wu Zheng-yi and P. Raven, eds. Flora of China. Science Press, Beijing and Missouri Botanical Garden, St. Louis. 15:34—38. Harvey, Y. and J. Pirpoty. 1995. Myrsinaceae. In: B. Stannard, ed. Flora of the Pico das Almas,Chapada Diamantina, Bahia, Brazil. Royal Botanic Gardens, Kew, Richmond, U.K Pp. 487-491. RICKETSON AND Pipoty, Myrsine in Mesoamerica 589 Mrz, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich 9(71V. 236):1—4 Pipoty, J. 1983. Contributions toward a oe of Cybianthus pine TL A revision of subgenus Laxiflorus. Brittonia 35: 1987. A systematic revision of sage: Ye bianthus subgenus Grammadenia (Myrsinaceae). Mem. New York Bot. Gard. 43:1— —___——*d'991. Systematic studies in the genus Myrsine L. (Myrsinaceae) in Guayana. Novon 1:204—210. 1992a. Estudios en el género Myrsine (Myrsinaceae) de Colombia. Caldasia L7Ly3—2 1992b. Notes on the genus Myrsive (Myrsinaceae) in Peru. Novon 2:392— 407. 1996. Contributions toward a new flora of the Philippines: I. A synopsis of the genus Mrs ee Sida 17:115—162. C. Cuen. 1995. Nomenclatural notes on the Myrsinaceae of China. Novon 51357361 Prance, G. 1982. Forest refuges: evidence from woody angiosperms. In: G. Prance, ed. Biological diversification in the tropics. Columbia University Press, New York. Pp. 137-156 tan, W.T. 1969. A synopsis of Jamaican Myrsinaceae. Bull. Brit. Mus. (Nat Hist.), . 18:884—900. Wire, F. 1968. Geographic variation and speciation in Africa with particular reference to es In: D. Nichols, ed. ad and geography. The Systematics Association . No. 4, London. Pp. 71 590 Stipa 17() BOOK NOTICE Copy, WILLIAM J. 1996. Flora of the Yukon Territory. (SBN 0-660- 16406-X, hbk; ISBN 0-660-15898-1, pbk). NRC Research Press, M-55, National Research Council of Cade, Ottawa, Ontario K1A ORG. $79.95 (U.S.), hbk. $49.95 (U.S.), pbk. 643 pp. Canada’s Yukon Territory, about the size of Alabama, Georgia, Mississippi, and Louisi- ana combined, is a rugged land of mountains (including Canada’s highest peak), plateaus, river valleys, and arctic coastal plain. Many botanists have explored and written about the Yukon. The author of this flora has made some 10,000 Yukon collections (probably the most extensive collection of the territory's plants ever assembled and from many previ- ously un- or cede collected areas). . of Cody’s professional life has been spent in pursuit of plants from north of the 60th parallel, a: only an occasional excursion into more equable climes. He is eminently eee to write this opu The Bok is well made and of attractive format. An introduction briefly describes the territory's terrain and vegetation, gives the number of species and genera in each of the Yukon’s plant families, and outlines the history of botanical exploration there. Then fol- low keys to major divisions and families. The main part of the book treats che families in to genera and species (which worked ~~ — phylogenetic order, with generic descriptions, fine for the 20 species I chose as test cases), and well-done generic and specific descriptions. All species are illustrated, the drawings coming mostly from Porsild Cody's Vascular Plants die ontinental Northwest Territories. Dot maps show the known Yukon distribution of ting the book are a glossary, a bibliography, and an index to scientific each species. Comple would have been more ial than the ce liog- and common names. A “literature cited” raphy,” most entries of which are not referred to in the main part of the book, thus making difficult a reader’s attempts to learn of sources for further reading about the included taxa. The Yukon flora has 80 families, 339 genera, and 1112 species of vascular plants. Fami- ) species (native and introduced) are Poaceae, Cyperaceae, and lies with more than are now treated in an up-to-date, well-done, and teraceae. These and the rest of the flora reasonably priced work. Kudos to Cody.—John W. Thier Sipa 17(3): 590. 1997 NOMENCLATURAL NOTES AND A SYNOPSIS OF MESOAMERICAN STYLOGYNE (MYRSINACEAE) JON M. RICKETSON Missouri Botanical Garden P.O. Box 299 St. Louis, MO 63166-0299, U.S.A. ricketso@mobot.ore JOHN J. PIPOLY II Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-40060, U.S.A. Jpipoly @ brit.org ABSTRACT nomenclatural summary is provided for the species of the genus Sty/ogyne occurring in A Mesoamerica. An updated description of the genus, along with a key to the species is Nisieieae A key to ee Ardisia, and Geissanthus from Stylogyne is also provided. e taxa are recognized, including the new combination Stylogyne turbacensis subsp. laevis (Oetsted) Ricketson & Pipoly. All taxa are enumerated, nomenclatural and taxonomic syn- onyms provided, and six binomials are newly relegated to synonym - RESUMEN Se ofrece un resumen de las especies mesoamericanas Satranen al ons Stylogyne. Se incluye una ay ame actualizada del género, acompafiada de una clave para identificar las especies. Se ofrece también una clave para distinguir Ardisia y as de Sinan, Se reconocen cinco taxa, caver a nueva combinacion Stylogyne turbacensis subsp. laevis (Oersted) Ricketson & Pipoly. enumeran los taxa, se ofrecen siné6nimos tanto nomenclaturales como taxonomicos, y se rele gan. Seis binémenes como nuevos sinénimos taxonomicos. INTRODUCTION The neotropical genus Stylogyne A. DC. comprises 63 species, of which 7 remain undescribed, owing to lack of adequate material. Understanding the systematic biology of this genus has long been problematic because of its sexual lability. Androdioecious, bisexual, polygamo-dioecious and dioe- cious species of Stylogyne have been documented (Pipoly 1989, 1991), as well as the consequent morphological variation due to sex expression. The genus has often been confused with Geissanthus J. D. Hooker (Agostini 1970) and Ardisia Swartz, from which it may be readily separated by the following key: Sipa 17(3): 591-597. 1997 S92 Stipa 17(3) . Calyx open in bud, the lobes 4—5, free or almost free, dextrorsely (rarely sinistrorsely) imbricate, quincuncial or contorted; corolla lobes dextrorsely = rare 2. Calyx and corolla lobes imbricate or quncuncial; corol in bud; ovules mostly pluriseriate, or at times appearing uniseriate when ovules few ina high ancthotactic spiral; inflorescence rachis opaque. ............ Ardisia Calyx and corolla lobes contorted; corolla lobes highly twisted in bud; ovules uniseriate or rarely biseriate; inflorescence rachis translucent pink y sinistrorsely) imbricate, quincuncial or contorted. la lobes not twisted —_~ to hyaline Stylogyne 1. Calyx ae d in bud, rupturing irregularly into 2-8 lobes in the same inflo- rescence at maturity, fused for varying lengths, valvate; corolla lobes val- vate or rarely imbricat Geissanthus In preparing a treatment of the Myrsinaceae for Flora Mesoamericana, analysis of the species in the region revealed that the two most commonly known entities, Stylogyne turbacensis and S. laevis, become inseparable when reviewed over their entire range. The two species, separated by terminal vs. axillary inflorescence in Flora of Guatemala and Flora of Panama (Lundell 1966a, 1971) are now known to overlap significantly in terms of inflores- cence position and morphology and occupy different habitats. Therefore, the latter is reduced to subspecific status. Below we provide an emended description of the genus, a key to the Mesoamerican species, and nomenclator. Stylogyne A. DC., Ann. Sci. Nat. Bot. 16:78. 1841. Dioecious, bisexual, androdioecious or polygamo-dioecious shrubs or small trees. Branchlets mostly glabrous, rarely with simple hairs or translucent lepidote scales on the buds. Leaves alternate; blades mostly glabrous; peti- oles marginate or canaliculate. Inflorescences axillary or terminal, rarely axil- lary and terminal, corymbose or paniculate with the branches racemose or short umbellate-corymbose, sessile or subsessile to long pedunculate, mostly glabrous, rarely with simple trichomes. F/owers unisexual or bisexual, corym- bose or umbellate, (4-)5-merous, pedicellate; sepals dextrorsely contorted and open in bud, free or short-connate basally, punctate and punctate-lineate, generally glabrous, rarely with simple hairs; petals dextrorsely contorted and highly twisted in bud, short-connate, commonly lineate, punctate and punctate-lineate, mostly glabrous; stamens of staminate and bisexual flow- ers usually shorter then the petals, the filaments slender, free from each other and adnate to the petals, inserted at petal base or above, the anthers oblong, subsagittate basally, dorsifixed or basifixed, twisted at anthesis, usually dehiscent _ atrorse longitudinal slits, rarely first by apical pores then widening by slits; staminodes of pistillate flowers similar to stamens but reduced in size, the antherodes devoid of pollen; pistil of pistillate and bisexial flowers obturbinate, the ovary ovoid, the style long, subequaling RICKETSON AND Pipoty, Stylogyne in Mesoamerica o93 or exceeding the stamens, the stigma punctiform, the placenta ovoid, or globose, basal; ovules 3-5, uni- or rarely biseriate; pistillode of staminate flowers lageniform, the ovary hollow, the style subcapitate, less than 1 mm long. Fruit drupaceous, 1-seeded, the endocarp crustaceous or osseous; seed globose or depressed, the endosperm corneous, excavate, not ruminate, the embryo transverse, elongate. A neotropical genus distributed from Mexico through Mesoamerica, the Andes and Amazonia to SE Brazil, and throughout the Caribbean. In Mesoamerica, there are four species known. In general, members of the genus are infrequently found along watercourses in gallery forests in savan- nas, in humid, moist, wet or pluvial cloud forests, premontane forests and lowland forests. Members of the genus tolerate very little disturbance and thus, their presence is useful as an indicator of environmental quality. KEY TO STYLOGYNE OF MESOMERICA 1. Leaf blades bullate, minutely puberulent, the margins serrate .............0.. 1.. S. hayesii 1. Leaf blades smooth, glabrous, the margins entire. 2. Inflorescence axillary, sessile or subsessile, corymbose or racemose. 3. Inflorescence racemose; pedicels 3—3.5 mm; anthers strictly longitu- dinally dehiscent; fruits 5.5-6.5 mm in diameter. ................ 2. S. darienensis 3. Inflorescences corymbose, pedicels 0.5-2 mm, anthers dehiscent by pores opening into longitudinal slits; fruits 4.0-5.0 mm diameter 3. S. glomeruliflora 2. Inflorescences terminal and/or bare paniculate. 4. Leaf blades obovate to ob: te, subcoriaceous, the } spicuous above; a ae nen paniculate, ere terminal, or occasionally terminal with few reduced ean inflorescences acensis subsp. turbacensis . Leaf blades elliptic to oblong, coriaceous, the punctations inconspicuous from above; inflorescences columnar-paniculate, mostly axillary or rarely axillary and terminal or pseudoterminal.4b. S. turbacensis subsp. laevis pee ted hayesii Mez in Engler, Pflanzenr. 91V. 236):272. 1902. Type: . Darien: In chick woods, Manmer Station [Manené], Panama Railroad, ‘ oe a S. Hayes 662 (HoLotyPE: G-BOIS; tsoryprs: BM, K, — Distribution and ecology—Lowland pluvial forests of the Chocé Floristic Province of Panama and Colombia, sea level—600 m elevation. Stylogyne hayesti is rare in Panama, but extremely common in the Andes, on the western slopes of the Cordillera Occidental at the junction of the Chocé and Antioquia Departments of Colombia. The small, axillary inflo- rescences and deeply to shallowly serrate leaf margins are distinctive. 2. Stylogyne darienensis Lundell, Wrightia 5:68. 1974. Typr: PANAMA. Darien: Slopes of Cerro Chucla, drainage of Rio Pavarando, 11 Feb 1972, A. Gentry 4261 (HOLOTYPE: LL-TEX; tsotypes: BM, MO). 594 Sipa 17(3) Distribution and ecology.—Lowland wet forest, ca. 100 m elevation, known only from the southeastern Darién of Panama. oe Na See cea Cuatrec., Revista Acad. Colomb. Ci. Exact. 326. 1951. :, COLOMBIA. Vite bEL Cauca: Drainage of Rio mien Rio Juan, below on ul, 1,300-1,500 m, 19 Mar 1947, J. Cuatrecasas 23847 (HOLOTYPE: F; isoryeE: COL) Distribution and ecology.—Premontane pluvial forest, 1OO—1,500 m el- evation, Choc6 Floristic Province of Panama and Colombia. Inclusion of this species here is the first report for Mesoamerica. 4. Stylogyne turbacensis (Kunth) Mez With the aid of recent collections, most notably from northern Colom- bian populations, we interpret Sty/ogyne turbacensis to include S. /aevis be- cause the inflorescence variation virtually eliminates the differences (ter- minal vs. axillary) used by Lundell (1966a, 1971) to separate them in his treatments of the family for Guatemala and Panama, and necessitating the new combination below. Fieldwork conducted by Pipoly in Nicaragua, Costa Rica, Panama and Colombia has shown that subspecies tvrbacensis tends to be locally common (the individuals clustered) on ridge or hilltops in low- land forests and in premontane forests, while subspecies /gevis is more eq- uably distributed through lowland wet or humid forests and in gallery forests surrounded by savanna vegetation. We follow the subspecies con- cept defined by Pipoly (1987), “groups of populations within a single lin- eage of ancestor-descendant populations that show variation by unique com- binations of plestomorphies, or homoplasic apomorphies, correlated with biogeography and/or ecology. This rank is primarily used to convey infor- mation regarding variation in the life histories of these populations and character state differences hypothesized to be the result of this variation, The subspecific rank in no way attempts to predict speciation events.” The differences regarding staminal exsertion indicated in Lundell’s treatment of the family for the Flora of Guatemala and the Flora of Panama treat- ments are a function of sexual expression, so that in each subspecies, stami- nate flowers have anthers slightly exserced, while pistillate and bisexual ones have them included in the corolla. On the other hand, pistillate and staminate flowers have long styles on the pistil and pistillode, respectively, while in bisexual flowers, the style is shorter. Stylogyne turbacensis (Kunth) Mez subsp. turbacensis. Ardisia tarbacensis Kunth in H.B.K., aa Gen. Sp. 3:244. es Tinus turbacensis (Kunth) Kuntze, Revis. Gen. Pl. 2:975. L891. Sty/og i turbacensis (Nunth) Mez in Engler, Pflanzenr. QTV, 236): 270. ae. Type. COLOMBIA. Antioguia: Near Turbo [Turbaco]. A. von Humboldt & A. Bonpland 1446 (AoLoTYPE: P-BONPL). RICKETSON AND PIPoLy, Stylogyne in Mesoamerica S95 ae tn Willd. ex Roem. & Schult., Syst. Veg. 3:248. 1827. Type. COLOM- nout further locality, aes od, 3005 (HOLOTYPE: B-WILLD, n.v.) Sn mont Blake, Contr. U.S. Natl. Herb. 24:16. 1922. TypzE. GUATEMALA. adas, 19-22 May 1919, H. Pittier — (HOLOTYPE: US; IsoryPE: NY). Paes oaxacana Lundell, Wrightia 4:72. ee SYN. Nov. Type. MEXICO. Oaxaca: istrito de Tuxtepec, C eee and oo 22 1, 1 Jul 1940, G. Martfnez-Calderin 19 (HOLOTYPE: LL-TEX; tsoryPe ’ US alae dete Lundell, Bull, pe Bot. Club 69:398. 1942. Syn. Nov. Type. E {BRITISH HO NDUR AS}: STANN Grass Creek District: Silk Grass Creek a 10 Sep 1939, P. H. Gentle 2990 (HoLotTyPE: MICH; tsorypes: K, LL-TEX, NY). Distribution and ecology.—On hilltops or ridgetops of humid lowland tropi- cal forest, to the premontane transition zone, 0—1,000 m elevation; Oaxaca and Veracruz, Mexico, southward through Mesoamerica to Colombia, then southward to Peru and eastward to the Venezuelan Andes. Populations corresponding to the type of Stylogyne guatemalensis are no- table only for their widely obovate leaf blades and terminal inflorescence relatively longer than most Mesoamerican populations of subspecies turbacensis. Stylogyne vaxacana was described largely because of its relatively smaller inflorescence and sepals, and longer style. The type of Sty/ogyne perpunctata has punctations and punctate-lineations more numerous and more brightly orange in color than the average for the subspecies, but is otherwise identical. All these variations overlap and no correlations can be found with ecology or phytogeography. 4b. Stylogyne turbacensis (Kunth) Mez subsp. laevis (Oerst.) Ricketson ipo y, comb. et stat. nov. Ardisia laevis Oersted, Vidensk. Meddel. Dansk Naturhist. Foren Kjabenhavn 1861:125. 1861. Tinus laevis (Oerst.) Kuntze, Revis Gen.PI. 2:974. 1891. cee laevis (Oerst.) Mez in Engler, Pflanzenr. 9([V. 236): 268. 1902. Type. COSTA RICA. Cartraco: “Monte [Irazt] Irasu, Jan 1847 A. Oersted 28 (LECTOTYPE by Lundell (196Gb): C). Ardista ramiflora Oerst., Vidensk. Meddel. Dansk Naturhisc. Foren Kj@benhavn 1861:125 Tinus ee (Oerst.) O. Kuntze, Revis. Gen. Pl. 2:975. 1891. Sty/ogyne ramiflora (Oerst.) Mez in Engler, Pflanzenr. 9(1V. 236):272. 1902. Type. NICARAGUA. pe prope Tortuga, Apr 1847, A. Oersted 31 (LicrorypsE by Lundell, Pasae C) zin Engler, ue QTV. 236):92. 1902. SYN. Nov., on Stylogyme eerie Blake, Contr. U.S. Natl. Herb. 24:16. 1922. Type. NICARAGUA. CHONTALES: Acoyapa, 15 Feb 18+ A] arn Aioyapa, misinterpreted by Mez}, E. von Friedvichsthal 857 (AOLOTYPE: ee reticulata Mez in Engler, Pace QTV. 236):269. 1902. SYN. Nov. Type. OLOMBIA. Cauca: Near Popayan, J. igh Ja (HOLOTYPE: P). Sm NICAVAQMENSIS ee ean 3:110. 1964. Syn. Nov. Type. NICARAGUA. Zelaya: La Esperanza, Rio Grande, 0-15 m, 10 Apr 1949, A. Molina 2125 (HOLo- TYPE: F) Stylogyne standley: Lundell, Wrightia 3:110. 1964. Syn. Nov. Typr. PANAMA. CANaL ZONE: Barro Colorado Island in Gatun Lake, less than 120 m, 18—24 Nov 1925, P. Standley 41048 (HOLOTYPE: US; isorypeE: BM). 596 Stipa 17(3) Distribution and ecology.—Moist to wet lowland tropical forest and gal- lery forest in savannas, 0-800 m. Veracruz, Mexico, southward through Mesoamerica to Colombia. Populations corresponding to the type of Stylogyne ramiflora have larger leaves and correspondingly smaller inflorescences, with stamens inserted higher on the petals than the average. Those features are common to stam1- nate plants of the subspecies in scattered locations throughout its geographic range. The type of Ardisia guatemalensis is in fruit, and thus, Mez may sim- ply have mistaken it for an Ardisia species, because it posseses no other notable feature. Populations represented by the type of Sty/ogyne reticulata Mez are diminutive in terms of inflorescence and leaf size. Populations from the eastern coast of Nicaragua corresponding to the type of Stylogyne nicaraguensis are notable only for their prominent orange punctations and punctate-lineations, rather than the more brownish colored punctations and lineations found more commonly throughout the range of the taxon. The type of Stylogyne standley? is notable only for its larger leaves and inflo- rescences. These variations, uncorrelated with habitat and based mostly on quantitative characters, all overlap when materials are examined through- out the range of the subspecies. ACKNOWLEDGMENTS We thank the Missouri Botanical Garden and the Flora Mesoamericana Project, for funding that allowed J. Ricketson (MO) to travel to Fort Worth, where he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Herbarium (LL-TEX), housed at the University of Texas at Austin. With- out access to that critical collection, assembled by C.L. Lundell over a pe- riod of nearly 60 years, the present study would not have been possible. We also thank the curators of the herbaria cited for loans of specimens. We are grateful to those who have been so instrumental in assisting us in our work, including Gerrit and Jeany Davidse, Linda Oestry, and Catherine Mayo, (MO), Barney Lipscomb, Lindsay Woodruff and Stacy Miles (BRIT), Billie L. Turner, Tom Wendt, Carol Todzia, Beryl Simpson, and José Panero (TEX). Reviews of the manuscript and helpful suggestions by William Burger (F), Gerrit Davidse (MO), Charlotte Taylor (MO) and Barney Lipscomb (BRIT) are also gratefully acknowledged. — REFERENCES AGostiNi, G. 1970. Notes on Myrsinaceae. I. Generic assignment of Conomorpha sodiroana Mez, Ardisia ambigua Mart., and related species. Phytologia 20:401—403. LUNDELL, C.L. 1966a. Flora of Guatemala: Myrsinaceae. Fieldiana, Bot. 24:135—200. 19G66b. Studies of the American Myrsinaceae- I]. Wrightia 3:192—-199. 1971. Flora of Panama: Myrsinaceae. Ann. Missouri Bot. Gard. 58:285—353. RICKETSON AND PIPOLy, Stylogyne in Mesoamerica DoT Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich 9(1V. 236):1—437. Pipoy, J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia a Mem. New York Bot. Gard. 43:1—76. 1989. Notas sobre el género Stylogyne A. DC. (Myrsinaceae). Ernstia 53:1—9. —________.. 1991. Stylogyne a at (Myrsinaceae): a new androdioecious species from Amcsoua. Novon 1:202- 598 Sipa 173) BOOKS NOTICES FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED Nations. 1994. FAO eed Review—1989-90. (ISBN 92-5-103608-X, pbk). Food and Agriculture Organization of the United Nations, Viale delle Terme di Caracalla, 00100 - Rome, ITALY. Price not given, 543 pp. From the cover: “This publication contains worldwide information on seeds and seed technologies such as varietal improvement and development; variety evaluation, registra- tion and release; quality control; production; processing; storage; marketing and distribu- tion, promotion; research and training. The information has been compiled from official FAO questionnaires sent to the authorities concerned in member countries. In total 86 countries returned the completed questionnaires. The information provided here will con- tribuce co better knowledge of seed status throughout the world.” Reeb, Richard K. 1995. Prophets of Agroforestry. Guarani Commu- nities and Commercial Gathering. (ISBN 0-292-77067-7, hbk). University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, U\S.A. 512-471-4032. $37.50. 264 pp, 8 b&w photos, 12 figures, 5 tables. “The indigenous Chiripa (Guarani) people of eastern Paraguay maintain ethnic autonomy and economic power through the practice of commercial agroforestry. In this study, Reed investigates the relation of this complex economy to the Chiripa social organization and offers ground-breaking insights i into the relations among indigenous groups, nation-states, and the international economy.” Contents include: Preface and Acknowledments; Intro- duction; Yerba, Society, and ne State in Mbaracay; Kinship, Households, and Commu- uty; Leadership and Religion; Chiripa Agroforestry; Patrones, Capataces, and Caciques; he lusions; Postscript: The Chiripa and Recent Changes; Notes; References; and Index. TUREKIAN, Kart K.. 1996. Global Environmental Change. Fourth Edi- tion. ISBN 0-13-478116-3, pbk). Prentice-Hall, Inc., Simon & Schuster/A Viacom Company, Upper Saddle River, NJ 07458, U.S.A. Price not given, 200 pp. “From the preface: “This book aims to address the issues of natural and human induced or accelerated environmental change on the global scale. This book consists of 11 chapters, each centered around a topic in global change that crosses over the clasic disciplinary boundaries.” Contents include: Preface; Prologue; The Changing Planet; Chronology; The Evolution of the Atomsphere; Temperature Variation over time; The Circulation of the Atmosphere and Oceans; Sea Level; Carbon Dioxide, Methane, and Global Warming; Chlo- rinated Fluorocarbons (CFCs) and Stratospheric Ozone; Acid Rain and Tropospheric Ozone; — Human Migration, Population Growth, and Environmental Change; Natural Catastro- phes and the History of Life; Epilogue; and Inc Stipa 17(3): 598. 1997 FLORISTIC COMPOSITION OF THE SAN JOSE DEL CABO OASIS, BAJA CALIFORNIA SUR, MEXICO JOSE LUIS LEON DE LA LUZ, RAYMUNDO DOMINGUEZ CADENA, MIGUEL DOMINGUEZ LEON, and JOSE JUAN PEREZ NAVARRO Division de Biologia Terrestre Centro de 1 i. Bioldgicas del Noroeste ». Postal 128, La Paz Baja California Sur 23000, MEXICO ABSTRACT The wetlands in the Baja California peninsula can be thought of as biological islands because they are surrounded by an arid environment. Also, because of the small area and regular exposure of the wetlands to — Hoods, endemic species are rare. The oasis of San José del Cabo is close to the ocean, but the water has relatively low levels of salinity. This oasis is the remnant of a small river chat existed at the beginning of this century but which dried up from anthropogenic causes and increasing aridity in the region. Since chis oasis is in one of the major tourist developments in Mexico, the aquifer will be affected in the future because of unregulated use to satisfy needs of tourism and of the increasing population. The objective of our study is to document the floristic composition of the wetland to serve as future reference. Our checklist contains 109 taxa of vascular plants. Of these, 78 (71%) are herbs (annuals and perennials). Only 4 (3%) of the taxa are ne distributed in the 30 hectares of this study (16 hectares of which are water). Kryworps: Oases, Baja California, Flora RESUMEN Los humedales de Baja California Sur pueden considerarse islas bioldgicas debido a que con sus relativamente elevados niveles de humedad se encuentran rodeados de territorios dridos; sin embargo, debido a la baja superficie que presentan y la exposici6n a inundaciones catastroficas, su composicion floristica es pobre en endemismos. El oasis de San José del Cabo se encuentra pr6ximo al mar, pero la intrusién de agua salobre es baja. Este oasis es el Ultimo vestigio de un rfo existente todavia a principios de este siglo, que se secé debido a causas antropogenas e incremento de la aridez en esta region. Debido a que €ste se encuentra dentro de uno zona de los grandes desarrollo turfstico de México, el acuffero que lo abastece sera afectado en los afios ee debido a la sobreextraccién de agua para satisfacer las demandas del turismo y de la poblacién en expansi6n. El objetivo del presente estudio es documentar la composici6n floristica actual en este oasis de modo que sirva como referencia para el futuro. El listado anexo comprende 109 taxa de plantas vasculares, de éstas el 78 (71%) son herbdceas (anuales y perennes), pero sdlo el 4 (3%) estan uniformemente distribuidos en las 30 hectdreas consideradas en este estudio (16 de ellas son de agua). Sipa 17(3): 399-614, 1997 600 Sipa 17(3) INTRODUCTION Baja California Sur is the Mexican state with the least available freshwa- ter. This is because of low rainfall and also the small number of permanent waterways (spring, seeps, pools, rivers, lakes) and reservoirs, some of which are locally called “pozas” or “oases” (Escurra et al. 1988; Grismer and McGuire 1993). Additional causes of the aridity are the predominance of coarse-textured soils that favor water percolation, the high levels of evapo- ration from the soil, the lack of high mountains, and the narrowness of the peninsula, which provides little surface area for hydrological basins. Some authors, such as Axelrod (1979) and Murphy (1983), suggested an evolv- ing climate on the peninsula. This change has allowed the climate to vary from a mesic subtropical environment through most of the Tertiary to the current xerophytic environment. If true, the oases could be interpreted as — relict and disjunct mesic habitats. The biological relevance of these small areas is in their biotic composi- tion, which can be found only in places with constantly available water, in mesic habitats outside the xeric peninsula, or on mountain tops where tem- peratures are cooler and precipitation is greater than in the surrounding desert. The social and economical importance of these areas is obvious. Currently, there are dozens of oases in the peninsula, some of them iso- lated and others grouped. Some of the best known are San Ignacio (27°25'N, 112°52'W, elev. 150 m), Comondti (26°02'N, 111°49'W, elev. 400 m), La Purfsima (26°10'N,112°07'W, elev. 150 m), Mulegé (26°53'N, 112°02'W, elev. 2 m), Las Pocitas system (around 24°30'N, 111°00'W, elev. 100 to 150 m), and San José del Cabo (23°03'N, 109°41'W, elev. | m). Some of these are close to the ocean; others are fed by water from the nearest moun- tains or by springs. There are no geo-hydrological studies from which to determine their dynamics. Other wetlands, oases, or estuaries have vanished in the last few decades, some because of water extraction in the hydrological basins to satisfy in- creasing demands of the local populations. It is possible that the disappear- ance of wetlands has been exacerbated by the combination of these de- mands with the natural process of increased aridity in the region (Axelrod 1979; Spaulding and Graumlich 1986). Such wetlands include Todos Santos (23°27'N, 110°18'W, elev. 5 m), Santiago (23°28'N, 109°42'W, elev. 350 m), and Santa Agueda (27°18'N, 112°21'W, elev. 230 m). Other areas may have been wetlands centuries ago, e.g., El Carrizal (23°51'N, 110°10'W, elev. 150 m) and La Ribera (23°34'N, 109°35'W, elev. 10 m). Lugo et al. (1988) pointed out, in a general classification of these areas, that because of the catastrophic floods and ecological limitations of such environment they have a low species richness, or low % diversity, com- De La Luz, er AL., Floristic composition of the San José del Cabo oasis 601 pared to adjacent habitats. Because of the catastrophic floods, the wetlands also have a relatively young structure. Moreover, they are influenced by certain environmental factors that seem to regulate the structure and other processes in the plant groups. Among these factors are the periodicity of floods and droughts, the kinetic energy of the water, the predominance of the water flux, and the nutrient concentration in the soil. The fundamental niche for each species in the wetlands could be affected to a large degree by water dynamics and nutrient factors. Cowardin et al. (1979) proposed a hierarchical system of classification for wetlands that was based on water regimen, water chemistry, and soil types. Each oasis can be considered a pa/ustrine system if it is less than 2 m deep, or a /acustrine system if deeper, if we assume that the water flows slowly and has a small surface area. Using this same classification, these must be designated as “forested wetlands” if palms dominate physiogno- mically, or as “emergent wetlands” if cattails (Typha), reeds, and forbs are dominant. In addition to the desire to preserve these communities for both ecologi- cal and socioeconomic reasons, the study of their plant composition is basic to the proposal of legislation for their use and management. Vegetation around the San José del Cabo oasis, according the Mexican official chart of vegetation (INEGI 1981), corresponds to the sarcocaulescent scrub. Shreve and Wiggins (1964) and Wiggins (1980) recognized this area as a disjointed segment of the Sonoran Desert inside the Cape Region. The oasis is physiognomically dominated by the sky-duster palm, Washingtonia robusta, an endemic species occupying riverside and stream banks in the southern half of the peninsula. In the past, this palm probably was widespread inland where moisture in the soil was relatively high. To- day, these lands are occupied by abandoned or presently cultivated fields and secondary vegetation. In natural conditions, the ground cover could change dramatically according to the incidence of catastrophic floods, which seem to occur at least once each decade. Our objective was to compile a floristic list of the higher plants growing in the vicinity of the San José del Cabo oasis. This list can serve as a baseline reference for additional studies seeking to record quality changes in the vegetation over time. The work was a part of a proposal for legislation to designate this community as a protected area. Such a designation was made by state legislation in 1993. METHODS During May, July, and September 1993, three trips were made to the oasis to collect plants. The oasis was crossed by foot along its river's edge. 602 Sipa 17(3) The river consists of a strip of saturated soil of variable width. We esti- mated a surface of about seven hectares for this area. About 16 hectares of water were traversed by boat to collect aquatic plants. Specimens collected are housed in the herbarium of the Centro de Investigaciones Biol6gicas (HCIB). References used for identification were Gould and Moran (1981), Lot et al. (1986), Shreve and Wiggins (1964), and Wiggins (1980). A checklist was devised beginning with our first field trip. It was considered complete after the third trip when no additional species were found. The life form of each species was noted based on the references cited above. Relative abundance was determined by using the density and distribution pattern shown in Table 1. Finally, because adja- cent land has been used for intensive agriculture, a proportion of the cur- rent Hora came from weeds associated primarily with the crops. Using in- formation from the references and voucher specimens collected elsewhere in the state and peninsula, we classified each taxon of the flora as either indigenous or non-indigenous. DESCRIPTION OF THE OASIS The San José del Cabo oasis is at 23°04'N and 109°41'W at the south- ern tip of the Mexican state of Baja California - According to Garcia (1973), the Koeppen climate formula is BW(h’)w(e). The climate is very dry and very hot, with a mean annual total precipitation (25 years) of 260 mm. Figure 1 shows the geographic location of the oasis. The oasis is separated from the ocean (Gulf of California) by a 40 m sandbar. The water level in the oasis is partially regulated by hydrostatic forces of the tides, but intrusion of salt water is small. The borders of this oasis (and others of the Sonoran Desert) seem to be contracting with the gradual increasing aridity during recente millennia (Van Devender 1990). In addition to depletion of the aquifer, occasional floods from the uplands are sometimes so violent that the oasis is almost swept clean. After such floods, the flora recovers but the conformation of the oasis is altered to some extent. Presently, water covers almost 16 hectares. Taber 1. Abundance critera used for the vascular plants of the San José del Cabo oasis. Value xx has greater dominance (evaluated as frequency, coberture and height) than value x. See appendix. Population Distribution Density Regular Clumped High XXXX XXX Low XX x De La Luz, er AL., Floristic composition of the San José del Cabo oasis 603 USA. Mexico Pacific ne. Ocean secondary vegetation from spring 1 sarcocaulecent r scrubland spring 20° - > Cultivated fields Urban Area enemas Fic. 1. Geographical location of the San José del Cabo oasis at the southern end of the Baja California peninsula, Mexico (upper). Delimitation of the oasis in relation to its suburbs (lower). 604 Stipa 17(3) Nelson (1921) noted, in 1905—1906, the existence of the “San José River,” a stream that arose from a spring about | km upland and flowed into a freshwater “lagoon” (the current oasis), which supported a flourishing agri- cultural area of around 300 hectares. The human population, then, was estimated to be 1600. Rice and sugarcane were some of the more promi- nent crops because of their high water requirements. Currently, this oasis is in one of the more important tourist develop- ments in Mexico, Los Cabos. There are around 6000 hotel rooms, each with a potential consumption of 400 to 500 liters day"! when occupied. There 1s also consumption for domestic purpose by almost 35,000 inhabit- ants. Reuse of water is highly recommended but not yet developed. At first, water to satisfy this demand came entirely from wells, all of them in the watershed of the oasis. In 1992 an aqueduct, which gathers the water pumped by several wells, some in agriculcural valleys upland, was completed to satisfy requirements of both tourists and inhabitants. It has a flow of 2000 to 3000 liters minute'. This expenditure is higher than re- covery by the aquifer and will inevitably drain it. CHECKLIST We did not recognize vegetation types based on physiognomic criteria. Palms, cattails, and reeds have an irregular distribution along the river’s edge. In the attached checklist, each entry is followed by its corresponding life form, abundance, and whether it is native or non-indigenous. Table 2 sum- marizes the life forms of the 109 taxa of vascular plants of this oasis. Most of them, 78 (71%), are annual or perennial herbs; 13 (12%) are shrubs or trees, and 18 (16%) belong to other life forms. Table 3 gives the relative abundance of the species of the checklist. Only four (3.6%) taxa have an abundance pattern of high density and regular distribution, marked in the catalog as XXXX; these are Washingtonia ro- busta, Typha domingensis, Bacopa monnieri, and Phragmites australis. Most of the species, 80 (73.4%), show a pattern of low density and regular distri- bution, marked XX. There is a notable difference between this oasis and the northern ones in the peninsula. This one has a low number of date palms, Phoenix dactylifera, a species introduced into the oases of the peninsula as a food resource by Spanish missionaries in the 16th and 17th centuries. This species survives now as feral groves or with minimal management. Our work establishes the distribution of several taxa not known before for this latitude on the peninsula, i.e., Echinodorus berteroi, Kosteletzkia depressa Phyla incisa, Ruellia inundata, Rumex pulcher, and Xanthinm strumavium, all known in wetlands from 4° to the north or in those tropical areas. Dr La Luz, Er AL., Floristic composition of the San José del Cabo oasis 605 Tabe 2. Life forms of the vascular plants in the oasis of San José del Cabo, Baja California Sur. Life forms No. of taxa Percentage Trees (Tr) 4 3.66 Shrubs (Sh) 9 8.26 Perennial herbs (Ph) 4] 37.64 Annuals (An) 37 33.93 Hydrophytes + Aquatics (Aq) 7 6.43 Vines (V1) 10 9.17 Parasites (Pa) 0.91 Total 109 100.00 Tas_e 3. Classification of the taxa of vascular plants according to their relative abundance in the oases of San Jose del Cabo, Baja California Sur. See Table 1 and appendix. Abundance code number of taxa percentage l (Xxxx) 4 3.66 2 (xxx) 8 7.34 3 (xx) 80 73.40 4 (x) 17 15.60 ~ 109 = 100.00 Only 49 (44.9%) of the species in this oasis are natives (in the appendix marked N). The remaining 60 (55.1%) are non-indigenous species indica- tive of early successional stages or clearly introduced weeds (marked IW) coming from cultivated fields or disturbed areas, such as Amaranthus fimbriatus, Argemone ochroleuca, Cynodon dactylon, Dactyloctenium aegypticum, Datura inoxia, Perityle californica, and Sida xantii. This implies that the wetland has been changed from its original struc- ture. The main sources of disturbances are the agricultural use of the adja- cent land, the cutting down of the sky duster palm for support and roofing of rustic “palapas,” and the severe floods regularly affecting the oasis. All these provide niches for species typical of early succesional stages. An addi- tional clue to support this stacement is the number of species tied to high levels of humidity (pointed out here as “aquatics” [Aq}, of which there are only seven [6.43% Because of the proximity of the ocean to this wetland, one might expect the presence of any of the five mangroves native to the Baja California peninsula, but there are none. For Baja California Hastings et al. (1972) and Wiggins (1980) identified Avicennia germinans, the uncommon Conocarpus erecta, Laguncularia racemosa, the more inland Mayrenus phyllanthoides, and Rhizophora mangle, some of which occur even in higher latitudes in the peninsula up to Bahia de los Angeles at 29°00'N. 606 Sipa 17(3) A possible explanation for the absence of mangroves in this wetland is given by Alvarez-L6pez (1988) and Lugo et al.(1978). Mangroves grow in warm waters of low kinetic energy with highly predictable cycles. In tropi- cal latitudes, they grow in water with low salinity. In subtropical latitudes, they are linked to highly saline water, a condition not prevailing here. From the types of freshwater vegetation in Mexico, Rzedowski (1978) cited the “popal,” “cular,” and “carrizal” as those occurring in hot and tem- perature climates. These last two are poorly represented in this oasis because of the dominance of, Phragmites australis, Scirpus americanus, Typha domingensis, and other monocotyledons. Floating and submersed species are not dominant here. Pofamogeton is a widespread aquatic form in the penin- sula, but it is absent from the oasis. CONCLUSIONS Grismer and McGuire (1993) studied 31 oases in the central Baja Cali- fornia peninsula. Because each has its own physical and biological features, each requires its own characterization. Ezcurra et al. (1988) studied the flora and phytogeography of 17 oases in the hyper-arid zone of the Sonoran Desert, almost all in natural condition. They reported only 26 species of vascular plants, none endemic. Most of these (31%) are taxa distributed in northwestern Mexico and southwestern United States. The hemicryp- tophytes (perennial herbs with their perennating buds at the soil surface) were the dominant life forms (58%). Felger (1980) studied life forms in several oases of the Sonoran Desert. He found 145 taxa, 34% of them annu- als. The life form spectrum found in our checklist is consistent with such results, but dominance of these probably is a sign of disturbance, since 28 of our annuals (73.7%) are considered as introduced weeds. Rzedowski (1978) indicated, in Mexico, that agricultural and industrial activities and water demand for domestic use have affected aquatic com- munities more than any other plant community type. The geographic po- sition of San José del Cabo oasis, i.e., within the tropics, and the permanent nature of the water would seem enough to allow a richer flora than that found. Undoubtedly, human disturbance, e.g., clearing of channels, dredg- ing, fumigation against mosquitoes, fires, sewage flow, and cutting down palms have modified the original plant composition drastically, contribut- ing to modification of both composition and structure. Additional evidence of disturbance in this place is the analysis of the associated fauna. Nelson (1921) quoted for this oasis the endemic rice mouse (Oryzomys cones? var. peninsularis), which Alvarez-Castafieda (1994) consid- ered to be now extinct. This oasis is a community which has largely lost its original composi- De La Luz, ET aL., Floristic composition of the San José del Cabo oasis 607 tion. Its existence is threatened unless the extraction of water is regulated in its watershed and the water is recycled after treatment for nonpotable use. Today, the oasis is an informal park where tourists go for a stroll. Our report represents one of the last opportunities for studying these special communities of the Baja California and the Sonoran Desert. ACKNOWLEDGMENTS This study was done with the funds of the former Delegaci6n SEDESOL of Baja California Sur and the project “Estado Actual y Potencial de Aprovechamiento de los Oasis del Noroeste Mexicano” (SIMAC 94/MA- 001) granted to Dra. Laura Arriaga, CIBNOR. The authors are indebted to Thomas F. Daniel for encouragement and editorial aid. Thanks also to Ellis Glazier for help with the English language text. APPENDIX Annotated checklist of the vegetation of the San José del Cabo oasis, Baja California Sur, México. Family/Species L.F. Abundance Residence MONOCOTYLEDONES Alismataceae Echinodorus berterot (Spreng.) Aq x N Fassett MDE 1502 Arecaceae Phoenix dactilifera L. Tr XX IW No Voucher Washingtonia robusta Wendell Tr XXXX N No Voucher Commelina Sale diantbifolia Del. Ph XX N JLLL 6094 Cyper Cc on ioicus I.M. Johnston Ph XX N JLLL 6116 Cyperis ferax L.C. Rich. Ph XX N JLLL 6117 gs see Brice. Ph XX N 486 C bes sp. Ph XX N JLLL 6075 Eleocharis oe Brite. Ph Xx N RDC 1729 — americanus Pers Aq XX N 489, RDC 1699 si ae (C. A. Mey) Pl ud, He 1723 a 1 Zz 608 Lemnaceae Lenna aequinoctialis Welw. JLLL 6049 Naiadaceae Natas marina 1, A JLLL 6114 yphaceae Typha domingensis Pers. RDC 239 Poaceae Arundo donax L. L998, 999 Cenchrus soe L. RDC 1706 Chloris =. Swartz be Cynodon dactylon (L.) Pers. oO voucher Dactyloctentum aegypticum (L.) eauy, RDC 1708 Digitaria ciliaris (Retz.) Koel JLLL 6090 Distichlis 5 al (L.) Greene JL 84 ae ‘eta microsperma (DC.) Kunth MDL 1479, RDC 1711 Panicum purpurascens Raddi RDC 1718 a sa ee vaginatum Sw. wLL 6115 ease australis Trin. No voucher Potamogetonaceae Ruppia maritima LL. MDL 517 DICOTYLEDONES Acant acea Elytraria ene ‘icata (Vahl.) Pers. JLLL 6092 Ruellia inundata Kuntze JLLL 6091 maranthaceae Amaranthus fimbriatus (Vorr.) enth. 1478 esi ee L. RDC fi ae —- (L.) Mog. RDC 1707 Aq Aq Ph Ph XXXX XXXX IW IW IW Sipa 17(4) De La Luz, et AL., Floristic composition of the San José del Cabo oasis Apiaceae Apinm leptophyllum (Pers.) Ph XX F. Muell. MDL 1477 Eryngium nasturtiifolinum . An XX Juss. ex Delar JLLL 6121 sclepiadaceae Cryptostegia grandifolia (Roxb.) Vi XX R. Br. RDC 1706 Cynanchun palmeri (S. Wats.) Vi XX S.F. Blake JLLL 6119 Boraginaceae Coldenia cuspidata Ph x I. M. Johnstc MDL 1472 Heliotrop YY ASSAVICH ‘ ropium curassavicum L. Ph XX Heliotropium macrostachyum Ph XX (DC.) ; JLLL 6123 emsl Chenopodiaceae Atriplex barclayana (Benth.) Sh Xx ietr. subsp. barclayana MDL 1462 Chenopodium ambrosioides L. An XX Chenopodinm murale L. An XX L 1469 Capparidaceae Cleome lutea Hook. subsp. An XX jonesii (Tidestr.) Macbr. JLLL 6122 Compositae Ambrosia ambrostoides (Cav.) XX N Payne An JLLL 6120 Ambrosia confertiflora DC. Ph XX MDL 1497 Aster exilis Ell. An x LLL 6105 Baccharis glutinosa Pers. An XX RDC 1715 Bidens aurea (Ait.) Sherff. An XX } 14 Conyza bonariensis (L.) Cronquist. An x Eclipta prostrata (L.) L. Ph XXX L 6102 Encelia californica Nutt. Ph XX 609 O10 Gnaphalinm i olor Bioletti MDL 150 Malachi xantit A. Gray MDL 99 any wali nica Benth. DC Paniile aurea 1 Ros MDL 1 Sonchus oleracens L. JLLL 60 Vientera tomentosa A.Gray JLLL 6078 Nanthium strumarim LL. MDL 1476 Cucurbitacea Cucumis i dpc Ehrenb. mee operculata (1..) Cogn, ntermedia Cogn. ex Rose or 1488 i ice ce IE. RDC 1, JLLL 6124 Convolvulaceae Cuscuta baie stris Yunck. DL 14 Ipomoea sp. JLLL 6110 Merremia quinguefolia (L.) Hallier JLLL 6109 Euphorbiacea Croton californ nicus Muell.-Arg. ae wDC Eatin DS is Mla Benth. JLLL 6103 oo communis LL. No voucher Gentiana ce Eustoma eal atum (L.) Griseb. RDC Labiatae iatae Salvia similts Brandegee RDC 1709 Leguminosae Cannavatlia villosa Benth. MDL 1487 C ‘rotalaria mncana LL. 713 Ne spnia ee na (L.) Benth. 47 1 Vi Vi Sh Sh IW Sipa 17( De La Luz, ET AL., Floristic composition of the San José del Cabo oasis Phaseolus atropurpurens Sessé An XX & Moc. MDL 1494 Rhynchosia minima (L.) DC. Vi XX 6106 Loganiac eon paneer A. Gray Sh Xx JLLL 6076 ytrhaceae Ammania coccinea Rottb. Ph XX JLLL 6077 aceae Abutilon incanum (Link) Sweet Ph Xx 1423 Abin sp. An x L 1425 G ne barbadense L. Sh XX 505 Herissantia crispa (L.) Brizicky Ph XX RDC 1698 Horsfordia alata (8. Wats.) Ph Xx A. Gray JL Kosteletzkya depressa (L.) Ph XX Blanch. Fryxell & Gates JLLL 60¢ Matlvastrum ee Ph XX (L.) Garcke MDL 1463 Sida rhombifolia L. Ph XX L 1434 Sida xantii A, Gray Ph XX MD 06 oa ambigua A. Gray var. An XX ambigua ay LLL 61 7 Sphaeralcea emnbate A. Gray var. An XX rosacede inz & ILM. cepa — JLLI Nyctaginaceae Boerhavia coccinea Mill. Ph x O71 olygo Sane Ui bound Michx. Aq XX MDL 1440 Rumex pulcher L. Ph XXX 5095 Onagraceae Gaura parviflora Douglas XX Ph ex Hook. JLLL 6118 IW IW 611 612 Ludwigia octovalvis Jacq.) Raven DC 1697 Ludwigia peploides (Xunth) Raven subsp. peploides JLLL 6060 Papaveraceae Arge zone ochrolenca Sweet JLLL 6053 Plumbaginaceae Plumbago scandens 5 MDL 14 Portulacacceae Portulacca pee L. MDL 142 Portulacca oleracea \.. MDL 1430 Primulacea aceae Anagallis arvensis L. JLLL 6027 Rubiaceae Houstonia arenaria Rose RDC 1670 Salicaceae Papas emt S Wats RDC 1728 Salix ene Benth. MDL 1437 Scrophulariaceae Bacopa monniert (L.) Wettst. JLLL 6124 Solanz Someta mmoxta Mill. DL 1498 Nin otiana glauca R. Graham No voucher Nicotiana 2 wae Dunal JLLL Physal ~ crass en ia Benth. DL 1501 Solanum ial A. Gray MDL | Verbenaceae Phyla incisa Sma JLLL 6101 Vit ia nf (L.) L. LLL 6 = Ph An XXXX IW IW IW Sipa 17(3) De La Luz, eT aL., Floristic composition of the San José del Cabo oasis 613 L. F. = Life Form Abundance Residen fr. Tree (See Tables 1 & 3) oe 7 “Introduced/ Weed Sh. Shrub N = Native Ph Perennial herbs An Annuals Aq. Aquatics/Hydrophytes Vi. Vines (annuals + woody) Pa. Parasites Collectors’ names: (All of the specimens are housed at the HCIB herbarium) MDL: Miguel Dominguez Le RDC: Raymundo ee Gilets REFERENCES ALVAREZ-CaSTANEDA, S.T. 1994. Current status of the rice rat, Oryzomys couesi peninsularis. uthw. Naturalist. 39:99-—100 aoe Lopez, M. 1988. Ecology a Prerocarpus officinalis forested wetlands in Puerto Rico. In: A.E. Luge, ed. Forested wetlands. Elsevier, Amster 127-149. AXELROD, 1979. Age and origin of Sonoran Desert vegetation. Ores. Pap. Calif. Acad. Sci. 132:1-74. Cowan, a V. Carter, EC. Goer, and E.T. LaRor. 1979. Classification of wetlands nd deep water habitats of the United States. U.S. Dept. of the Interior, Fish and Wild- life Service, Office of Biological Service. Washington, D.C. Ezcurra, E., R.S. FeLGer, A.D. Russett, and M. Equinua. 1988. Freshwater islands in a desert saad sea: the eee flora and phytogeography of the Gran Desierto oases of northwestern Mexico. Desert Pl. 9:2:35—44, 55—57. Fricer, R.S. 1980. 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Flora of Baja California. Stanford University Press, Stanford, California. CONTRIBUTIONS TO THE FLORA AND ECOLOGY OF THE NORTHERN LONGLEAF PINE BELT IN RANKIN COUNTY, MISSISSIPPI MICHAEL WAYNE MORRIS Department of Biology North Georgia College & State University Dahlonega, GA 30597, U.S.A. ABSTRACT A study of beech-magnolia forests and adjacent habitats in southern Rankin County, Mississippi, in 1995, resulted in the documentation of twelve species of angiosperms con- sidered rare or imperiled in Mississippi. In addition, populations of several vascular plant species previously known only from localities considerably farther south, either in the southern Longleaf Pine Belt or Coastal Pine Meadows physiographic regions, were discov- ered during the course of field surveys. The rare or unusual species peportedt oie from the extreme northern edge of the Longleaf Pine Belt in central Mississippi are adnatus, ypericum galioi serosum Pe aie sae aninaaiel Ludwigia linearis, ae bien. “Mikania cordifoli id, 1, Persea palustris, Platanthera cristata, Pluchea foetida, Polygala boykinit, ae aie Das: Sabatia Lapa ara, 8 Schisandra glabra, Stewartia malacodendron, Triphora trianthophora, and Verbesina walters. Iris cristata is reported as occurring at the southern periphery of its range in Rankin County. T he associated plant communities of these taxa are also discussed. RESUMEN Un estudio de los bosques de is ieee ag Magne panei y habitats adyacentes en el sur del Condado de Rankin, Mississippi, en 1995, sult sdocucientacign de doce especies de of angiospermas consic ae raras o en peligro en Mississippi. Ademas, fueron descubiertas poblaciones de varias especies de plantas vasculares previamente conocidas s6lo en localidades mucho mas al sur, tanto en el sur de las regiones de Longleaf Pine Belt o de Coastal Pine Meadows, durante los muestreos de campo. Las especies raras 0 poco frecuentes que se citan aqui del extremo norte del Longleaf Pine Belt en Mississippi central son Asver ee Hypericum galtoides, H. setosum, ee dlata, Wlictum floridanum, Ludwigia linearis, Melanthinm — ae cordifolia vax quinguefolinm, Persea palustris, Platanthera cristata, Pluchea foetida, Polygata boykinit, cr ae hystrix, Sabatia sk celia Schisandra glabra, is artia malacodendron, eats trianthophora, y Verbesina walteri. Iris cristata es citado en el extremo sur de su drea en el Condado de Rankin. Se discucen también las comunidades vegetales asociadas de estos taxa. INTRODUCTION Rankin County is located in central Mississippi, due east of and adjacent to Hinds County, in which the city of Jackson is located (Fig. 1). The south- ern part of Rankin County is included in the Longleaf Pine Belt physi- Sipa 17(3): 615-626. 1997 616 Sipa 17(3) Fic. 1. Location of Rankin County, Mississippt. ographic region (Lowe 1921). The flora of this area, as well as other areas in the northern Longleaf Pine Belt of Mississippi, is poorly known. Botanical investigations conducted in the southern part of this physiographic region include Harper (1914), Carter and Jones (1968), Mills and Jones (1969), and Rogers (1977); therefore, a considerable body of information concern- ing vascular plants already exists for that portion of the state. The main purpose of this study was to inventory beech-magnolia forests, referred to as Southern Mixed Hardwood Forests by Ware et al. (1993), and adjacent habitats at the northern periphery of che Longleaf Pine Belt in southern Rankin County, especially for vascular plant species listed as rare or imper- iled by the Mississippi Natural Heritage Program (1996). Springs, bogs, and bay swamps at or near the sources of streams and longleaf pine-scrub oak communities on the surrounding ridges were surveyed as well as the Southern Mixed Hardwood Forests. Characteristic species of each commu- nity type are listed in Tables 1, 2, and 3. Field work in this area has re- sulted in the discovery of additional populations of twelve species of plants considered rare or imperiled in Mississippi, as well as new stations for sev- eral species at the peripheries of their ranges. — Morais, Flora and ecology of the northern Longleaf Pine Belt 617 Ea 4 a > “ wel bed —as YA ae = sat "28 Ne rhs yo 5 Rock outcrops aN S. apes (:Loess> Na — Ses = XS ON - \ Kisatchie Polen 7 a? os = ok 5 Wen 3 : ~ —_ SO eae a Citronelle Formation 7 RS Be ae as yan a ¥ . eS ey =n one wes : 5 SSeS ~ K itman ~ PAS (tronstone at contact) : Awe OY oi eS ES Savannah Quit beg - ae ee ee . ti Le oh 7 ye Spee oe = 20 hee TAS - al = > __ Kickville = nd = en . ee ~~ ~ wo “Sandstone 8 7 yw s cea CN om ce > 4 siltstone outcrops = \\ aes os 7 gt Pe E Kirkville 5 eS 7 ‘oe , a ie, ~~ if wee ae. WOE = ae =e iat ot oe Fat 70% Fa la Formation —s Bae ON Me ah fo “0° , CaLgnon” T! Bi. =" er a 7 ee “ot oo cee mil ie = Z ce a Ni att te ep ab ae Alluvium “On f x : x yor x x Vicksburg Formation x x Z Sree —— i aa aes, a 424 Forest Hill Formation Yazoo Formation Fic 2. Soils and geologic formations in southern Rankin County, Mississippi. Soils in the study area are primarily of the Smithdale-Providence- Kisatchie, Smithdale-Providence, Savannah-Quitman, and Quitman- Kirkville associations (Cole et al. 1987). Providence soils are derived from loessial deposits; Smithdale soils are included in the Citronelle Formation; Kisatchie and Savannah soils comprise the Catahoula Formation in this area; and both Quitman and Kirkville soils are classified in the Vicksburg Formation (Fig. 2). According to Cole et al. (1987), Smithdale-Providence- Kisatchie soils are mostly deep, well-drained, medium acid to extremely acid soils on rugged dissected upland with narrow ridgetops, steep hill- sides, and short drainageways; they also state that shallow to deep ravines and outcrops of sandstone and siltstone are frequently associated. Perme- ability is moderate to very slow, available water capacity is moderate to low, and texture ranges from clay to silt, loam, and sand. Smithdale-Provi- dence soils are mostly well-drained, medium to very strongly acid soils on rolling to hilly upland ridgetops and steep hillsides. Permeability is mod- erate to moderately slow (in the fragipan), available water capacity is mod- erate, and texture is mostly loamy to silty. Savannah-Quitman soils are mostly deep, moderately well-drained, strongly to very strongly acid soils on gently sloping stream terraces and sloping uplands. Permeability is moderate to moderately slow, available water capacity is moderate, and texture is mostly loamy. Quitman-Kirkville soils occupy nearly level areas on stream terraces and floodplains and are mostly deep, moderately well- drained, strongly acid to very strongly acid soils. Permeability is moderate 618 Sipa 17(3 ~— to moderately slow, available water capacity is moderate, and texture is usually a fine sandy loam. Elevation above sea level in southern Rankin County ranges from ap- proximately 220 feet in the floodplain of the Pearl River to about 612 feet on Shiloh Hill (Cole et al. 1987). Both the Pearl River and its tributary, the Strong River, drain this region, and a ridge dividing the two watersheds crosses the area in a southwest-northeast direction. Due to urban expansion in western Rankin County near the city of Jack- son, agricultural practices, and the lumber industry, much of the native vegetation has been drastically altered; however, many sites still exist that are minimally disturbed, especially in areas that are too steep to clear. South- ern Mixed Hardwood Forests (beech-magnolia forests) are well- developed primarily in deep ravines, on north- and west-facing slopes and bluffs above streams, and in rich creek bottoms, The terminology of physiographic regions in Mississippi in the discus- sion of noteworthy vascular plant species follows Morris (1989) as adapted from Lowe (1921). Other works consulted in the compilation of this paper in addition to those specifically cited in the following discussion include Clewell (1985), Godfrey (1988), and Radford et al. (1968). Herbarium abbreviations follow Holmgren et al. (1990) except MMNS (Mississippi Museum of Natural Science, Jackson, Mississippi), NGC (North Georgia College, & State University, Dahlonega, Georgia), and mwm (Michael Wayne Morris, pers. herb.). NOTEWORTHY SPECIES ARALIACEAE Panax quinquefolium L. Mississippi. Rankin Co.: ca 10 km SE of Star, 25 Aug 1995, Morris 4484 (MMNS). Ginseng occurs here in rich hardwoods on a gentle, N- facing slope in the relatively { sa Dabbs Creek bottom. The species is listed as rare or uncommon in Mississippi (MNHP 1996), and it is found at widely scattered localities mainly northern and meee portions of the state in the Loess Bluff, North Central Plateau, Inte- rior Flatwoods, Pontotoc Ridge, Black Prairie, Tennessee River Hills, and Jackson Prairie Regions (Morris et al. 1993). It is now known south to Wilkinson, Hinds, Rankin, Smith, and Lauderdale counties (MNHP 1996). ARECACEAE Rhapidophyllum hyscrix (Pursh) Wendl. & Drude. Mississippi. Rankin Co.: ca 9 km W of Puckett, 26 Aug 1995, Morris 4497 (BE, MMNS). Needle palm is classified as rare in Mississippi (MNHP 1996), and this site on the undulating terrace of Campbell Creek is at the northwestern periphery of the species’ entire range in the southeastern United States (Clancy 1988). This population consists of one large and two small crowns of the same genetic individual on the upper bank of a large stream near its confluence with Campbell Creek. Rankin County was not included in the cotal range of needle palm by Clancy (1988). Morais, Flora and ecology of the northern Longleaf Pine Belt 619 Taste 1. Characteristic species of Southern Mixed Hardwood Forests in southern Rankin County. Acer barbatum A. rubrum avya glabra C. tomentosa Castanea dentata Fagus grandifolta Ligqu ee styraciflna Liriodendron tulipifera Magnolia grandiflora yssa sylvatica Pinus glabra Aesculus pavta Amelanchier ine aes parviflora . triloba es arpinus caroliniana Chionanthus virginicus Cornus florida Erypbrina } herbs Gelseminm ee virens Halesia diptera Hamamelis virginiana cates eae quercifolla a um hypericoides ium ae hes var. ambigua I. ambigua var. monticola I. decidua 1. opaca Antennaria solitaria Arisaema dvacontium A. gquinatum Aristolochia serpentaria CO amaelivinm luteum Desmodium nudiflorim Epifa; Mitchella repens Overstory Pinus taeda Prunus serotina Quercus alba . falcata . laurifolia O. michauxit iia americana Understory Trees, Shrubs, and Woody Vines Uicinm floridanum Lindera benzoi = Magnolia map Ostrya virgin Oxydendrum ar an eum Prunus mexicana Prelea trifoliata Rhamnus carolinie Rhapidophyllum re SEFIX Rhododendr on canescens Smilax i a S. sma Stewartia — odendron Styrax grandifolia Symplocos tinctoria \ APCIVTTSIY arbor, V. elliottii Herbs Panax quinquefolinm hye Q Phryma leptostachye Spigelia marilandica Thelypteris hexagonoptera T. kunthii Tipularia discolor Trillium cuneatum Uvularia perfoliata U. sessilifolia Viola affinis V. palmata V. walteri 620 Stipa 17(3) TABLE 2. Characteristic species of seeps, springs, bogs, and bay swamps in southern Rankin C ounty. Woody Species Acer rubrum Persea Aronia arbutifolta Pinus taede Cephalanthus occidentalis Oren laurifolia Decumaria barbara Q. nigra Hypericum crux-andreae R a ee OM CANESCENS Itea virginica Smilax laurifolta Lyonta ligustrina Toxtcodendron vernix Magnolia virginiana Vaccinium corymbosum Myrica certfera Viburnum dentatum M. heterophylla Vi nudum Nyssa sylvatica var. biflora Herbs Aletris aurea Lysimachta lanceolata Aster sericocarpotdes Melanthium virginicum Bartonta paniculata Mitreola petiola Cacalia ovata M. sesstlifolia Carex a escens Osmunda cinnamonea a buvi O. regalis Cc ‘aananedtied laxum Oxypolis rigidtor Cyperus haspan Panicum microcarpon Drosera capillaris Platanthera ciliaris Eleocharis sone P. clavellata E. tuberculosa P. cristata Elephantopus nudatus Pluchea foetida Eryngium integrifolium Polygala cruciata E palo tum frstulosum P. mariana 1, perfoliatum Rhexta mariana E. votundtfolium R. virginica E. semiserratum Rbynchospora globularis ‘MIVena SQuATV OSA R. glomerata Gratiola pilos R. gracilenta Hydrolea ovata R. inexpansa Hypericum gymnanthum R. variflora A. setosum Sabatia campanulata Hypts ala Solidago patula J uncis bie S. rugosa Lechea Spiranthes praecox Le oo itt Viola primulifolia Liatris spicate Woodwardia areolata Linum striatum Nyris difformts Ludwigia hirtella N. widifolia L. linearis N. torta Lycopodium alopecuroides Morris, Flora and ecology of the northern Longleaf Pine Belt 621 TABLE 3. Characteristic species of longleaf pine-scrub oak communities in southern Rankin County. Woody Species Carya pallida Pinus taeda C. tomentosa Prunus umbellata Castanea pumila Quercus falcata G eanothus ¢ americanus Q. hemisphaerica Celtis tenutfolia Q. marilandica Ilex vomite Q. stellata Malus Se Rhus copallina Myrica cerifera Toxicodendron toxtcarum Nyse sylvatica Vaccinium arbore Pinus echinata V. stamineum P. pa is Herbs Agave virginica Lespedeza hirta Asclepias amplexicantis L. stuevet ticillata Liatris squarrosa Aster adnatus L. squarrulosa A. concolor Mo onde punctata A. linariifolius Onosmodium virginianum Aureolaria pectinata ILVOpsis graminifo 1a Bonamia humistrata Polygala nana Crotonopsts elliptica Preridium aquilinum Cyperus plukenetii Sabatia brachtata anthonia serviced Sn zachyrinm scopavinm Desmodium laevigatum Seymeria casstoides Eryngium yuectfolinm Solidago odora Enpatorinm album Lephrosia virginiana Gymnopogon ambiguus Viola pedata ASTERACEAE adnatus Nutt. Mississippi. Rankin Co.: ca 3 km SSW of Star, 1 Aug 1995, Morris 4390 (IBE). Chainleaf aster is locally infrequent at this locality near sandstone and siltstone outcrops on a longleaf pine ridge. Previously this interesting species was known from sandhills in Jasper (Carraway 1990) and Smith (Lowe 1921) counties and south to the Gulf of Mexico. The Rankin County population extends the documented range of chain] on aster northwestward in Mississippi by about 70 km nia cordifolia (L.f.) Willd. Mississippi. Rankin Co.: ca 4 km NE of Puckett, 21 ee 995, Morris 4459 (IBE, MMNS, NGC, SWSL). Characteristic of bluffs and thickets in rich hardwood forests, this hempweed is considered rare to uncommon in Mississippi (MNHP 1996); and it is present at the northern edge of its range in Rankin County. Otherwise, it is found in Mississippi from Jones County{(Morgan 1979); (2 Aug 1978, es 1076, IBE)} in the Longleaf Pine Belt and Claiborne County (MNHP 1996) in the Loe uffs southward. Based on herbarium records, the species appears to be most com- mon in oe southern Loess Bluff Region. At the Rankin —— site, M. cordifolia is found in rich hardwoods on the natural levee of Purvis Creek hea foetida (L.) DC. Mississippi. Rankin Co.: ca 8 km SW of Johns, 7 Aug 1995, Morris 4406 BE, NGC). This species is present at the margin of longleaf pine woods ina 622 SIpA 17(3 wa seepage area — a gas pipeline. Prior to this collection, P. foetida was only documented in Mississippi from Clarke County [(17 Aug 1892, Boydstrum s.n., MISSA); (19 Jul 196 Jones 145061 © Jos MISS)} to Simpson County (15 Jul 1970, Jones 19253, MISS) and south to the erbesina ee Shinners. Mississippi. Rankin Co.: ca 2 km SE of Puckett, 9 Aug 1995, Morris 4423 UBE, NGC). Verbesina walteri occurs here at the edge of damp decidu- ous woods in the floodplain and on the natural levee of the Strong River. This composite appears to be restricted to the shaded margins of moist hardwood forests in the Longleaf Pine Belt in Mississippi from Smith (11 Oct 1968, Temple 10761, MISS) and Rankin to Claiborne (Lowe 1921) counties and south to Forrest County (Rogers 1977), except for one curious old record from Tishomingo County in the extreme northeastern corner of the state (Lowe 1921). CLUSIACEAE Bypencam galioides Lam. Mississippi. Rankin Co.: ca 6.5 km SE of Brandon, 24 Jul 1995, Morris 4340 (BE, NGC). This population is ina seepage area along a stream and constitutes the northernmost record of H. galioides in Mississippi. Prior to this collection, the species was documented from Lawrence {(24 Jun 1957, Ray 8283, MISSA); (6 Jul 1966, Jones 7, MISS)} to Wayne counties and farther south. Thus, this is a range exten- sion of about 75 km from the nearest locality. ypericum setosum L. Mississippi. Rankin Co.: ca 8 km SW of Johns, 28 Jul 1995, Morris 4303 (BE, NGC). This St. John’s-wort occurs at the margin of longleaf pine wooc — i 7) in a seepage area ea a gas pipeline. Previous records of H. sefoswm in aineaecrias are from Lawrence (31 Aug 1966, Jones 9982, MISS) to € Rankin County ete extends he species’ known range northwestward in the state by about 65 km. GENTIANACEAE Sabatia campanulata (L.) Torr. Mississippi. Rankin Co.: ca 8 km SW of Johns, 28 Jul Clarke counties and southward. Thus, the 1995, Morris 4370 (IBE, NGC). A local, but dense, population of this rose gentian is at the margin of longleaf pine woods in a seepage area along a gas pipeline. Representing a significant range extension in Mississippi, t a station is approximately 100 km northwest f o of the nearest populations in Lamar (27 Jun 1967, Jones 13616, MISS), Forrest (Rogers 1977), and Wayne (21 Jul 1955, Ray 5050, ak counties. ILLICIACEAE Hlicium floridanum Ellis. Mississippi. Rankin Co.: ca 2.5 km E of Puckett, 14 Aug 1995, Morris 4439 (IBE, NGC, SWSL); ca 10 km SE of Star, 25 Aug 1995, Morris 4482 (BE). The first population of star anise cited here is present ina ridge bottom forest in the floodplain of the Strong River. This ridge bottom is adjacent to several small swampy depressions, and I. fovidenum locall y dominates the understory in the transition area to the depressions. The latter population of star anise occurs in rich hardwoods and around low- a. areas in the undulating Dabbs Creek bottom and also locally dominates the under- ory. These Rankin County sites are che norchwesternmost stations recorded thus far for J. ren in Mississippi. Elsewhere in the state, it is known from Copiah (29 Apr 1956, Ray 5468, MISSA) to Lauderdale (22 Nov 1969, Clonts 464, IBE) counties and souchward. IRIDACEAE Iris cristata Ait. Mississippi. Rankin Co.: ca 3 km E of Star, 17 Aug 1995, Morris 4445 (BE). This collection documents the southernmost population of crested dwarf iris known in Mississippi. At this site, /. cristata is on the lower slope above a bay swamp in a deep ravine. The very slender rhizomes and rel: vavely broad leaf blades of this 1. cristata speci- Morris, Flora and ecology of the northern Longleaf Pine Belt 623 men helped avoid confusion with J. verna, which has much thicker rhizomes and usually more narrow leaf blades. In addition, I. verna is only known to occur south and east of Rankin County. The closest voucher specimens of J. cristata are from Carroll County in the Loess Bluff Region (16 Apr 1990, Barbour 7033, IBE),and Winston (24 Apr 86, Smith 1982, IBE) and Kemper (6 Apr 72, McDaniel 15863, IBE) counties in the North Central Plateau. Each of these collections is at least 150 km north or northeast of the Rankin County population. LAMIACEAE Hyptis alata (Raf.) Shinners. Mississippi. Rankin Co.: ca 10 km W of Puckett, 2 Au- gust 1995, Morris 4392 (IBE, NGC). Usually found in bogs, flatwoods, wet thickets and clearings (Godfrey & Wooten 1981), musky mint occurs here in a boggy depression at the edge of longleaf pine woods. This is as far north as H. alata has ever been recorded i aad Jones (1976) shows that H. a/ata is mostly distributed from Amite to oe ne counties and southward in Mississippi except for one population in Copiah County (4 Aug 1966, 1 Temple 3808, MISS), about 60 km southwest of the Rankin County site. There are three additional peripheral populations, one in Covington County about 50 km due south- east of the Rankin County station (30 Jun 1962, McDaniel 3265, IBE), and two in Jones County about 85 km distant (21 Aug 1974, McDaniel 19180, IBE; 1 Sep 1978, Morgan 1178, TBE). LAURACEAE ersea palustris (Raf.) Sarg. Mississippi. Rankin Co.: ca 5S km NNE of Johns, 25 Jul 1995, Morris 4343 (IBE); ca 9 km SW of Johns, 27 Jul 1995, Morris 4355 (BE, NGC). The first collection was made in a branch bay between ridges with dry longleaf pine woods, and this documentation of P. palustris is the most northern record of the species in Missis- sippi. At this locality, some individuals of swamp bay are at least 7 m in height. The second collection was made from a small 5 m tree in a well-developed bay swamp within — Southern Mixed Hardwood Forest. Prior to these records, the documented range of swamp bay in Mississippi was from Simpson (8 Mar 1986, ou art 1780, IBE) and Jones [(Morgan 1979); (28 Dec 1978, Morgan 1544, IBE)} counties southward to the Gulf Coast. LILIACEAE Melanthium virginicum L. Mississippi. Rankin Co.: ca 3.5 km E of Star, 7 Aug 1995, Morris 4409 (IBE, MMNS). Considered imperiled to rare in Mississippi (MNHP 1996), Virginia bunchflower is present at this site in a bay swamp at the base of steep, sandy ravines; of 46 plants in this population, 20 had undergone anthesis and were forming capsules. The species is known in Mississippi from the following physiographic regions: Tennessee River Hills (MacDonald 1996), North Central Plateau (Morris 1987, 1988; Winstead 1990), Longleaf Pine Belt (Carter & Jones 1968, Rogers 1977), and Coastal Pine Meadows (MNHP 1996). ONAGRACEAE Ludwigia linearis Walt. Mississippi. Rankin Co.: ca 8 km SW of Johns, 30 Jul 1995, Moe ee & Morris IBE). The population of this seedbox consists of scattered individuals a low, wet sage area at the margin of longleaf pine woods along a gas pipeline. es ae occurs here at the northwestern periphery of its range in Mississippt. Prior to this study, it was known from Simpson to Lauderdale counties and farther south in the Longleaf Pine Belt and Coastal Pine Meadows, with the exception of two records of the species in Monroe County in the Tennessee River Hills ones 1974; MacDonald 1996). 624 Sipa 17(3) ORCHIDACEAE Platanthera cristata (Michx.) Lindley. Mississippi. Rankin Co.: ca 10 km SE of Bran- don, 31 Jul 1995, Morris 4384 (IBE, MMNS). Approximately 105 individuals of this species co-occur with an equally large population of P. c//iaris in moist to wet flatwoods along a gas hata and adjacent to dry-mesic i ae scattered longleaf pines. Crested fringed orchid is listed as rare in Mississippi (MN 1996); favored habitats are pitcher plant bogs, bay swamps, relatively open os Peni along spring branches, and wet flatwoods primarily in the eastern and southern portions of the state excluding the Black Prairie and Jackson Prairie regic Triphora trianthophora (Swartz) Ryd b. Mississippi. Rankin Co.: ca 10 km SE of Star, 25 Aug 1995, Morris 4483 Mixed Hardwood Forest on a gentle, N-facing slope in the undulating Dabbs Creek bot- mwm). Three birds orchid is found at this site in Southern “a tom. It is an imperiled to rare species in Mississippi and has been documented only at widely scattered localities in the northern part of the state south to Smith and Clarke counties (MNHP 1996). Rankin is at the southwestern periphery of its range. POLYGALACEAE Polygala boykinii Nutt. Mississippi. Rankin Co.: ca 6 km SE of Brandon, 24 Jul 1995, Morris 4338 (BE, NGC, mwm). Growing along che border of upland pine- an hickory woods at this locality, P. boykznii appears to be restricted to the Longleaf Pine Belt, eastern Jackson Prairie, extreme southeastern North Central Plateau, and southern Black Prairie regions in Mississippi based on herbarium records at IBE, MISS, and MISSA. Thus, with the addition of this collection, the species ranges to ee and } a (Lowe 1921) coun- ties in the northwest and Noxubee (11 Jun 1966, Marler 23, MISS; 14 Jun 1966, Marler , MISS) and Lowndes (Lowe 1921) counties in che nae SCHISANDRACEAE Schisandra glabra (Bickn.) Rehd. Mississippi. Rankin Co.: ca 10 km SE of Star, 24 Aug 1995, Morris 4480 (IBE, MMNS). Scarlet woodbine is very local here, consisting of a population of 8 vines on a rich, well-drained terrace in the Dabbs Creek bottom. This woody vine is rare to uncommon in Mississippi MNHP 1996). Although known from the Longleaf Pine Belt (Rogers 1977) and the western North Central Plateau (Winstead 1990), S. glabra is most common in the Loess Bluffs (Morris 1987, 1988). THEACEAE Stewartia malacodendron L. Mississippi. Rankin Co.: ca 11 km SW of Florence, 26 Jul 1995, Morris 4348 (BE, MMNS, NGC, SWSL); ca 9 km SW of Johns, 27 Jul 1995, Morris 4353 (IBE, MMNS); ca 11 km ESE of Brandon, 5 Aug 1995, Morrs 4399 (IBE, MMNS). Silky camellia is a rare to uncommon shrub in Mississippi (MNHP 1996), anc these Rankin County populations mark the northernmost known occurrence of the species [on in Mississippi. Previously, $. malacodendron was documented from Copiah to Smith and Wayne counties and southward in the Longleaf Pine Belt (MNHP 1996). The first Rankin County station 1s an extensive system of rich ravines with many N-facing slopes above Bear Creek; a 0 tal of 48 stems, some of which are 5-6 m tall, in four colonies comprise the population of si Iky camellia here. The northernmost population is on the lower slope of a N-facing ravine on Shiloh Hill, the highest point in the county, and the population south- west of Johns is on a steep, N-facing slope above a bay swamp. ACKNOWLEDGMENTS Ken Gordon, Mississippi Natural Heritage Program, is gratefully ac- knowledged for providing information concerning the distribution and Morais, Flora and ecology of the northern Longleaf Pine Belt 625 status of some of the species discussed in this paper. Dr. Sidney McDaniel (IBE and MISSA) and Dr. M. B. Huneycutt (MISS) are thanked for their cooperation concerning the herbarium collections cited herein and Dr. McDaniel also for the verification of Iris cristata. Field work was supported by a Mississippi Wildlife Heritage Fund 1995 Research Grant. Field assis- tance by Suzette Morris is deeply appreciated, and thanks are due Cornelia A. O’Neal and Mary L. O’Neal for facilities used in the preparation of herbarium specimens. Publication cost was paid by a North Georgia Col- lege & State University Faculty Research Grant. REFERENCES Carraway, D.T. 1990. A floristic study of the Tallahala Wildlife Management Area of Bienville National Forest, Mississippi. Master's thesis, Department of Biological Sci- ences, Seep State University, Mississippi State. Carter, J.W. and S.B. aaa — 1968. The vascular flora of Johnson State Park, Missis- Sippi. ee 33219 LANG 1988 ee cee anatomy, and demography of the Needle Palm Riga Lum By strix), Dissertation, Department of Biological Sciences, Mississippi State University, Mississippi State CLeweLl, A-F. 1985. Guide to the vascular plants of the Florida panhandle. Florida State ee Press, Tallahassee. Cote, W.A., R.W. Situ, M.L. SpANN, and D.C. Stamps. 1987. Soil survey of Rankin County, Mississippi, United States Department of Agriculture, Soil Conservation Service 1n co- operation with Mississipp! ee Sapien and Forestry Experiment Station Goprrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida aia adjacent Georgia and Alabama. he Wekeag of Georgia Press, Athe GoprreEy, R.K., and J.W. Wooten. — . Aquatic and eae plants of southeastern United Saas ‘onli Vol. . University of Georgia Press, Athens Harper, R.M. 1914. A superficial ee of the pine-barren vegetation of Mississippi. Bull Torrey Bot. Club 41:551—567. HoimGren, P.K., N.H. HoitmGren, and L.C. Barnett (Eds.). 1990. Index herbariorum. Part I: the herbaria of the world, 8th ed. New York Botanical Garden, Bronx Jones, S.B., JR. 1974. ae flora. II. Distribution and identification of the Oneetee Castanea 39:370—379. JONES, 8.B., JR. eo eae Hora V. The mint family. Castanea 41:41—58. Lowe, E.N. 1921. Plants of Mississippi, a list of flowering plants and ferns. Mississippi State Geol. Surv. Ball 17. Hederman Bros., Jackson, MS. MaeDenins, | 1996. A survey of the flora of Monroe County, Mississippi. Master’s thesis, Department a Biological Sciences, Mississippi State University, Mississippi State Mitts, R. and S.B. Jones, Jr. 1969. The en a of a mesic southern mixed hardwood orest in souk Mississippi. Castanea 34:62— Mississipp! NATURAL HERITAGE PROGRAM. i ae plants & animals: by county. Mu- seum of Natural Science, Mississippi Department of Wildlife, Fisheries & Parks. Jack- n, Mississippi. Moraan, D.R. 1979. A floristic study of northeastern Jones County, Mississippi. Master's thesis, Department of Biological Sciences, Mississippi State University, Mississippi State. Morris, M.W. 1987. The vascular flora of Grenada County, Mississippi. Master’s thesis, Department of Biological Sciences, Mississippi State University, Mississippi State. 626 Sipa 17(3) sect M.W. 1988. Noteworthy vascular plants from Grenada County, Mississippi. Sida 3:177-186 eis MW, 1989. Spiranthes (Orchidaceae) in Mississippi. Selbyana 11:39—48. Morris, M.W., C.T. Bryson, and R.C. Warren. 1993. Rare vascular nian and associate cane communities yo the Sand Creek Chalk Bluffs, Oktibbeha County, Mississippi. Castanea 58:250—259. Raprorb, A.E., H.E. none and C.R. BELL. 1968. Manual of the vascular of the Carolinas. The Ulver iy of North Carolina Press, Chapel Hill. Rocers, K.E. 1977. Vascular flora of che Ragland Hills area, Forrest and Perry counties — erga Sida 7:51-79 RE, S., C. Frost, and P.D. Doerr. 1993. Southern mixed Soaehe forest: The form long af pine forest. In: Martin, W.H., $.G. Boyce, and A.C. Echternache, eds Biodiversity of the southeastern United States: Lowland Terrestrial eee John Wiley & Sons, Inc., New York. WINSTEAD, R. 1990. A taxonomic and ecological survey of the plant communities of Attala County, Mississippi. Master's thesis, Department of Biological Sciences, Mississippi State University, Mississippi State. DOCUMENTED CHROMOSOME NUMBERS 1997.1. CHROMOSOME NUMBERS IN COMPOSITAE FROM MOROCCO AND SPAIN JOHN L. STROTHER University Herbarium, 1001 Valley Life Sciences Building University of California Berkeley, CA 94720-2465, U.S.A. LINDA E. WATSON Department of Botany, 316 Biological Sciences Building Miami University Oxford, OH 450506, U.S.A. ABSTRACT 18 populations, which represent 14 species (1 represented Chromosome numbers from 4 tribes, of Compositae, confirm reports for the plants. by 2 varieties) from 11 genera, RESUMEN Los nimeros cromosémicos de 18 pobl iclones, que representan 14 t especies (1 con 2 variedades) de 11 géneros, 4 tribus, de Compositae, confirman lo registros para estas plantas. Although chemical methods increasingly dominate the current vogue, chromosome numbers continue to be important indicators of evolutionary (and taxonomic) relationships among plants (cf. Stebbins 1993). MATERIALS AND METHODS Source-plants reported on here were collected and identified by Watson. Voucher specimens have been deposited in the herbarium of Miami Uni- versity (MU). Chromosome counts were made by Strother from meiotic figures in acetocarmine-stained squashes (in Hoyer’s mountant) of sporo- phytic (pollen parent) cells from floral buds fixed in a solution of 3 parts 95% ethanol: | part acetic acid (see Radford et al. 1974; Sharma and Sharma 1980) RESULTS Chromosome counts reported here confirm previously reported numbers for the counted taxa (as cited in standard indices, e.g., Goldblatt and Johnson 1996 and references therein). Sipa 17(3): 627-629. 1997 628 Sipa 17(3) Anthemideae Aaronsohnia pubescens (Desf.) K. Bremer & Humphries. 27 = 9 I. Mo- rocco, S of Geurcif, along Rte. $329 at Fritissa, Watson 95-19A. Anacyclus clavatus (Desf.) Pers. 27 = 9 II. Morocco, ca. 8 km NW of Ifrane along Rte. $309, Watson 95-12C; Spain, SE of Seville at Villaluenga del Rosario, Watson 95-29C. Anacyclus radiatus Loisel. 2n = 9 II. Spain, NE of Seville, S of Lora del Rio at Guadalquiver River, Watson 95-4E; Spain, E of Cadiz at Medina- Sedonia, Watson 95-37A. Anthemis arvensis L. 2n = 9 Il. Spain, SE of Seville at Villaluenga del Rosario, Watson 95-29B. Chamaemelum mixtum (L.) All. 2n = 9 I. Morocco, along Rte. P24 be- tween Ifrane and Azrou, Watson 95-13G. Chamaemelum nobile (L.) All. 2n = 9 II. Morocco, between Ito and Azrou, Watson 95-14C; Spain, SE of Seville, at Villaluenga del Rosario, Watson 95- 30B Chamaemelum nobile var. discoidea (L.) All. 2n = 9 I. Morocco, along Rte. P24 between Ifrane and Azrou, Watson 95-13B. Chrysanthemum coronarium L. 2n = 18 (8 II + 21, chain of 3 + 7 ID + 1/7, etc.). Spain, SE of Seville at Montellano, Watson 95-8C Chrysanthemum segetum L. 2n = 9 I. Morocco, coe, Rte. P24 between Ifrane and Azrou, Watson 95-13A. Cladanthus arabicus (L.) Cass. 2n = 9 IL. Morocco, along Rte. P1 between Oujda and El-Aloun, Watson 95-20A. Glossopappus macrotus (Durieu) Brig. 2” = 9 II. Spain, S of Seville be- tween Arcos and El Bosque, Watson 95-26C. Otanthus mavitimus (L.) Hoffmanns. & Link. 2” = 9 II. Spain, W of Algeciras at Cabo de Trafalgar, Watson 95-30A. Calenduleae Calendula incana Willd. subsp. agarbiensis (Boiss.) Ohle. 2” = 16 11. Spain, W of Algeciras between Zahara de Los Atunes and Punta Comarinal, Watson 95-34B. — Inuleae Pallensis spinosa (L.) Cass. 2n = 5 Ul. Morocco, NE of Fez at J. Zalagh, Watson 95-10. Lactuceae Hypochaeris achyrophorus L. 2n = 6 Il. Morocco, along Rte. $309 ca. 15 km NW of Ifrane, Watson 95-11B. STROTHER AND WATSON, Chromosome numbers in Compositae 629 ACKNOWLEDGMENTS We gratefully acknowledge Marco Antonio Mateos for field assistance and Benito Valdes and Moh Rejdali for assistance with travel arrangements and collection permits. Research supported, in part, by National Science Foundation grants DEB-9408019 and DEB-9596274 to L.E.W. REFERENCES Go psBLatT, P. and D.E. JoHNson. 1996. Index to plant chromosome numbers 1992-1993. Monogr. Syst. Bot. Missouri Bot. Gard. 58:i-x + 1-276 Raprorp, A.E., W.C. Dickison, J.R. Massey, and C R. BELL. 1974. Vascular plant system- atics. New York: Harper & Row SHARMA, A.K. and A. ne 1980. Chromosome techniques: Theory and practice. Ed. - London: Butterworths. StepBins, G.L. 1993. Concepts of species and genera. In: Flora of North America Editorial Committee, eds. Flora of North America North of Mexico. London & New York: Ox- ford University Press. Vol. 1, pp. 229-246. NOTES CHAENORRHINUM MINUS (SCROPHULARIACEAE) NEW TO TEXAS Chaenorrhinum minus (L.) Lange {Linaria minor (L.) Desf.}, an erect glandu- lar-pubescent annual native to Europe (Fernandes 1972), has long been known as a widespread weed of disturbed and waste places in the eastern United States, particularly in the ballast along railroads (Robinson & Fernald 1908 (as Linaria); Fernald 1950; Gleason & Cronquist 1963; Radford et al. 1968; Strausbaugh & Core 1978). It has also been reported relatively far to the west in states such as Missouri (Steyermark 1963) and Oklahoma (Blair 1978; Taylor & Taylor 1994), and from eastern Kansas west to eastern North Dakota (Holmgren 1986). However, it was not included in the Texas flora by Correll and Johnston (1970), Hatch et al. (1990), or Jones et al. (1997). While working on the forthcoming Shinners & Mahler's Ulustrated Plants of North Central Texas (Diggs et al.), two Texas collections of Chaenorrhinum minus from just south of the Red River border with Oklahoma have be- come known. One (Taylor & Taylor 10570, previously at DUR, now at BRIT) was collected along a roadside in Fannin County in 1972. A more recent collection (Diggs 5748, BRIT) was obtained from railroad ballast in Grayson County in 1994. Because of the weedy nature of this taxon and available appropriate habitat, additional Texas records are expected. The species, which is unusual in having a spurred corolla, can be recog- nized by the following description: Erect, glandular-pubescent, often branched annual herb with stems to 40 cm tall; leaves opposite below to opposite or alternate above, the blades linear to oblong-lanceolate, 5-20 (—35) mm long, 1—3(—8) mm wide, entire; flowers in terminal, indistinct, bracteate racemes; pedicels conspicuous, 3-20 mm long in fruit; calyx lobes 2-5 mm long, subequal, linear to linear-spatulate, obtuse; corollas 4.5—9 mm long, pale lavender or lilac with yellow palate, with tube, 2-lipped limb, and a straight, cylindrical, 1-3 mm long spur; stamens 4; capsule 3— 6 mm long, porate; seeds numerous; flowering May—July.— George M. Diggs, Jv., Department of Biology, Austin College, Sherman, TX 75090, U.S.A. & Bo- tanical Research Institute of Texas, gdiggs@austine.edu; Constance E.S. Taylor, R. John Taylor, Southeastern Oklahoma State University, Durant, OK 74701, U.S.A, ctaylor@sosu.edu. REFERENCES BLair, A.P. 1978. New plant distribution records in eastern Oklahoma. Proc. Oklahoma Acad. Sci. 58:106-107. Sipa 17(3): 631. 1997 632 Sipa 17(3) Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Diccs, G.M. Jr., B.L. oe and R.J. OKENNON. Shinners & Mahler’s illustrated plants of North Central Texas. To be published in early 1998 by the Botanical Research Insti- tuce of Texas, Fore Worth. FERNALD, M.L. 1950. Gray’s manual of botany, 8th ed. Reprinted 1987. Dioscorides Press, Portland, Oregon. FERNANDES, R. 1972. Chaenorrhinum. In: T.G. Tutin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H. Valentine, $.M. Walters, and LD . Webb, eds. Flora Europaea. Cambridge Univ. Press, eek England. Vol. 3, pp. 224-226. GLEASON, H.A. and A. CRonguist. 1963. } eee “OF the vascular plants of northeastern United States “and adjacent Canada. Van Nostrand Reinhold Company, New York Harcu, S.L. Vee ae L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas A&M Ag ixp. Sta. Misc. Publ. 1655. College Stati HOLMGREN, N.H. 1986. 5 rophulaiacea In: Great Plains Flora eee Flora of the Great Plains. Univ. Press of Kansas, Lawrence. Pp. 751-797. Jones, $.D., J.K. Wiper, and P.M. eo 1997. Vascular plants of Texas: A com- nensive checklist including synonomy, bibliography, and index. Univ. of Texas Press, pre Austin. Raprorb, A.E., H.E. AHies, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The Univ. of North Carolina Press, Chapel Hill. ROBINSON, B.L. and M.L. i ERNALD. 1908. Gray's new manual of botany, 7th ed. American Book Company, New STEYERMARK, J.A. 1963. ae of Missouri. The lowa State Univ. Press, Ames. STRAUSBAUGH, P.D. and E.L. Corr. 1978. Flora of West Virginia, 2nd ed. Seneca Books, Inc., Granetsville, West Virginia. Tayior, R.J. and C.E.S. Taytor. 1994. An annotated sperms and flowering plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Univ., Durant. — ist of the ferns, fern allies, gymno- Sipa 17(3): 632. 1997 NOTES 633 HY POCHAERIS GLABRA (ASTERACEAE), A NEW RECORD FOR TEXAS Hypochaeris glabra L., a + scapose annual native to Europe (DeFilipps 1976), is known in the southeastern United States as far west as Louisiana and Arkansas from disturbed and waste places, especially in sandy soils (Radford et al. 1968; Cronquist 1980; Gandhi & Thomas 1989). However, it was not included in the Texas flora by Correll & Johnston (1970), Hatch et al. (1990), or Jones et al. (1997), nor is it known from Oklahoma (Taylor & Taylor 1994). While working on the forthcoming Shinners & Mahler's Ilustrated Plants of North Central Texas (Diggs et al.), a collection of H. glabra from a fre- quently mowed hay field in Williamson Co. in the southern part of North Central Texas (T.J. Watson 1620, 1993, BRIT), identified as H. microcephala var. albiflora, came to our attention. Guy Nesom kindly confirmed our identification of the specimen as H. glabra. Shinners (1966) first documented the occurrence of the genus Hypochaerts in Texas following his discovery of the South American H. microcephala (Sch.Bip.) Cabrera var. albiflora (Kuntze) Cabrera in southeast Texas. Sub- sequently, two additional species, the Eurasian H. radicata L. and the South American H. brasiliensis (Less.) Griseb., have been added to the state’s flora (Tomb 1974). Keys separating the four Hypochaeris species in Texas and the southeastern U.S. can be found in Shinners (1966), Cronquist (1980), and Gandhi & Thomas (1989). Hypochaeris glabra can be recognized as follows: Annual 10-60 cm tall, usually glabrous or nearly so; stems leafless or only sparsely and minutely bracteate, unbranched or branched; basal rosette leaves numerous, oblan- ceolate, toothed or pinnatifid; heads 5—15 mm wide; corollas white to yel- lowish, 1(—1.5) times as long as involucre, only ca. 2 times as long as broad, pappus of an outer series of short, merely barbellate bristles and an inner series of much longer, plumose bristles; outermost achenes usually beakless, the other achenes usually with a well-developed beak. Flowering Feb-? (in southeastern U.S. flowering mostly in spring, continuing until fall, or al- most all year in protected areas) (Shinners 1966; DeFilipps 1976; Cronquist 1980; Gandhi & Thomas 1989). The genus name has sometimes been spelled Hypochoeris (e.g., Shinners 1966; Tomb 1974; DeFilipps 1976), probably based on the spelling by Linnaeus (1754) in Genera Plantarum. However, the accepted spelling is Hypochaerts (Greuter 1993), based on Linnaeus’ (1753) original spelling in Species Plantarum.—George M. Diggs, Jv., Department of Biology, Austin Col- SIDA 17(3): 633. 1997 634 Sipa 17@G lege, Sherman, TX 75090, U.S.A. & Botanical Research Institute of Texas, gdiges@austinc.edu; Robert J. O'’Kennon, bokennon@brit.org, and Barney L. Lipscomb, barney@ brit.org, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102, U.S.A REFERENCES Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Cronaquist, A. 1980. Asteraceae. In: Maer ar tae of the southeastern United States. The Univ. of North Carolina Press, Chapel H Diccs, G.M. Jr., B.L. Lipscomp, and R.J. O° a NNON. Shinners & Mahler's illustrated plants of North Central Texas. To be published in early 1998 by the Botanical Research Insti- e of Texas, Fore Worth. DE Fi Lipps, R.A. 1976. Hypochoeris. In: Tutin, T.G., V.H. Heywood, N.A. Burges, D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Flora Europaea. Cambridge Univ. Press, Cambridge, England. GANDItL, K.N. and R.D. Tac 1989. Asteraceae of Louisiana. Sida, Bor. Misc. 4:1—202. Harcnu, S.L., K. - ri and L.E. Brown. 1990. Checklist of the vascular ee of Tex esas A&M Agric. Exp. Sta. Misc. Publ. 1655. College Station. tome ‘S D., J.K. Wiper, and P.M. Montrcomery. 1997. Vascular plants of Texas: A com- prehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press, ustin LINNAEUS, C. 1753. Species Plantarum.... 2 vols. Stockholm. —.«d'754. Genera Plantarum, ed. 5. Stockholm. Rabrorb, A.E., H.E. Anes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The Univ. of North Carolina Press, Chapel Hill. SHINNE RS, L.H. 1966. Hypochoeris microcephala var. ee (Compositae) in southeastern € New to North America. Sida 2:393—3¢ Taytor, RJ. and C.E.S. Taytor. 1994, An annotated list of the ferns, fern allies, gymno- sperms ee Gomes plants of Oklahoma, 3rd ed. Southeastern Oklahoma State Uni- versity, Durant. Toms, A.S. 1974. Hypochoeris in Texas. Sida 5:287—289. WERNER, G., R.K. BRUMMITT, E. Farr, N. Kirtan, P.M. Kirk, and P.C. Siva. 1993, Names in current use for extant plant genera. Regnum Veg. Vol. 29. Sipa 17(3): 634. 1997 Nores 635 LECTOTYPIFICATION OF CLITORIA AUSTRALIS (FABACEAE) Bentham (1864) published an Australian flora and in it, newly described was Clitoria australis Benth. Currently, the legume treatment of a modern Flora of Australia is nearly completed. The objective of this note is to meet a request to provide documentation on the lectotypification of Bentham’s species name. Clitoria australis is an Northern Australian endemic subshrub or suffrutescent herb with both 3- and 1|-foliate leaves, small (2.5—3 cm) white flowers, a short, silky pubescent calyx tube (6-8 mm long) with lobes (9— 11 mm long) conspicuously longer than the tube, and fruits constricted between the seeds. Fantz (1979) assigned this species to C/:toria section Tanystyloba Fantz. Other species in this section include C. cordiformis Fantz (Thailand), C. hanceana Hemsl. (China, Kampuchea, Thailand, Vietnam), C. javanica Mig. (Thailand), C. /inearis Gagnep. (Laos), and C. macrophylla Wall. ex Benth. (Myanmar, Thailand, Vietnam). Bentham (1864) cited one collection of C. australis saying “N. Australia. Arnhem S. Bay, R. Brown (Herb. R. Brown). I have been unable to adopt Brown’s MS name {= C. a/ba} which is now preoccupied in the genus [= C. alba Don, 1832}.” have examined five specimens (E:2 sheets, F:herb. Small, K, NY) of this species collected by R. Brown from Arnheim Bay. They are regarded as original material. Stafleu and Cowan (1979) reported that Robert Brown's Herbarium came into posession of J.J. Bennett in 1858, and was later split upon the latter's death in 1876, with collections going to BM, K and E. Bentham examined these specimens after his revision (Bentham 1858) of C/itoria that lacked C. australis, but prior to his publication in 1864. Presumably, the speci- mens were examined when they were part of Bennett’s Herbarium. No specimen has been located at BM. Two specimens (E-108', K) bear a printed label citing “R. Brown, Iter Australiense, 1802-1805. [Presented by direction of J.J. Bennett, 1876].” Label data included “Arnheim S. Bay,” a collection number of 4237, and “Clitovia alba” R. Br. (Robert Brown's name) with “C. australis, Benth.” written on the label below Brown’s name. The remaining three specimens (E-109, E, NY) lacked the Bennett label, Brown’s species name and collec- tion number, but were distributed with Bentham’s species name. The E- 109 specimen had the same locality and date, whereas the F specimen bore 'Specimens from E lacked accession numbers. Cited are the accession numbers found be- low 60/74 {loan number} in the center of an inked circular stamp with “Royal Botanical Gardens, Edinburgh” around the margin. Sipa 1703): 635. 1997 636 Sipa 17(3) only “Australia, 1802-5” and the NY specimen bore only “Australia, Arnheim S. Bay.” In selecting a lectotype, the F and NY were excluded immediately from consideration, as they lacked the Bennett label, Brown’s manuscript name C. alba, and complete label data. In addition, these two specimens appear somewhat distinct in appearance to me from the other three specimens, in the age of the flowers, color of vegetative parts, and in overall stature. It seems probable to me that this pair of specimens were collected on ee ent dates, are not true duplicate collections of the specimens (Brown 4237) deposited at E and K, and possibly not even examined by Bentham. f the remaining three specimens, the K specimen was designated as the lectotype. It is a good representative specimen of the species, agrees closely with the diagnosis by Bentham, bears Brown's name and the spe- cific locality name discussed in the protologue, and was annotated by Bentham. Bentham (1864) did not cite Brown's collection number, but Bentham (1858) has used this practice previously when he described other species of Clitoria (e.g. C. flagellaris, C. nana). Location of these types re- vealed a collector's number not cited by Bentham in the protologue. The F and NY specimens are probable original material, but are not regarded by me as duplicates of the lectotype collection; therefore, they were annotated as lectoparatypes instead of isolectotypes. In conclusion, the citation for this species is as follows: Clitoria australis Benth., Fl. Austral. 2:242. 1864. [Clitoria alba R. Brown, nom. in sched. ,non Don (183 2)}. Type: AUSTRALIA. Norruern ees Arnheim S. Bay, 1802-1805, R. Brown 4237 (ecroryrE, designated h -J.J. Bennett herb.; lsoLtecrorypes: E-108! & 109). AUSTRALIA, lic., R. Brown s.n. ae TOPARATYPES: F-Hb. Small, NY —Paul R. Fantz, Dept. of Horticultural Science, Box 7609, North Carolina State University, Raleigh, NC 27695-7609, U.S./ REFERENCES BENTHAM, G. 1858. Synopsis of the genus C//roria. J. Linn. Soc., Bot. 2:33—44. Bentuam, G. 1864. Flora Australiensis. Lovell Reeve & Co., London. Vol. 2, p. 242. Fantz, P. 1979. A new section of Clitoria subgenus Newrocarpum (Legumit and a new species endemic to Thailand. Brittonia S3L:LLS-118. STAFLEU, FL and R. Cowan. 1976. Taxonomic literature. Bohn, Scheltema & Holkema, Utrecht. Vol. 1, p. 364. ~ — Sipa 17(3): 636. 1997 Notes 637 NAMING A SOUTHWESTERN MALANIS (ORCHIDACEAE) Recently, Coleman (1997) brought to my attention the fact that Watson (1883) had described a specimen collected in Arizona by the Lemmons, naming it Microstylis purpurea. Ridley (1888) revised Microstylis and Malaxis. Since Lindley (1849) had used the specific epithet purpurea to describe a Microstylis from Ceylon and Java, Ridley renamed the taxon Microstylis porphyrea. In this revision, he rejected the idea that the taxon was the same as M. ebrenbergit. Ridley’s comment was “M. porphyrea has no distinct fovea; the lip is concave at the base but not saccate.” This latter refers to Reichenbach’s (1849) description of the lip of M. ebrenbergii as “gibbere acuto in medio parte basilari,” i.e. acutely gibbous in the middle basal part. Kuntze (1891) placed all Microstylis species under Malaxis, so the taxon became Malaxis porphyrea (Ridl.) Kuntze. In their consideration of Malaxis nomenclature, Ames and Schweinfurth (1935) decided that M. porphyrea was synonymous with M. ebrenbergii, and subsequent authors have followed their lead (e.g., Correll 1950; Luer 1975 et al.). I have examined the holotype of M. porphyrea from the Gray Herbarium. It has papillate floral segments and narrowly sagittate lip which easily dif- ferentiate it from M. ehrenbergii with glabrous floral segments and a broadly triangular-hastate lip. The species should be listed as follows: Malaxis elec eee Kuntze., Rev. Gen. Pl. 2:673. 1891. Microstylis porphyrea Ridl., J. Linn. Soc., Bot. 24:320. 1888. Macrostylis purpurea 8. Watson, Proc. Amer. Acad. ae 18:195. 1883, non Microstylis purpurea Lindl. Type: U.S.A. ARIZONA. Cochise Co.: Tanner's Canyon, Hyachuca Mts., Jul 1882, /.G. &5P Lemmon 2881 (HOLOTYPE: G There is a difference that appears in Ridley’s and Reichenbach’s descrip- tions that has not been considered in previous justifications for separation of the two species. Ridley states the stem to be “superne laxe racemosus,” i.e., loosely racemose above. Reichenbach, in contrast, says, “racemus plurimiflorus,” i.e., raceme very many flowered. This difference is readily seen in Figures 1 and 2, where each dorsal sepal is about 2 mm long. The ranges of the two do not overlap. The southernmost sites of M. porphyrea are probably in northern durango and Sonora. The northernmost site of M. ehrenbergii is in southern Hidalgo. The gap between is about 1200 km ACKNOWLEDGMENTS Iam grateful to the Curator of the Gray Herbarium for the expeditious loan of a holotype specimen and thank all the friends and colleagues who have provided comments and suggestions. Sipa 17(3): 637. 1997 638 Stipa 173) Fic. 1. Malaxis porphyrea. New Mexico. Fig. 2. Malaxis ehrenbergii, Oaxaca, Mexico. Photo by T.K. Todsen. Photo by E.W. Greenwood. —Thomas K. Todsen, Department of Biology, New Mexico State University, Las Cruces, NM 88003, U.S.A. REFERENCES Ames, O. and C. ScuweinrurrH. 1935, Nomenclatorial studies in Malaxis and Spiranthes. s:116. CoLeMAN, R. 1997. Personal communication. Correwt, D. 1950. Native orchids of Norch America. Chronica Botanica. Kunrze, O. 1891. Revis. Gen. Pl. 2:67. LinpLEy, J. 1849. Microstylis purpurea sp. nov. Gen. et Sp. Orch. 20. Lure, C. 1975. The native orchids of the United States and Canada. New York Botanical Garden. REICHENBACH, H. 1849. Microstylis ehrenbergi7 sp. nov. Linnaea 22:835. Riptey, H.N. 1888. A revision of the genera Mecrosty/is and Ma/axis. J. Linn. Soc. Bot. 24:320. Watson, S. 1883. Description of some new western species. Proc. Amer. Acad. Arts 18:195. SIDA 17(3): 638. 1997 BOOKS NOTICES SmitrH, Bruce D. 1995. The Emergence of Agriculture. (ISBN 0-7167- 5055-4, hbk.). W.H. Freeman and Company, 4419 West 1980 South, Salt Lake City, UT 84104, U.S.A. (801) 973-4660, (801) 977-9712 FAX. $32.95. 230 pp, numerous b&w, color photos, and maps. From the cover: “In The chi sleuth well-known archaeologist Bruce Smith explores the initial emergence and early expansion of agriculture, and the transformations in human society that it a possible. He haeologists have come to recognize that agri- culture didn’t just emerge in one or two places, from whence it spread to others, but was independently invented several times over, in widely separated parts of the world. In his book, Bruce Smith charts the course of the agricultural revolution as it occurred in the Middle East, Europe, China, Africa, and the Americas, showing, too, how basic archaeo- logical methods and modern technologies such as plant analysis, radiocarbon dating, and ampling are used to investigate this pivotal event. Although in the popular mind the agricultural revolution is often seen as a one-step transition from hunter-gatherer soci- eties to farming ones, Smith shows how truly varied were the patterns of animal and plant domestication in different parts of the world.” Contents include: Preface; In Search of Origins; Creating New Plants and heal New Technology and the Seach for Agricul- tural Origins; The Fertile Crescent; Europe and Africa; East Asia; Middle and South America; Eastern North America and the Southwest; Epilogue: The Search for Explanations; Pur- ther Readings; Sources of Illustrations; and Index. VENTOCILLA, JORGE, Heractio Herrera, and VALERIO Nunez (Edited by Hans Roeder, translated by Elisabeth King). 1995. Plants & Ani- mals in the Life of the Kuna. (ISBN 0-292-78725-1, pbk; 0-292- 78726-x, hbk.). University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, U.S.A. 512-471-4032. $12.95 (pbk.), $25.00 (hbk.). 160 pp, 5 b&w photos, 33 line drawings, 3 maps. The earth is the mother of all things...’ begins this original and accessible book on how the Kuna Indians of Panama relate to the natural world. An integrative project in- volving Kuna traditional leaders and trained scholars, and fully illustratec by a Kuna artist, this translation of Plantas y animales en la vida del pueblo Kuna focuses on Kuna plant and animal life, social life, and social change as a means of saving traditional ecological knowledge and ‘returning’ ic to the community.” Contents include: Foreward by James Howe; Acknowledgments; Baba’s Creation; Ready to Change; The Kuna; Rivers and Moun- tains; Terrestrial Fauna; Hunting in Gangandi; Money Creates Hunger; Submarine “De- forestation;” The Spirit of the Uaga; Medicinal Plants; The Uerwk Palm; To Be or Not to Be; Epilogue; Appendix A. (Scientific and Vernacular Names of Plants and peer ); Ap- pendix B. (Kuna Communities); Glossary; Bibliography; Authors; and Index. The book is nicely illustrated. Stpa 17(3): 639. 1997 640 Stipa 17(3) Ren, S., Eprror. 1994. Multilingual Dictionary of Agronomic Plants. (SBN 0-7923-2970-8, hbk.). Kluwer Academic Publishers, 101 Philip Drive, Norwell, MA 02061, U.S.A. $120.00. 286 pp. From the Preface and Introduction: “This ig nai contains the names of nearly 2500 plant species, in some cases including subspecies and varieties. Tropical crops from all parts of the world receive the same attention as those ee the temperate zone. Common names are given not only in English, French, German, Portuguese and Spanish but also in many other languages, where such names are used in the technical literature. ‘Agronomic plants’ comprise not only the agricultural and horticultural crops but also pasture plants, green manure, soil covers, trees used in agroforestry, and major weeds. Purely ornamental plants have been excluded. Also included are plants which are pres- ently being discussed as new crops, with considerable economic value. The Dictionary includes five languages: English, French, German, Portuguese, and Span- ish. The a eee wil a meet the needs not only of scientists in agriculture, botany and geography but also those of agricultural extensionists, merchants in agricultural products and professional translators.” Hunter, MALcoL L. Jr. 1995. Fundamentals of Conservation Biology. (ISBN 0-86542-371-1, hbk.). Blackwell Science, 238 Main Street, Cambridge, MA 02142, U.S.A. (617) 876-7000, (617) 876-7022 Fax. $42.95. 482 pp, numerous b&w illustrations. “The book focuses on what can be done to maintain biodiversity through management of ecosystems and populations. Starting with a succinct look at what conser- From the cover: vation and ene really mean and progressing to more complex topics such as mass extinctions, | rabitat degradation, and over-exploitation, Hunter creates a context in which the principles of conservation biology can be readily understood. Discussions of the social, political, and economic aspects of conservation biology issues are interwoven throughout the text and addressed independently in their own chapters. With a careful eye towards providing coverage that is balanced geographically, taxonomically, and environmentally, Hunter provides a super ative book that sets the standard in this increasingly ee subject.” Contents include: Preface; Conservation and Conservation Biology; Wha Biodiversity; Species Diversity; Ecosystem Diversity; Genetic Diversity; Mass ae and Global Change; Extinction Processes; Habitat Degradation and Loss; Over exploita- tion, Exotic Species; Managing Ecosystems; Managing Populations; Zoos and Gardens; Setting Priorities; Social Factors; Economics; Politics and Action; Epilogue; Literature Cited and Author Index; Species Index; and Subject Index SipA 17(3): 640. 1997 BOOK REVIEWS Don E. WILSON AND ABELARDO SANDOVAL (eds) MANU. 1997 The Biodiversity of Southeastern Peru; La Biodiversidad del Sureste del Pert. ISBN 1-56098-710-3, pbk). Smithsonian Institution Press, 470 LEnfant Plaza, Washington, DC 20560. $35. 679 pp. Numer- ous figures, maps. and b/w plates. This volume of 26 articles by 93 authors and collaborators represents an up-to-date summary of the inventory protocols. flora, and fauna present in the Manu MAB Biosphere Reserve. Located in the Departments of Cusco and Madre de Dios, the area comprises over 1.5 million hectares tropical forest, and has been the site of intensive field biological re- search since 1987. The book i is divided into three sections: General floristics, flora and fauna. The general floristics not only provides a detailed summary of what is known regarding the Baa but a detailed explanation of the sampling protocol employed. The floristic section cludes two articles dealing ith the mapping and inventory protocols used by the BIOLAT and Smithsonian/MAB office of Biodiversity Programs. These are articles written in an mate clear style and will be standard references for a long time to come. The Flora ection contains seven articles, ranging from fee dealing with forest dynamics, family accounts (Chavez, Londofio, Herrera), and finally a report on the bryoflora (Gradstein). All are first-class treatments, written in a Ke agora, easily usable style, a are exhaustive in their coverage. Finally, the Fauna section contains two subsections, inverte- brates (8 articles) and vertebrates (12 ae The aati ee section is dominated by entomological and arachnological studies, pointing to the need for studies of other inver- tebrates, such as annelids, and mollusks. The vertebrate section contains icthyological, herpetological, ner ee and mammalogical studies, with one study of a dipteran. rs of the eee subsection, the Silva & Coddington paper on the spiders, and the Mor ales & McDiarmid paper on amphibians, are particularly notable be- cause of their statistical rigor, relevance to the floristics section, and their foci on compara- tive analyses with other areas of Peru. The paper on genetic ae of amphibians, by Cérdova et al. is particularly timely and provides baseline data critical for monographers to consider because karyotipic oo within one general area is discussed. In summary, this volume is a timely and clearly written summary of our knowledge of the biological peas present in cs mammoth Biosphere. It will not only provide baseline data for the taxonomic groups treated, but will provide fodder for many a grant an and the ere so clearly explained, make ic an excellent choice for a biodiversity m suring and monitoring course. The editors are to be congratulated on still another foe product, and the two program directors involved, Dr. Wilson of the Smithsonian Biodiversity Programs office, and Dr. Dallmeier of Smithsonian/MAB, are to be com- nded for an outstanding example ad term collaboration. At $35 foe cloth, and less . ae the book is more than a bar —John J. ae, bt Botanical Research Institute of as, 509 Pecan Street, Fort Worth, TX 76102. 4060, U Sipa 17(3): 641. 1997 642 Sipa 173) KELLER, ROLAND. 1996. Identification of Tropical Woody Plants in the Absence of Flowers and Fruits (ISBNO-8176-5184, pbk). Birkhauser-Verlag, P. O. Box 133, CH-4010 Basel, Switzerland. $69.95. This book is the result of careful field observations by Keller in Venezuela, Madagascar, and Indonesia and to a lesser extent, in French Guiana, Guadeloupe, and Cameroon. Its principal purpose is to facilitate identification of sterile or juvenile woody plant specimens (living or, with greater difficulty, herbarium ww to the familial or hopefully, generic leve I have tested this book with taxa from Colombia, Ecuador, Peru, the Guianas, Philip- pines and Papua New Guinea, and can unequivocally state that it is, indeed, a handy eference work. I was able to find only one glaring omission, that of the lianous genus Embelia (Paleotrop.) in the Myrsinaceae. It should key out opposite the Annonaceae entry, in Key T, page 71, with a couplet as follows: “Bark not aromatic. Plant not twining, starting out as a scandent shrub, later maturity into ae without hooks.” Apart from confusing portions of Key X (couplet 13 in particular), the 26 keys are clear and easy to follow. The specialized Keys, composed of X, Y and Z are particularly helpful and desper- ately needed. The selection of families is wide and reasonable. | would have added the Clethraceae and Gesneriaceae, because of their great importance in such regions as the Western Cordillera of Colombia and adjacent Ecuador, the Amazonia/Hylaea interface of Ecuador and Peru, and Eastern Malesia (especially New Guinea). The Glossary, Notes and Illustrations section comprises 59% of the book’s bulk. Section , geographical distribution”, is essentially a list of important woody families from each area conforming to the author's categories. Sections 2-20 comprise an illustrated glossary, which is very good. I would have rephrased Section 6 because my understanding of sympo- dial branching is that it occurs by either substitution or apposition. Whether the resultant branches are plagio- or orthotropic does not affect the process of sympodial branching, but describes the final resultant branches orientation. Thus, e and f on page 6 should have the word “sympodial” added in front of the terms apposition and substitution, respectively. Perhaps that can be further clarified in the next edition of the book. In Section 7, I see little difference between continuous and diffuse ramification. I suggest that any perceived difference between the two is strictly spatial and not morphogenetic. I think the same thought process applies to the Location of the ramification on the unit of extension, which I have found to be quite variable in highly dynamic forests (with many windstorms, etc.). Also, it would have been helpful to remind the reader who is not fluent in architectural ire about differences between monoaxial and polyaxial systems in monocaulous runks. This is often a point of confusion, and it bears repeating in Section 7. From there, in aie 6, I would have preferred to seen a line or two about ptyxis vs. vernation in vegetative organs, but the examples here from Cullen are quite good. For Venation, Sec- tion 17, | would have preferred more close adherence to the standards proposed by Hickey in the Anatomy of the Dicotyledons, 2nd ed., Vol I. The same comment would hold true for the chapter on trichomes of the same volume, here included as Section 20. ” art Il, “Principal families of tropical woody Dictyledons...” is extremely useful for many families, but needs the following corrections: leaf with linear lyso-schizogenous canals (punctate-lineations), rather than leaf with resiniferous ducts, Stipa 17(3): 642. 1997 Book REVIEW 643 : leaf with hydropotes , instead of underside of blade with glandular trichomes in- ae at the bottom of small cavities The author is to be commended for successful completion of a Herculean task. Doubt- less, there will be groups other than those mentioned in the forward that have slipped by him, but on the whole, there is no other resource like this book. While I have pointed out but a few suggestions, I highly encourage all tropical ecologists, dendrologists, agrono- mists and systematists to buy this book and put it to the test in their area of geographic dare If the community can respond constructively and the minor corrections made, s field guide will be a truly unique standard for everyone’s use “The paper is high quality, as well as the binding. While my copy is already dog-eared, the spine has survived quite well. Its price is somewhat prohibitive, but its usefulness makes it a bargain, given its utility. I unhesitatingly recommend it for everyone.—John Pipoly I, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102- 4000, U.S.A Sipa 17(3): 643. 1997 Documented chromosome numbers 1997.1. Chromosome numbers in Con Morocco and Sp JobiL. Strother ae Linda EB. Watson 7 62 Notes Chaenorrhinum minus (Scrophulariaceae) new to Texas George M. Diggs, Jr, Constance E.S.Taylor, and R. fobn Taylor ) O31 Hypochaeris glabra (Asteraceae), a new record for Texas George M. Diggs, Jr, Robert J. O'Rennon, and Barney Lipscomb 033 Lectotypification of Clitoria australis (Pabaccac) Paul Rk. Faitz ) O35 Naming a southwestern Malaxis (Orchidaceae) Thomas kK. Todsen 03 Book reviews and notices 550, 568, 590, 598, 639-043 Vr N ISSN 0036-1488 IpOsikk ie from Sipa 17(4) INTRODUCTION The grass genus Erionenron was described by Nash in Small (1903) based on Erzoneuron pilosum (Buckl.) Nash. Species of this genus have been placed in several genera that are discussed in Valdés-Reyna (1985) and Peterson et al. (199?). Erionewron and Dasyochloa species were submerged in Tridens for many years (Parodi 1934; Hitchcock 1913, 1951; Gould 1951). Tateoka (1961), ina study of Tridens presented cytological, morphological, and ana- tomical evidence to support recognition of two distinct genera. Based on these results Tateoka recognized the genus Erioneron with five species, and suggested affinities with the genus Munroa, rather than with Tridens. That classification was accepted by agrostologists and used in several grass floras (Correll & Johnston 1970; Mico: 1973; Gould 1968, 1975, 1979: Anton 1977; McVaugh 1983; Zuloaga et al. 1994). The genus Dasyochloa Willd. first eared in Steudel (1840) as a syn- onym of Uralepis. Two species of Dasyochloa were listed, D. avenacea Willd. and D. pulchella Willd., both as “nomina nuda.” Although Rydberg (1906) validated Dasyochloa with an English description as part of a key, it has been regarded as a synonym of Erionewron. Caro (1981) transfered the spe- cies of the genus Erioneuron (excluding FE. pilosum) to Dasyochloa, based on the lemma morphology. Sanchez (1983) studied Dasyochloa reporting that anatomical information formed the most important differences between Ertonenron and Dasyochloa. She accepted Dasyochloa as a oo North American genus, consisting of D. pilchella Willd. ex Rydberg Anatomical characters of both leaf epidermis and leaf ae cross section were reported by Valdés-Reyna and Hatch (1995) and support the conclu- sions of this study. Dasyochloa has U-shaped leaf blades and sclerenchyma that are well-developed in association with the vascular bundles. Erionenron leaf blades are V-shaped and the sclerenchyma strands are absent or few in association with the vascular bundles. Enzyme electrophoresis can help delimit the occurrence of what appeared to be intermediates within the E. avenaceum-pilosum complex. Suspected natural hybridization between these species has raised the question con- cerning relationships. In addition, electrophoretic information may be important for the acceptance or rejection of the monotypic North Ameri- can genus Dasyochloa (Caro 1981; Sanchez 1983). MATERIAL AND METHODS Plant materials were obtained from field locations in the southern United States and northeastern México from 1983 to 1993. Plants were also grown from seed in the greenhouse. Samples of 10 plants per site were collected from 60 naturally occurring populations. Field notes were taken for each population sample concerning location, growth habit, general appearance, A REVISION OF ERIONEURON AND DASYOCHLOA (POACEAE: ERAGROSTIDEAE) JESUS VALDES-REYNA Departamento de Botdnica Universidad Autonoma Agraria “Antonio Narro” Buenavista, Saltillo, Coahuila 25315, MEXICO STEPHAN L.HaAlrCH S.M. Tracy Herbarium Department of Rangeland Ecology and Management Texas AGM University College Station, TX 77843-2126, U.S.A. ABSTRACT A taxonomic treatment of the New World genera, Erionenron and Dasyochloa, is pre- sented. Materials for morphological, electrophoretic and anatomical studies were collected from naturally occurring populations, and from herbarium specimens throughout the range of the species. A morphological analysis of specimens from 60 populations of Erzonenron and Dasyochloa is given. Phenetic and basic biostatistical techniques were used to delineate and define taxa. Based on these results four species were defined. An electrophoretic survey was conducted on seedlings representing 33 natural populations. Individuals were assayed for five enzyme systems: PGI, GOT, IDH, PGM, and MDH. The results show that varia- tion at the molecular level is concordant with the morphological characters for the separa- tion of taxa. Based on these studies and anatomical data three species of Erionenron are recognized and a lectotype is designated for FE. pi/osam. This research supports the recogni- tion of Dasyochloa (D. pilchella) as a monotypic North American genus. RESUMEN Se presenta un estudio 5mico de los géneros del Nuevo Mundo Ervonenron y Dasyochloa. o Los materiales para los estudios morfolégicos, electroforéticos y anatémicos Riecon tomados de ate naturales y de ejemplares de herbario en todo el rango de distribucion de las especies de estos dos géneros. Fue realizado un andlisis morfol6gico de muestras de 60 cree Fueron usadas técnicas fenéticas y bioestadisticas basicas para delimitar y definir los taxa. Basados en los resultados fueron dletintdes cuatro grupos. Pue realizado un muestreo electroforético en plantulas que oe 33 poblaciones naturales. En los individuos fueron analizados cinco sistemas enzimaticos: PGI, G ue IDH, PGM, y » MDH. con los caracteres + Los resultados muestran que la variacion al a molecular mortfol6gicos para la separacién de los taxa. Basados en estos estudios se reconocen tres especies de Evionenron, se designa el lectotipo de E. pilosum. Esta investigacion apoya el reconocimiento de Dasyochloa (D. pitchella) ¢ un género monotipico para Norteamérica Stipa 17(4): 645-666. 1997 CONTRIBUTIONS TO BOTANY FOUNDED BY LLOYD H. SHINNERS 1962 Sf AS Wm. F. Mahler Publisher 1971-1992 © @ Barney L. Lipscomb Botanical Research Institute of Texas, Inc. 509 Pecan Street Fort Worth, Texas 76102-4060, USA 817 332-4441 / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/siday John W. Thieret Prof. Dr. Félix Llamas Associate Editor Contributing Spanish Editor Biological Sciences Dept. Dpto. de Botanica, Facultad de Biologia Northern Kentucky University Universidad de Leén Highland Heights, Kentucky 41076, USA E-24071 Leon, Spain Guide ad for contributors a are available aie request and on the inside back cover of the last issue of each volume. subscription per year: $25. Individual, $40. USA nes $50. Outside USA; numbers issued twice a 4 fa) a © SIDA, CONTRIBUTIONS TO BOTANY, Volume 17, Number 4, pages 645-873. Copyright 1997 Botanical Research Institute of Texas, Inc rinted in the United States of America ISSN 0036-1488 CONTENTS A revision of Erioneuron and Dasyochloa (Poaceae: Eragrostideac) Jestis Valdes-Reyna and Stephan L. Hatch O45 Geographic spatial autocorrelation of morp hakuunensis (Liliaceae) jological characters in Hemerocallis Soon Suk Kang, Ki Bae Park, and Myong Gi Chung 607 From. Yunnan, China Fan Guo-Shene and Deng Li-Lan 677 New species of Rhamnaceae Leaf-print analyses: An ecologically friendly methodology for plant identification RJ. Ferry Sr., R. Foroughbakhch, L.A, Hauad, S. Contreras, J.V. Star, M.H. Badii, and H. Gamez 681 —— Suggested modifications of the Salisbury stomata index devised from a study o Stanhopea (Orchidaceae) RJ. Ferry Sr., R. Foroughbakhch, L.A. Hauad, H. Gamez, J.V. Star, 8. Contreras, and M.H. Badii 691 A synopsis of the genus Gentlea (Myrsinaceae) and a key to the genera of Myrsinaceae in Mesoamerica Jon M. Ricketsun and John J. Pipoly HI 697 New species of Psychotria Subg. Heteropsychotria (Rubiaceae: Psychotrieae) from South America Charlotte M. Taylor The vegetation of Cucphuong National Park, Vietnain Nguyen Nghia Thin TY A new species of Eucrosia (Amaryllidacce) from northern Peru Alan W. Meerow and Abundio Sagastegu i Alva 7O1 Nomenclatural notes on Neotropical Clusicae (Clusiiceae) Jobn J. Pipoly Il CONTRIBUTIONS 705 TO BOTAN) VOLUME 17 NUMBER 4 DECEMBER, LOO7 Vatpfs-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 647 Taste 1. Abbreviations for the 46 characters used in phenetic analysis. Level of accuracy, binary state (BS) and multistate (MS) characters are give Character Abbreviation Accuracy Vegetative sae) STL 1.0cm Culm height CUH 1.0 cm Culm diameter CUD 0.1 mm Culm nodes vertiture NOV Culm vestitur CUV MS Leaf sheath length LSL 1.0 mm Leaf sheath margin vesticure SMV MS Leaf sheath collar vestiture SCV 0.1 mm Leaf ligule length LLL 0.1 mm Leaf blade length LBI 1.0mm Leaf blade width LBW 0.1 mm Leaf blade adaxial vestiture ADV MS Leaf blade abaxial vestiture ABV MS Leaf blade folded or involute LBF BS Leaf blade c aiec esl margins BCM MS Leaf blade base vestiture LBV MS Inflorescence nflorescence length INL 1.0 cm Inflorescence width INW 1.0 mm Nodes per inflorescence INN MS Spikelets per branch SPB MS Inflorescence color INC BS Inflorescence position INP BS Spikelet Spikelet lenght SPE 0.1 mm Number of florets/spikelec NEI MS First glume length FGL 0.1 mm Second glume length SGL 0.1 mm Lemma length LLE 0.1 mm Lemma awn length LAL 0.1 mm Lemma cleft length LCL 0.1 mm mma base vesticure LB [s ma margin vestiture LMP s Lemma lateral awns LLA BS Lemma lateral nerves LNL 0.1 mm Palea length PAL 0.1 mm Palea vertiture PAP MS Spikelet pedicel vesticure SPP MS Anther length ANL 0.1 mm Caryopsis length CAL 0.1 mm Caryopsis width CAW 0.1 mm Ratios Inflorescence length/width RINF oa First glume length/spikelec length REGS oe Second glume lenght/spikelec length RSGS —— First and second glume/spikelec lenght RAGS — Lemma cleft length/lemma length RLEC —— Lemma awn length/lemma length RLAL — 648 Sipa 17(4) time of flowering, soil type, elevation, date, and associated species. Infor- mation on population variation, species variation, and flowering character- istics was recorded at each site. Field collections were supplemented by herbarium specimens obtained on loan from the following major herbaria: A-GH; ANSM; ARIZ; BAF; BRIT; COCA; COLO; CORD; ENCB; MEXU; MICH; MO; NMC; NY; SAT; SD; TAES; TEX-LL; US; WYAC. Voucher specimens collected during this study are deposited at ANSM and TAES. Several populations were collected by collaborators and used in this study. Herbarium specimens obtained from Argentina that were not collected specifically for this study were used as a “real population.” These artificial populations were assigned different numbers to distinguish them from other populations in the phenetic analysis. Phenetic Analysis of Populations. —Populations were collected and identi- fied in the laboratory prior to the phenetic analysis and observations made on the variation for a number of characters. Forty six characters (Table 1) were identified that appeared to show variation among specimens. Sixteen vegetative characters, six from the inflorescence, eighteen spikelet charac- ters, and six derived ratio characters were measured. ee As the characters were measured, each specimen and population were given an @ priori symbol. This was done because certain multivariate tech- niques (e.g. MANOVA) require the use of « prior? designations for analysis. These symbols were given according to certain morphological characters which are listed below. A = Specimens with slender wiry stolons, blade involute, inflorescence not well-exserted, spikelets in leafy fascicles (Dasyochloa pulchella). B = Specimens with inflorescences long exserted, blades conduplicate, lemma acuminate, entire or witha notch 0.1—0.5 mm deep (E. pilosum). = Specimens with lemma truncate with a notch 1.0—2.5 mm deep (E. avenaceum-pilosum complex). O D = Specimens similar to “C” except with the lemma awn longer than the lobes, the spikelets 10-15 mm long (E. avenaceam-grandiflorum complex). Specimens similar to “C” except with the lemma awn scarcely longer than the lobes, the spikelets less chan 10 mm long (E. avenacenm-nealleyi complex). les] T Two methods were used to analyze the variation present among the mor- phological characters. Statistical Analysis Syscem (SAS) [Ray 1982} was used to provide basic statistical data and to perform discriminant analysis. The Numerical Taxonomy System of Multivariate Statistica Programs (NT- SYS) {Rohlf ec al. 1980] was utilized to provide Principal Components Analysis (PCA) and Cluster Analyses were performed with UPGMA and distance matrices. — VaLp£s-ReyNA AND Hatcu, Revision of Erioneuron and Dasyochloa 649 All populations were analyzed using univariate methods to determine groupings and relationships within the taxa, and for selection of characters to be used in multivariate analyses. Cluster analysis and principal compo- nents analysis were used to compare all populations. Electrophoretic Analysis, —Materials for this study were obtained by ger- minating seeds harvested from individual plants from 33 naturally occur- ring populations, from across the geographic range of this genus in North America. A total of 285 individuals were assayed. Table 2 lists the locali- ties where the populations were collected. The electrophoretic sample consisted of crude homogenate tissue from the three uppermost leaves taken from the seedlings grown under artificial light. Starch gels were formed in 22.5 cm x 15.5 cm X 0.8 cm plastic molds similar to those described by Cardy et al. (1980). Two buffer systems were used. Buffer system I consisted of a Tris (0.083M)/citric acid (0.005M) gel buffer (pH 8.7) and an electrode buffer (pH 8.1) of 0.3M boric acid and 0.056 sodium hydroxide. Gels used with system I were of a 8.8% starch (Connaught) concentration. Buffer system II consisted of a histidine (O0.004M)/citric acid (0.0005 M) gel buffer (pH 6.5) and an electrode buffer (pH 6.5) of 0.0064M histidine and 0.0008M citric acid. Gels used with system I were of a 10% starch concentration. Filter paper wicks (12 mm x 2 mm - Whatman #3) con- taining homogenate from each machine-ground sample of leaf tissue were inserted in gels of both buffer systems, 30 samples per gel were examined. Gels were run at 30mA and 250vy for buffer system II, and 30mA and 10Qv rising to 250v for buffer system I. The wicks were removed after 30 minutes. Electrophoresis continued for a total of six hours. Two slices were taken from buffer system I, and three slices from buffer system H. Those taken from buffer system I were stained for the enzymes PGI (phosphoglucoisomerase), and GOT (glutamate-oxaloacetate transaminase). Buffer system II was stained for IDH (isocitrate dehydrogenase), PGM (phos- phoglucomutase), and MDH (malate dehydrogenase). In order to obtain a quantitative measure for genetic interpretation of the phenotypes observed in these species, and to provide a better under- standing of allozyme variation, observational data were taken from routine electrophoretic screens of the taxa. These data included the occurrence and frequencies of isozyme phenotypes. The phenotypes were coded as qualita- tive multistate characters and the NT-SYS package of Rohlf et al. (1980) was employed to perform Cluster Analysis (UPGMA), and PCA. From these analyses, a taxonomic classification of both genera 1s presented. RESULTS AND DISCUSSION Phenetic Analysis. —Characters for which there were a high accounting 650 Stipa 17(4) TABLE 2. Specimens of Ersonenron and Dasyochloa used tor electrophoretic analyses, listed by = country, state, collector, and collection number. Specimens are deposited at ANSM and TAE Erioneuron avenaceum (H.B.K.) Tateoka (O: CoanutLa: La Noria, Va/dés-R. 1568. SAN Luis oS Guadalcazar, Va/dé R. a Enrique Estrada, Valdés-R. 1694, 1695: 10 mi S of San Lusi Potos{, Weldés-R. 1097, 1698. Erioneuron grandiflorum (Vasey) Tateoka AEX IC “0: CoanulLa: Saltillo, Valdé-R. 1507, 1544. Nuevo Leon: El Salero, Valdés-R. 1623, 1627 UNI te D STAT ES: Texas. Presidio Co.: Valdér-R. 1687. a nealleyi (Vasey) Tateoka CO: Sec HLA! eee lo, Valdés-R. 1455, 1531, Hatch 5015. Durango; Poanas, S. nee 2812, 2792. Nugvo Leon: El Salero, Va/dé-R. 1620. TAMAULIPAS: Bustamante, Vialdés-R. 1655, UNITED STATES: Texas. Presidio Co.: Valdéis-R, 1689, Erioneuron pilosum (Buckl.) Nash [EXICO: CoaHulsa: conn Valdés-R. 1502, 1701. Nurvo Leon: Arroyo Mojarras, Valdés-R. 1632. San Luis Porosf: Enrique Estrada, Valdé-R. 1695.5 UNITED STATES: New Mexico. Grant Co.: Va/dé-R. os TEXAS. Briscoe Co.: » ALC. Johnston s.n. Jim Wells Co.: C. Coffey 599. Kimble Co.: C. € ‘offey A-17 Dasyochloa pulchella (H.B.K.) Willed. ex Ryc MEXICO: Coanuta: Saltillo, Hatch et al 905 5a. SAN Luis se Enrique Estrada, Valdé-R. 1696. Zacatecas: Concepcién del Oro, Vald&-R. UNITED STATES: Texas. Presidio Co.: Valdéi-R. 1688. 169] (50%) of interspecies variation were STL, CUH, NOV, LSL, LBW, ADV, ABV, LBF, BCM, INL, INN, SPB, INP, LCL, LLS, LLA, RFGS, RSGS, RAGS, RLEC, RLAL (Table 1). A number of characters not mentioned in previous treatments are con- tributed to explain the variation on the PC axes. These include CUD, LBL, LBP, and ANL. A principal component analysis of 60 populations shows a clear separation of all populations of group A (Fig. 1). The first two compo- nents account for 55.5% of the variation present in this first analysis (31.9 and 23.6 percent of trace, respectively). Specimens of group A are small plants of usually not more than 10 cm tall. A phenogram derived from a correlation of all populations is presented in Figure 2. Group A is sepa- rated from the rest of the groups morphologically. The remainder of the populations analyzed are shown in Figure 3, two clusters are observed, although without a clear separation. The small cluster corresponds to populations of group E. T — ve other cluster corresponds to a complex of groups B, C, D, and some E. Group E was fairly distinct as a sepa- rate cluster. The major characters accounting for variation were size, spike- let characters, and the ratios. Groups B, C, and D appeared to form a con- tinuum in their variation with no clear discontinuity between populations. VaLpEs-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 651 | i@ ‘oO lq | if Fic. |. Principal components analysis of all populations in Ervonewron sensu lata. The clus- ter cae uu to ome right of the main group represents group A (labeled 1, 6, 14 4, 36, 39, These results estimate the overall var soy among the species and in- dicates groupings based on morphological similarities. Group A, which re- presents Dasyochloa pulchella, can be separated distinctly from the rest of populations, which represent the genus Eronenron. Groups B, C, and D cannot be consistently separated on the basis of morphological characters. In gen- eral specimens of group B and C tend to be separated, on some instances, on the basis of lemma awn and lemma cleft length. However, hybridization ts suggested along a continuum of morphological variation given the pres- ence of group D which represents E. grandiflorum. Groups B and C repre- sented E. pilosum and E. avenaceum respectively, and in some instances these taxa can be differentiated. However, intergradation is suggested. Group E, 652 Sipa 17(4) et t t +——____—. | l : ik l l : —— | ———— | call ) l — | ne | eee 3 — LE ! [= | | esachcinaniiciie I i | j ee ! ! | a ! ee { { | (tae , aa : tre. | [ | ! Ly ———-+1_. t t t -———+ Fic. 2. Cluster analysis of all populations, showing three main groups. Phenon line is represented by the dashed line. VaLbEs-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 653 4 we | 4 16 io \ PT 9 | a7 6 a 48 i 1 41 | 7 45 : a f 9 46 I] | a a 8 ' LV | 4 VECTOR 3 CR a ee — Fic. 3. Three dimensional representation of groups B, C, D, and E analyzed with MANOVA. which represents E. nealleyz, is separated from the complex of groups B, C, and D. It is a fairly homogeneous group closely related to group C (E. avenaceum). Electrophoretic Analysis. —Figure 4 shows the electrophoretic activity found in the different enzyme systems assayed, and the following information was gathered. PGI (Fig. 4a): The phenotypes found in this survey showed PGI as a dimeric enzyme; with two loci that appear to be compartmentalized. GOT (Fig. 4a): This system, typically dimeric, appears to have two loci with each locus having two alleles. [DH (Fig. 4b): In this system phenotype | was observed exclusively in D. pulchella. The E. avenaceum-pilosum complex has high variation within this system. However, phenotype 5 appears to be exclusive to “typical” E. pi/oswm from populations in the southwestern United States and northern Mexico. Phenotypes 2 and 4 were common in E. nealleyz, as well as E. avenaceum. Erioneuron grandiflorum had phenotypes 4 and 6, again suggesting a heterozygous condition. PGM (Fig. 4c): This system appeared 654 Stpa 17(4) PGI GOT} — Vd ee, _—_— _—_—e —_ —— —_—_ #?1 3 Penne oii deere 1 2 aeees 00 dd ———_ TE Ce ee eam CO mmm ee GoTu weeee Be ee aenee Bb memes bb rman bb os — wo 1 2 4 5 6 8 10 1 2 (OH eonee — ee — ayy jae 7 z errery — ’ 2 3 ‘4 P r 5 PGM MDH 1 G0 mom, —1~, — Cc Serco nt HOU ae eas od vet a +e-- a — —_ — eas ere — a 5 1 2 3 4 5 1 Fic. 4. 4a. Schematized electrophoretic banding patterns for PGI and GOT. The allelic designations given to the left of some bands are presumed on the basis of observed varia- tion within that enzyme system. 4b. Diagrammatic electrophoretic banding patterns of IDH. 4c. Schematized electrophoretic banding patterns for PGM, and MDH to be monomeric, where the genetic interpretation seemed more obvious. Phenotype | again was exclusive to D. pulchella, phenotype 2 almost exclu- sive to E. avenaceum, and all others species presented a mixture of both. The heterozygous condition (phenotype 3) occurred only in E. grandiflorum. MDH (Fig. 4c): This system showed a dimeric condition with two loci. Dasyochloa pulcbella had phenotype | only, whereas E. vea/ley/ appeared to have pheno- type 2, E. avenaceum and E. grandiflorum had phenotype 4, and E. pilosam had phenotypes 2 and 4. In this system the presence of several unresolved bands suggests new phenotypes. The limited sample number within spe- cies does not allow a stronger differentiation of phenotypes. VaLpEs-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 655 The PCA of 208 individuals assayed shows a clear separation of the D. pulchella individuals as a distinct group. The Erionenron species form two groups without a clear separation. Figure 5 shows the PCA for all species, the populations are represented by one individual. The first two compo- nents account for most of the variation present in the analysis (53.66 and 16.92 percent of trace, respectively). The enzyme systems PGI, GOT, IDH, and MDH contributed to explain the separations of groups. All the D. pulchella samples remained distinct from Erionevron species. Within Evioneuron a cluster representing E. nea/leyi was observed, whereas the E. avenaceum-pilosum complex were mixed. The results suggest that allozyme variation is concordant with the mor- phological analysis for the separation of Erionexron species. Electrophoretic evidence supports Dasyoch/oa as a distinct monotypic genus. In the Erzonenron species, the electrophoretic evidence suggests that “typi- on cal” E. avenaceum and E. pilosum are distinct groups. Erionewron nealleyz might be considered as a distinct group, although closely related to E. avenaceum. Cytological studies indicate that E. avenacenm and E. pilosum are diploid (2n=16), and E. grandiflorum tetraploid (2n=32) [Valdés-Reyna, 1985]. Hybridization in E. grandiflorum is suggested by the electrophoretic data which is concordant with cytological information. CONCLUSIONS The results of these and previous studies suggest that the genus Erzonenron has three species with seven varieties distributed in the United States, Mexico, Bolivia, and Argentina, and also supports the recognition of the genus Dasyochloa. Morphologically che groups identified represent 1) involute leaf blades with inflorescences included in leafy fascicles (D. pulchella), and 2) condu- plicate leaf blades with exserted inflorescences (Erionewron). Dasyochloa has a different electrophoretic, anatomical, and morphological pattern. Within the genus Erionenron, E. nealleyi is a relatively uniform group with deeply cleft lemmas having truncate lobes and long glumes. Erioneuron avenaceum and E. pilosum are two relatively homogeneous taxa, with E. grandiflorum intermediate between them. Electrophoretic analysis and chromosome number reports suggest that E. grandiflorum is a result of hybridization between these two species. TAXONOMY |. Inflorescences long-exserted; stolons when present not ascending to prostrate; blades conduplicate Erioneuron |. Inflorescences included in and surpassed by fascicles of leaves at top of short naked culms; stolons slender, ascending, wiry, abundant; blades involute Dasyochloa Stipa 17(4) TH oe yt -O 1 Fic. 5. Principal components analysis showing first t for all groups. Groups are distinguished as: (group A = Dasyochloa), (group B = Erionenron ne (group C = E. avenaceum - pilosum complex), (group D E. avenacenm - grandiflorum complex), (group E = E. avenaceum - nealleyi complex) ERIONEURON Erioneuron Nash in Small, Fl. S.E. U.S. 143. 1903. Type (Buckl.) Nash. : Evionenron pilosum Tufted perennials, occasionally stoloniferous. Culms erect, leaves mostly basal. Inflorescence an open or contracted panicle or a raceme. Spikelets with several florets, the upper florets staminate or neuter, disarticulation above the glumes and between the florets. Glumes membranous, thin, subequal, acute or acuminate, 1-veined. Lemmas broad, rounded on the back, 3-veined, conspicuously long-hairy along the veins at least below, the central vein often extended into a short awn, the two lateral veins near the margins sometimes extended as small mucros; lemma apex narrowly margined, toothed, or obtusely 2-lobed. Palea slightly shorter than the lemmas, ciliate on the keels, long hairs on the basal portion between che veins. Lodicules two, adnate to base of the palea. Stamens 1—3, having the VaLbis-REYNA AND Hartcu, Revision of Erioneuron and Dasyochloa Goi monandrous condition common in two species. Caryopsis oblong, glossy and translucent; embryo half the caryopsis length. Seedlings with a shaggy- white-villous indument. Basic chromosome number x=8 A New World genus of three species with 7 varieties including the typi- cal; distributed in the United States, Mexico, Bolivia and Argentina. KEY TO THE SPECIES 1 Lemmas entire or with a small notch 0.1—0.5 mm deep; lemma midvein extending into an awn 0.5—2.0(—2.5) mm long; glumes both shorter _ owermost floret — . E. pilosum . Lemmas a0 a notch 1.0—2.5 mm deep; lemma midvein extending ae an awn 2—4 mm long; glumes (at least the second) equal to or longer than lower most floret 2 — 2. Lemma lobes obtuse to broadly acute; lemma notch 1-2 mm deep; lat- eral veins of lemma not extending as small mucros; plants 10-30 cm tall, stolons common 2. E. avenaceum 1.5-2.5 mm deep; lat- eral veins of lemma extending as small mucros to 0.5 mm long; plants 30-65 cm tall, stolons absent 3. E. nealleyi 1. Erioneuron pilosum (Buckl.) Nash in Small, Fl. S.E. US. 144, 1327. 1903. Uralepis les Buckl., Proc. Acad. Natl. Sci. Phila.14:94. 1862 face U, ae Buckl. 95. 1862 = _ Tridens muticus (Tore. ) Nash}. Szeg/ingia pilosa (Buckl.) Nash oS cae Ill. Fl. N. U.S. 3:504. 1898. mh le il Hees ) A.A. Heller, ct. N. Amer. Pl. ed 2. 28. re ei piles Bo )M DA. Cite. 32:9. a. hidens pilosus (Buckl.) A. S. Hite ont 15. Natl. a P3572 1915; middle Texas, eek SM. pes nee not seen; LECTOTYPE, here desig- nated: Wright 781 (PH!)}. i) Lemma lobes rounded to truncate; lemma notch Tricuspis acuminata Munro ex A. Gray, Proc. Acad. Nat. Sci. Phila. 14: ao 1863. Triodta acuminata (Munro)Benth. ex Vasey, Dept. Agric. sien Rep. é3: . 1883. “nomen nudu asey, U.S.D.A. Div. Bot. Bull. 12(2):pl. 32. 1891. a acuminata (Munro) Kuntze, Revis. Gen. Pl. 2:789. 1891. pe: western Texas, Wright 781. (HOLOTYPE: GH!; isoryPe: US!). nna perennials, occasionally stoloniferous. Culms erect, (6—)10— 30(—40) cm tall, glabrous to hispidulous, usually one internode showing above the dense ae of basal leaves, the nodes glabrous to villous. Sheaths smooth, striate, keeled, margins hyaline and often erose, tuft of hairs typi- cally present at the throat, the hairs up to 2 mm long. Ligules 2.0—3.5 mm long, a fringe of hairs. Blades (1—)3—6(—9) cm long, (0.5—)1—1.5(-2.5) mm wide, conduplicate, keeled, rarely flat, usually basal, striate, the margins strongly cartilaginous, the apex acute, pilose near the collar, both surfaces sparsely pilose to glabrous, grayish-green. Inflorescences a compact panicle or raceme, (1—)1.5—4(—6) cm long, (0.5—)1.0—2.0(-2.5) cm wide, ovoid, exserted above the basal leaves (occasionally in overgrazed areas the inflo- rescences borne among the basal culms), with 3—9 short pedicellate spike- lets per branch. Spikelets (6—)8—12(-15) mm long, with (S—)6—12(—20) — 658 Sipa 17(4) Horets, laterally compressed, purplish when immature to pale or pale with a purplish tinge when mature, frequently light green, disarticulation above the persistent glumes. Glumes subequal, glabrous, acuminate, |-veined, the first (2.5—)4—5(—7) mm long, the second (3—)5—6(—7) mm long, shorter than the spikelet, pale, the margins and apex membranous. Lower lemmas (3-)4—S(—6) mm long, acuminate; 3-veined, the lateral veins near the mar- gins, green or purplish green becoming stramineous with age, densely pi- lose on the base of the keel and margins; the midvein extending into a short, straight awn from an entire or minutely notched apex; the notch 0.3—0.5 mm deep, awns 1—2(—3) mm long. Paleas 2—3(—4) mm long, ellip- tic, pilose on the back and margin below. Stamens with anthers 0.3—1 mm long. Caryopsis 1—-1.5 mm long, oblong, translucent. Chromosome num- ber 27=16. Distribution.—Dry rocky hil nn and mesas, often in oak and pinyon-juni- per woodlands. United States southern parce of Nevada, Utah, Colorado, and Kansas, south through Arizona, New Mexico, Texas to northern Mexico in Chihuahua, Coahuila, Nuevo Leon, and Tamaulipas. Present also in Ar- gentina where it is represented by two varieties in the semi-arid zones in the central and west regions north to La Patagonia. KEY TO THE VARIETIES oO? 1. Lemma awns 2—2.5 mm long; plants from me E. pilosum var. ee 1. Lemma awns 0.5—2 mm long; plants from the U aed States, Mexicc a ee 2. Lemma awns to 0.5 mm long; first glumes to 2.5 mm long, second glumes to 4mm long; plants from Provincia de Mendoza, Argentina Ss. ptlosum var. mendocinum ong; firsc glumes 4-5 mm lone, second glumes 5-6 mm long; plants from United States and Mexico 4 2. Lemma awns l—2 mm abs , ptlosum var. pilosum Erioneuron pilosum ae .) Nash var. ee ee (Parod1) Nicora, arwiniana 18:10 1973. Basionym: Tridens pilosa (Buckl.) Hitche. var, mendocina Parodi, Rev. ee Agron. 4:250. 1937. T ypE: ARGENTINA. Mendoza, Las Heras, Potrerillos, estancia El Salto, Ragouese 94 (HOLOTYPE: type, since Parodi (193 / : not seen; the holo- 7) did not designate one when it was described, has not been located. This was also mentioned by Anton (1977). Anton (1977) followed Parodi’s (1937) variety with the geographic dis- tribution restricted to the Provincia de Mendoza. A fragment of Kurtz 10036 (CORD), which Anton (1977) examined, was studied. The taxon belongs to E. pilosum and the varietal description given by Parodi (op. cit.) is as follows. Caespitose perennial. Culms 8-15 cm tall, usually one internode show- ing above the dense tuft of basal leaves, the nodes glabrous. Ligules a fringe of hairs. Blades 2-5 cm long, to 2 mm wide, conduplicate, pilose on the abaxial surface. Inflorescences a panicle 1-2 cm long, contracted, ovoid. VAaLDES-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 659 Glumes |-veined, keeled, acuminate, the BIE 2.5 mm long, the second 3 mm long. Lemmas acuminate, without cleft, 4 mm long, with an awn to 0.5 mm long. Stamens 3, anthers 0.2—0.5 mm long. Ib. Erioneuron pilosum (Buckl.) Nash var. longearistatum (Kurtz) Anton, Kurtziana 10:51—67. 1977. BasionyM: Triodia avenacea vat. ci Sekei Kurtz, Rev. Mus. La Plata $:299. 1893. Typr: ARGENTINA. Colinas secas ce a Estancia La Era, al SO de San ae del Chanar, Kurtz 6729. ee CORD!) i a ee oe ee Kuntze, Revis. Gen. Pl. 3: cna a“ . a (Buckl.) Hitche ar. argentina (Kuntze) Parodi, ve es yeas 937. Erionenron jae eae ) ae var. argentinum (Kuntze) Nicora seca es 18:104. 1973. Type: ARGENTINA. Prov. Rio Necro: Rio Calsada. Lorentz s.n. (HOLOTYPE: “LP?: ISOTYPE: CORD). Ertoneuron ptlosum (Buckl.) Nash var parodianum Sanchez, Darwiniana 22:171. 1979. syochloa argentina (Suntze) Caro var. parodiana (Sanchez) Caro, Dominguezia 2:7. 1981. Type: ARGENTINA. Depro. Linve. CaLeL: La Pampa, Lihuel Calel, Krapovickas 3633 (HOLOTYPE: BAF!; isorype: CORD). Dasyochloa argentina (O. Kuntze) Caro var. aristiglumis Caro, Dominguezia 2:4. 1981. Erioneuron pilosum (Buckl.) Nash var. arsstiglamis (Caro) Sanchez, Lilloa 36:135. 1983. TYPE: ARGENTI NA. Depro. Rivapavia: La Rioja, entre Potrerillo y Porongo, Kurtz 20690 (HOLOTYPE: BAF!). Caespitose oo Culms 15—30 cm tall, leaves basal. Blades 5--10 cm long, 2.0-2.5 mm wide, conduplicate, pilose on the abaxial surface. Inflorescences a panicle, 2-3 cm long, to 2 cm wide, short, dense, con- tracted spikelets 8-10 florets. Glumes acuminate, the first 4-5 mm long, the second 5—7 mm long. Lemmas to 5 mm long, with a notch to 0.5 mm deep, lemma awns (1—)2—2.5 mm long. Stamens 1—3, commonly 1—2, an- thers to | mm long. This is a variety that can be differentiated by the length of the awns which is 2—2.5 mm long, as reported by Parodi (1937) in the original yi ae ae Qa. description. Distribution. —This is a common variety distributed in the central and western parts of Argentina. It is also frequent in the mountains in the Provincia de Cordoba (Anton 1977). Ertonenron pilosum var. parodianum, an endemic variety from Sierra Lihuel- Calel de la Pampa, described by Sanchez (1979) on the basis of lateral veins of lemmas that are pilose only in the lower one third, is included here as a synonym. The character of vestiture of the lemmas was included in the phenetic analysis of this study, and significant differences were not found in the populations analyzed at the inter- and intraspecies level or at the interpopulation level. Therefore, this variety is a synonymy of E. pilosum var. /ongearistatum, which Sanchez (op. cit.) reported as anatomicaly similar to the var. parodianum. A similar situation exists with Dasyochloa argentina var. aristiglumis Caro (1981) which is based on the acuminate glumes pro- —_— 660 Sipa 17(4) jecting into an awn 0.5—1 mm long. In the numerical analysis no signif- cant differences were found for this character. It is important to mention that during the analysis of the paratype of this variety (Stwckert 1761) one stoloniferous specimen was observed. 2. Erioneuron avenaceum (Kunth) Tateoka, Amer. J. oe 48:572. 1961 Trtodia avenacea Kunth, Nov. Gen. Sp. Pl. 1:156 pl. 48. 1815. Uralepis avenacea (Kunth) Kunth, Revis. Gramin. 1:108. 1829. Basso nena (Kune) Will eX Steud., Nom. Bot. ed 2. 1:484. 1840. Tricaspis avenacea Thu x 7, Proc. Acad. ie Sci. eee oes 14:335. 1863. Sh nen ean Revis. Gen. Pl. 2:789. 1891. Tridens avenacens (Kunth) Hitche., Contr. U.S. Natl. Herb. aes 191 3. Type: See ‘Crescit in cele oi exicana inter montem Chapoltepec et Penol de los Banos, alt. 1190 hexap,” Humboldt and Bonpland s.n. (HOLOTYPE: P not seen). Uralepis avenacea (Kunth) Kunth var. v/ridiflora Fourn., Mexic. Pl. 2:110. 1886, “nomen nudum.” Type: MEXICO: San Luis Potosi, Vrer 1379. Triodia avenacea Kunth var. breviartstata Kurtz, Rev. Mus. La Plata 5:301. 1893. Type: MEXICO: Valle de Mexico, Schaffner 158 (HOLOTYPE: US!). a aus Vasey, Contr. U.S. Natl. Herb. 1:59. 1890. Sveglingia avenacea (Kunth) ze var. grandiflora (Vasey) L. H. Dewey, Contr poe Natl. Herb. 2:538. 1894. Ho sy grandiflora (Vasey) — Grasses N. Amer. 2:471. 1896. Tr oe grandiflorus (Vasey) Wooton and Stand. Mexico Agric. Exp. = oe Bull. 81:129. 1912 Evtonenron i re ee Tateoke, Amer. J. Bot. 48:57 2. a “Ero nealleyi (Vasey) Tateoka var. “bors aie (Vasey) Beetle, Phytologia 27:442. 1974 - Ersonenron avenaceum (Kunth) Tateoka var. grandiflorum (Vasey) Gould, ae 26:60. 197 Type: UNITED ST. ne Texas. Presidio Co.: Chenate Mts., Nea/ley 823 nena US!; pararyee: NY!; Toporype: MO!). Tufted perennials, stolons rare to abundant (more common on central Mexico populations). Culms (7—)10—30(—40) cm tall, (0.4—)0.7—1.0 mm diam., glabrous, leaves basal, a single elevated culm node with a reduced leaf, the nodes glabrous to villous. Sheaths smooth, striate, keeled, margins hyaline, a tuft of hairs present at the throat, the hairs up to 2 mm long. Ligules, a — of hairs, to 0.5 mm long. Blades folded (conduplicate) keeled, rarely flat, striate, margins cartilaginous, (1.5—)3—5(—8) cm long, (O.5—)1—1.5(—2.5) mm broad, pilose near the collar, both surfaces sparsely pilose. Inflorescences a panicle, 2-8(—-10) cm long, 1—3 cm wide, compact or open, ovoid, exserted above the basal leaves, stramineous to purple, with 2—10(-16) short pedicellate spikelets per branch. Spikelets 6B—8(—10) mm long, with (4—)6—12(—20) florets, laterally compressed, keeled, purplish, disarticulation above the persistent glumes. Glumes ae glabrous, acuminate, l-veined, the first 4-7 mm long, the second (4—)6—9 mm long. Lemmas 4—6(—7) mm — ong, acuminate, 3-veined, the lateral veins near the margins, densely pilose near the base to the middle, purplish green, lemma lobes obtuse to bluntly acute, the midvein extends into an awn from a notched apex 1—2 mm deep, awns 2-4 mm long. Paleas 2.0—3.5 mm long, elliptic, pilose on the back and margins below. Stamens 1—3, anthers 0.4— Vatpfs-ReyNa AND Hatcu, Revision of Erioneuron and Dasyochloa 66 1.0 mm long or when monandrous the anther to 1.3 mm long. Caryopsis 1.0-1.4 mm long, oblong, translucent. Chromosome number 27=16, 32. Distribution.—This species is a highly variable, common in rocky areas in southwestern United States, to central Mexico, also in Bolivia and Ar- gentina. Included here as a synonym is E. grandiflorum, which is not signif- cantly different based on the phenetic analysis, and the electrophoretic analy- sis. The electrophoretic analysis also suggests a hybrid origin. In Argentina three varieties are recognized following Anton (1977). — KEY TO THE VARIETIES 1. Lemma lobes 0.7—1 mm long; glumes shorter than the spikelet 2a. E. avenaceum var. kurtzianum 1. Lemma lobes 1.5—2 mm long; glumes as long as or longer than the spikelet (at least lowermost floret) 2. Second glumes (7—)8—11(-13) mm long; plants from northwest Argentina E venaceum var. longiglume 2. Second cone 5.0-6.5 mm long; plants from southwestern United States, MEO: r the Andean highlands of Argentina and Bolivia . Lower ian mas 3.5—4.5(—5) mm long; plants from the Andean high cee of Argentina and Bolivia 2b. E. avenaceum var. pygmaeum . Lower lemmas 5—6 mm long; plants from the southwestern United E. ave jon pe sein and Mexico aceum var, avenaceum N ~ . Erioneuron avenaceum Ce Tateoka var. kurtzianum (Parodi) Anton, Kurtziana 10:62. 1977. Bas! IONYM: Tridens avenacea eae Hitche. var. pols Parodi, Rev. Argent. Agric. 4:256, 1937. Type: ARGENTINA. Prov. SAN JUAN. Depto. Zonda: Precordillera entre Barreales, Tontal y ae entre Agua Sine Pedernal, Kurtz 9850 (HoLotyYPE: US!; isorypes: CORD, BAF!). This is a variety Parodi (1937) considered as a possible link er Ee avenaceum and E. pilosum, due to the lemma cleft depth. He suggested the possibility of a hybrid origin. Anton (1977) examined this variety and agreed with the intermediate position. She justified the variety rank on the basis of the lemma lobes, as Parodi (op. cit.) did, adding inflorescence branch pubescence and inflorescence shape. It is considered an endemic variety known only from the type locality. Anton (op. cit.) discussed the differences between typical E. avenaceum and the three varieties that occur in Argentina, since Parodi (op. cit.) suggested that the South American forms might be considered as different species. She concluded that the Argentine material did not possess stolons when com- pared with the type specimen of E. avenacewm from central Mexico. Erionenron avenaceum var. kurtzianum seems most allied with typical E. avenaceum from North America. Both posses glumes that are shorter than the spikelet, and the palea never reaches the lemma awn base. The differences between vari- eties are that E. avenaceum var. kurtzianum has a lemma cleft that is more shallow, and the size of the awns is longer than typical E. avenaceum. ee 662 Sipa 17(4) Study of the type collection leads to the same conclusion that was out- lined by Parodi (op. cit.) and Anton (op. cit.). However, more representa- tive specimens of this variety, need to be studied. In the analysis of the populations from o American E. avenaceum var. avenaceum the lemma cleft was from 0.9-2.2 mm deep, and awn length from 2—4 mm long. These are not fate differences and neither is inflorescence shape or branch vestiture. — 2b. Erioneuron avenaceum ene Tateoka var. pygmaeum (Hackel) nton, Kurtziana 10:63. 1977. Basionya: iy gcraes var. pygmaea Hackel, in R.E. Fries, Nov. Acta Regiae. Soc. Sci. Upsal., ,1:179. 1905. Tridens avenacea var. pygmaea (Hackel) 7 Rev. Argent. on a 3. 1937. Type: oh (HOLOTYPE: W ?; ISOSYNTYPES: COR Variety pygmaewm has a distribution restricted to the Andean highlands of Argentina and Bolivia, usually above 2500 m in elevation. Parodi (1937) characterized the variety as low plants, (less than 10 cm high), with long glumes. Anton (1977) reported the similarity with the var. /ongig/wme, where many intermediate specimens make it difficult to separate the two variet- ies. She suggested the geographical distribution as the most important point. A single collection from Bolivia, Cardenas 204 (GH) was determined as var. pygmaeum. However, as Anton (op. cit.) suggested, a close relation- ship exists with var. /oneze/ume. Therefore, the need for a definitive study on these two varieties exists. —_— 2c. oo avenaceum a ‘Tateoka var. longiglume (Parodi) , Kurtziana 10:65. 1977. Basionym: Tridens avenacea var. longielumis P aa . ev. Argent. Agron. 4: eu 1937. Dasyochloa /ongiglumis (Parodi) Caro var. longiglumis Caro, Dominguezia 2:8. 1981. Type: ARGENTI . PRov. CATAMARCA, Dpto. Andalgala: Las ie ene 1718 (HOLOTYPE: 7; IsoTryPES: BAF! GH! MO! US!). Dasyochloa loneiglumts (Parodi) Caro var. cabrerae Caro, Dominguezia 2:13. 1981. Erronenron avenaceum var. cabrerae seed Sanchez, Lilloa 36:135. 1983. Type: “AR GENTI Prov. La Rioja. Depto. Famatina: Quebrada de la Asai, Cabrera 18096 (HOLO- TYPE: we This variety is characterized by the length of the glumes 7—12 mm long (sometimes equal), and lemma lobes that are oblique anc Distribution.—Between the 1800-3000 m elevation, in northwest Ar- gentina. Included here as a synonym is Dasyochloa longiglumis var. cabrerae which is distinguished by the hair at the base of the lateral veins of the lemmas. As mentioned in the phenetic analysis the vestiture character of the lemmas did not show significance difference. The results of the morpho- logical analysis show that this variety is closely related to the E. nealleys from North America based upon the glumes sizes. on) obtuse. VaLpEs-REYNA AND Hartcu, Revision of Erioneuron and Dasyochloa 663 3, Erioneuron nealleyi (Vasey) Tateoka, ane ee 48:572. 1961. Triodia nealleyi Vasey, “nomen nudum” Vasey, Bull. Torrey Bot. Club 15:49. 1888; Vasey, S§.D.A. Div. Bot. Bull. 12:pl. 36. 1891. Sieglingia ate (Vasey) L.H. Dewey, Contr. U.S. Natl. Herb. 2:538. 1894. Tricuspis alae kes: Heller, Cat. N. Amer. Pl. ed 2. 28. 1900 edie ies (Vasey) Woot. and Standl., New Mexico Agric. Exp. Sta. Press Bull. 8 29.1912. Erioneuron eee var. nealleyi (Vasey) Gould, ae 26:60. 1974. Typr: UNITED STATES. Texas. Pre aa Co.: Cibolo Canyon, Chenate Mts, Neafley s.n. (sorype: US!; paratype: NY!, TopoTtyPEs: GH! NY! US!). Perennials, rarely stoloniferous (only in populations from central México; San Luis Potosi). Culms (15—)35—65 cm tall, 0.8—2.0 mm diam., glabrous to hispidulous, the nodes glabrous to densely villous. Sheaths smooth, stri- ate, keeled, margins hyaline, tuft of hairs present at the throat, the hairs up to 2(—3) mm long. Ligules a fringe of hairs, 0.2-0.6 mm long. Blades 5—10 cm long, 2.0—2.5 mm wide, folded (conduplicate), when moist conditions flat, striate, margins cartilaginous, pilose near the collar, both surfaces pi- lose to villous, green. Inflorescences a compact panicle 5-10 cm long, 1—3 cm wide, narrow, rarely open, ovoid to usually 2—4(—G) times as long as broad, occasionally interrupted in the lower half, light green to purple, with 5-17 short pedicellate spikelets per branch erect to appressed. Spike- lets 7-11 mm long with 3—15 florets, laterally compressed, keeled, pur- to pallid. Glumes subequal, glabrous, acuminate, 1-veined, the first 5—7 mm long, the second 6—9 mm long and generally as long as or longer ie lowermost floret, margins and apex membranous. Lemmas 4—6 mm long, with a notched apex 1.5—2.5 mm deep, the lobes rounded to trun- cate, 3-veined, the lateral veins near the margins, extending into a small mucro less than 1 mm long, scarsely to densely pilose toward the base, the midvein extending into an awn 1—3.5 mm long. Paleas 1.5—3 mm long, elliptic, pilose on che back and margins below. Stamens one (monandrous), anthers 1.0—1.5 mm long. Caryopsis |.3-1.5 mm long, translucent. Chro- mosome number 27= 1 Distribution.—Southwestern United States to northern Mexico. DASYOCHLOA Dasyochloa Willdenow ex Rydberg, Fl. Colorado, Colorado Agric. Exp. Sta. Bull. 100:18,37. 1906. Dasyochloa Wi desea ex Steudel, Nomencl. Bot. ed. 2, 1:484. 1840. “nomen nudum.” Type: Dasyochloa pulchella (Kunth) Willd. ex Rydberg. Stoloniferous or mat-forming, perennials, with tufts at ends of ascend- ing stolons which gradually bend down and take root “mop habit” (Arber 1934). Culms usually less chan 10 cm tall. Blades involute. Inflorescences a raceme included in leafy fascicles. Lemmas notched ca half the length. As 664 Stipa 17(4) in the genus Erzonevron the seedlings becomes shaggy-white-villous. This indument disappears after washing in water and is apparently composed of myriads of hair-like water-soluble crystals which are products of transpira- tion. This condition is previously cited by Parodi (1934) and Johnston (1943). The name is from the Greek “dasys” shaggy, hairy thick-haired, and “chloe” grass. Basic chromosome number x=8. A monotypic genus distributed from central México to California, Ne- vada, Colorado, Arizona, New Mexico, Utah, Wyoming and Texas. The most common desert grass in the creosote-tarbush scrub. ak Sp. 12 55 + 47 ie 1815. Riders la cot es Syst. Veg. 1:332 2, 1825. Uralepis pulchella (Kunth) Kunth, . Gram. 1:108. 1829. pas bulohelle (Kunth) Willd. ex Steud. “nomen on Nomenc! Bot. ed. 484, 1840. Tricuspis pulchella (Kunth) Torr., Pacif. Railr. Rep. Par oe Bot. 4: 156, re Sieglingia pulchella (iunth) Kuntze, Revis. Gen Pl. 2:789. 1891. ini pulchellus (Kunth) Hitche., in Jepson a Calif. L:141. 1912 “Fria pulchellum OXunth) Tateoka, Amer. J. Bot. 48:572. 1961. Type: MEXICO: subfrigidis, siccis, apricis regnit Mexicani inter Cumnaauts Mina de Tee et Cubilete,” Humboldt & Bonpland s.n. (HOLOTYPE: P not seen; IsoTyPe: B-W no 2046, type fragment US!). Trichodiclida prolifera Cerv., Naturaleza 1:346. 1870. Type: near Mexico city “In collibus de Guadalupe, ec de Moctezuma prope Mexicum,” Cervantes 5.2. (HOLOTYPE: ??) Ba See ld aed mean Kuntze var. parviflora Vasey ex Beal, Grass. N. Amer. 2:468. 8906. Type: southern California, Orcvt/ s.n, (HOLOTYPE: US? isorype: TAES, type frag- ment US). Low, strongly stoloniterous, tufted perennials; stolons 3—7(—1 1) cm long, wiry. Culms (1-)4—10(—15) cm tall, scabrous or puberulent, consisting of an elongated internode topped by a fascicle of leaves, which gradually bends down and takes root, producing other culms. Sheaths striate, margins scari- ous, tuft of hairs present at the throat, the hairs up to 2 mm long. Ligules a fringe of hairs 3-5 mm long. Blades (1—)2—6 cm long, strongly involute, with a pointed apex, often curved, abaxial surface scabrous, adaxial surface scaberulous. Inflorescences a panicle, or a short raceme 1.0—2.5 cm long, |.O—-1.5 cm wide, small, capitate, usually not exceeding the blades of the fascicle, light green or purple tinged, appearing white-wooly, with 2—4 short pedicellate spikelets per branch. Spikelets (S—)6—9(-10) mm long, with (4—)6-L0 florets, laterally compressed, disarticulation above the per- sistent glumes. Glumes subequal, glabrous, acuminate, l-veined, awn tipped, the firse 6.0-8.5 mm long, the second 6.5—9.0 mm long, and as rok as or longer than the florets, pale, margins membranous. Lemmas 4.0—5.5 mm long, conspicuously pilose toward the base, 3-veined, dee ply cleft, cleft (I—)3.0—3.2 mm deep, the midvein extending into a straight — Va.pfs-REYNA AND Hatcu, Revision of Erioneuron and Dasyochloa 665 awn from between clearly obtuse lemma lobes, awns (1 5—)2.5-4 mm long. Paleas (2—)2.5—3.5 mm long, long pilose below, ciliate above the keels. Stamens 3, anthers 0.2—0.4 mm long. Caryopsis 1.0-1.5 mm long, translucid. Chromosome number 27=16. Distribution. —Rocky soils in desert regions from southern California east to southern Colorado and south to central México. In México reported from Aguascalientes, Baja California Norte, Chihuahua, Coahuila, Durango, Guanajuato, Hidalgo, Jalisco, Querétaro, México, Nuevo Leén, San Luis Potosf, Sonora, Tamaulipas, and Zacatecas. ACKNOWLEDGMENTS Research reported here was conducted by the Texas Agricultural Experi- ment Station. Financial support in part was received from the Consejo Nacional de Ciencia y Tecnologia (CONACyT) and the Texas Higher Edu- cation Coordinating Board - Advanced Research Program. REFERENCES ANTON, A.M. 1977. Notas criticas sobre gramineas de Argentina. II El género Ersonenron en Argentina. Kurtziana 10:57—67. Arper, A. 1934. The Gramineae. A study of cereal, bamboo, and grass. Repr. J. Cramer Weinheim 1965. Carpy, B.J., C.W. Sruper, and M.M. Goopman. 1980. Techniques for starch gel electro- phoresis of enzymes from maize (Zea mays L.). Inst. Statistics. Mimeo No. 1317, North Carolina State Univ., Raleigh. Caro, J.A. 1981. Rehabilieac én del género Dasyochloa (Gramineae). Dominguezia 2: 17, f. 1-4 — Correll, D.S. and M.C. na 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Ren Duvatt, M.R., P.M on ae A.M.H. Curistensen. 1994. Alliances of Mublenbergia OACEAE) within the New World E ee aaa are identified by re analysis of eee restriction sites from plastid DNAs. r. J. Bot. 81:622-62 GouLp EW. 1951. Grasses of southwestern ie coon Vol. XXII, No. 1. Biol. Sei. Bull. Univ. ns i No. 7. 1968. Grass systematics. McGraw-Hill, New York. 1975. Grasses of Texas. Texas A&M Univ. Press, College Station. 1979. A key to the genera of Mexican grasses. Texas Agric. Exp. Sta. MP-1422. Grnvren W., H. M. Burpet, W.G. CHALoner, V. DEMou in, R. Grote, D.L. HAwKswortH, . NICHOLSON P.C. Sinva, F.A. S11 , E.G. Voss, and J. McNEILL. 1988. Interna- eee code of botanical hme oe Veg. 118. Koeltz Scientific Books, Kénigstein, Germany. Hircucock, A.S. 1913. Mexican grasses in the United States National Herbarium. Contr. U.S. Nat. ete 181-389. 1951. Manual of the grasses of the United States. 2nd ed. rev. A. Chase. A. fie ie . 200, JOuNSON: B.L., D. ak and O. Hai. 1967. Analysis of genome and species relation- ships in the ply ploids wheats by protein electrophoresis. Amer. J. Bot. 54:1089-1098. 666 Sipa 17(4) Jounston, I.M. 1943, Plants of Cox mnt, eastern Chihuahua, and adjoining Zacatecas and eee ~ J. Arnold Arbor. 24:375—421 McVaucnu, R. 1983. Flora Novo-Gi aliciana: A descriptive account of the vascular plants of western ei The Univ. of Michigan Press, Ann Arbor. 14:436 Nicora, E.G. 1973. Novedades agrostologicas patagonicas. tes 18:80—LOG, f. 1—6. Paropl, L.R. 1934. Las gramineas del genero Muvnroa. Rev. Mus. La Plata 34:17 1— 93 1937 7 RENISIOn las gramineas del genero Tridens de la flora argentina. Rev. opens Agron. 4:258- Pererson, P.M., R.D. ee bee J. Vatpts-Reyna. (In Press). Genera of New World Eragrostideae (Poaceae: Chloridoideae). Smithsonian Contr. Bot. Ray, A.A., Ed. 1982. SAS user's guide: statistics. SAS Institute Inc., Cary, North Carolina. Rour, FJ., J. Kispavcu, and D. Kirk. 1980. NT-SYS. Numerical taxonomy system of multivariate statistical programs. University of New York at Stoney Brook. (“Manual” printed from pice! ram.,). RypsBerc, B.A. 1906. Flora of Colorado. Colorado Agric. Exp. Sea. Bull. LOO. SANCHEZ, E. 1979, Anatomia foliar de las especies y variedades argentinas de los géneros Tridens Roem. et eee / a Nash (Gramineae-Eragrostoideae- Eragrosteae). Darwiniana 22:159— 4, . 1983. eae Willdenow ex Rydberg (Poaceae), género monotipico de Norteameérica. Lilloa 36:131—138. SMALL, J.K. 1903. Flora of the southeastern United States. Published by the author, New York. STEUDEL, E.T. 1840. eras botanicus I. Typis et sumptibus J.G. Cortae, Scuttgartiae et ae pe 731 —~ TATEOKA, T. eel A. ee study of Tridens (Gramineae). Amer. J. Bot. AQ 48 : VaLbEs-RreyNna, J. 1985. A biosystematic study of the genus Erionenron Nash we cna aoe Ph.D. Dissertation, Texas A&M University, College Statio Vatpts-Reyna, J. and $.L. Harcu. 1995. Anatomical study of eae and Dasyochloa (Poaceae: C Hioddedae be ons in North America. Sida 16:4 2 ZULOAGA, EO., E.G. Nic , Z.E. RUGOLO bE AGRASAR, O. To tenn 7 PENSIERO, and A.M. CIALDELLA. 1994. Caso os la Familia Poaceae en la Republica Argentina. Monogr. Syst. Bot. Missouri Bot. Gard. « GEOGRAPHIC SPATIAL AUTOCORRELATION OF MORPHOLOGICAL CHARACTERS IN HEMEROCALLIS HAKUUNENSIS (LILIACEAE) SOON SUK KANG Department of Biology Gyeongsang National University Chinju 660-701, THE REPUBLIC OF KOREA KI BAE PARK Department of Horticulture Ansung National University Ansung 456-749, THE REPUBLIC OF KOREA MYONG GI CHUNG Department of Biology Gyeongsang National University Chinju 660-701, THE REPUBLIC OF KOREA ABSTRACT Spatial autocorrelation analysis of 12 quantitative mong 30 populations of Hemerocallis hakunnensis was conducted to understand better their peor variation patterns. There are two types of ee between the mean values of the characters measured and their spatial autocorrelations: (1) significant heterogeneity of means w - significant autocorrelation (mostly floral characters); and (2) significant heterogeneity of means with no significant autocorrelation (mostly vegetative characters ). Perianch size shows significance north-south clinal variation and width of inner and outer perianth show a typical monotonic decline from significant positive autocorrelation at eeppslation cee of 67 to 107 km to significant negative autocorrelation at distances of 221 to 357 km. TI ults might result from the combinations of gene flow, genetic drift, and/or selec- tive Ge coe in populations of H. hakuunensis. RESUMEN alizé un andlisis de correlacién espacial de 12 caracteres Cuantitativos en 30 soblecienks de Hemerocallis hakuunensis para comprender mejor sus patrones de variacion geografica. Hay dos tipos de relaciones entre los valores medios de los caracteres medidos y sus autocorrelaciones espaciales: (1) heterogeneidad significativa de las medias con autocorrelacién significativa (la mayorfa en caracteres florales); y (2) heterogeneidad significativa de las medias sin autocorrelaci6n significativa (la mayoria en caracteres jon vegetativos). E] tamafio del perianto muestra una variaci6n clinal norte-sur significativa y la anchura del perianto interno y externo muestra una disminuci6n monotonica tipica esde una autocorrelacién positiva significativa en distancias interpoblacionales de 67 a 107 km hasta una autocorrelacién negativa significativa en distancias de 221 a 357 km. Los resultados podrian derivarse de las combinaciones de flujo genético, deriva genética y/ o fuerzas selectivas que operan en las poblaciones de H. hakuunensis. Stipa 17(4): 667—675. 1997 668 Sipa 17(4) INTRODUCTION In Korea, five hae of Hemerocallis are recognized on the basis of mor- phological analysis (1 2 quantitative and seven qualitative characters), field- work, and examination of Japanese herbarium specimens (Chung & Kang 1994; Kang & Chung 1997a). These are H. hakuunensis Nakai, H. thunbergii Baker, H. middendorffii Tr. et Mey., H. hongdoensis M. Chung & S. Kang, and Hf. taeanensis 8. Kang & M. Chung. Hemerocallis bakuunensis is Korean endemic species that commonly grows at the margins of pine-oak forests, grasslands, and disturbed habitats such as chest-nut orchards in southern, central, and northwestern Korea (Chung & Kang 1994). Among the four Korean Hemerocallis species, populations of H. hakuunensis are the most variable in their inflorescence and floral morphology (Kang & Chung 1994). In addition, Chung and Kang (1994) noted that populations of H. hakuunensis with larger flowers are generally located in the central Korean Peninsula, while populations with smaller Howers occur in southern Korea. These patterns of geographic variation in morphological characters of the species require further study for a better understanding the species. A detailed study of the patterns of geographic variation of quantitative morphology in H. hakuwunensis is reported here. The principal method of analysis for this purpose is spatial autocorrelation analysis. Spatial autocorrelation statistics are summary measures of the de- pendence of the value of a particular variable at one location on the value of that same variable at other nearby locations. The most commonly used measure 1s Moran's (1950) autocorrelation coefficient I, in which geographic neighbors are compared in terms of their deviation from the mean of al observations. The method has been applied in a variety of fields including biogeography, political science, ecology, archaeology, and econometrics (Cliff & Ord 1981; Griffith 1988). Recently, this method of character analysis has been employed in the analysis of morphological characters and the spa- tial dispersal patterns of genetic variations because it can provide a detailed picture of geographic variation in characters (Sokal & Oden 1978a,b: Sokal et al. 1986; Jensen 1986; Chung 1995, 1996). Sokal and Oden (1978a) described five correlogram patterns of geographic variation of characters. Cline is a gradual change from the highest positive / to the lowest negative I, across all distance groups. Depression is nothing but a circular cline, with the lowest negative / between the shortest and longest distances, not the longest distance. Double depression is on the whole quite similar to that of the depression, with low / in a certain distance from which another low J was separated. Intrusion patcern shows positive /s at a certain dis- tance, because of the homogeneity within che distance, with a sharply de- —_— KANG, ET AL., Hemerocallis hakuunensis 669 creasing I bordering the distance. Crazy quilt pattern shows high Js sur- rounded by low Is, vice versa, which yields no significant Js at all. The purpose of this study is to determine whether or not the 12 quanti- tative characters recorded for H. hakuwunensis by Chung and Kang (1994) and Kang and Chung (1994) show any geographic patterns such as clines, depression, double depression, intrusions, and crazy quilt patterns (Sokal & Oden 1978a). MATERIALS AND METHODS Thirty populations Sea: the geographic range of H. bakuunensis were used for this study (Fig. 1). Measurements of 12 quantitative charac- ters (Table 1; also see Table 2 in Chung & Kang 1994) were taken on each of 10 randomly selected individuals directly from their natural habitats from 1991 to 1995. Voucher specimens of all collections are deposited at the herbarium of Gyeongsang National University (GNUC). For spatial autocorrelation analysis, mean values were assigned to each population for the 12 characters. Every possible pair of populations was considered as a join or a connection and was assigned to one of six distance classes based on the geographic distance between them. These six distance classes were constructed by equalizing sample sizes among the classes. The distance classes are 0<76, 76<102, 102<136, 136<169, 169<221, and 221<357 km. Moran’s J values were calculated for interpopulational dis- tance classes by aN SUV ZOE 1] i] (Sokal & Oden 1978a). N 1s the num weighting matrix, where Wj; is set as one if 7th and ych population are in the distance class and zero otherwise, Z; = Xj - X, Z; = Xj; - X, the variables Xj and Xj are the mean scores for /th and sth ar respectively, and X is the mean score for all populations. The value of J ranges between + | = der of populations, Wj; is the join on (complete positive autocorrelation, i.e., paired populations have identical values for all characters) and -| ample negative autocorrelation). Each / value was used to test significant deviations from the expected values, E(/)=- 1/(N-1) (Cliff & Ord 1981). The neighboring populations in the distance class considered, with a significant positive value of Moran’s J have similar scores, whereas those with a significant negative value should have differ- ent scores. Overall significance of individual correlograms was tested using Bonferroni's criteria (Sakai & Oden 1983). All calculations and statistical analyses were performed using the SAAP program (ver. 4.3) written by D. Wartenberg. Sipa 17(4) 670 Alig P te, f , ¢ * . it tr i * het 4 é ® v * + + 7 ig + + a 4 mer * a ” oa " * < x ae in “i Pte ae re » ” x af “ “ ra KOREA a iv 2 g a @ G ‘ Arena @ ‘ oat e r FL ) e \ . e e @ ‘Nua @ : @ @ e t se @ 4 of Pree - x 5 a” °¢ ee, ‘ ‘< et éj Pal PUES QE Ny QS 7. . 48 3 aw ame Li JAPAN VA : x Pear Fic. 1. The location of 30 examined populations of Hemerocallis hakunnensis. KANG, ET AL., Hemerocallis hakuunensis 671 TasLe 1. List of 12 quantitative characters used in the spatial autocorrelation analysis. Acronym Character derivation Unit or category PSH Plant (scape) height m LIF Length of inflorescence minus flowers cm NBS Number of bracts/scape NFS Number of flowers/scape it LLB Length of the lowest bracts cm WLB Width of the lowest bracts cm LPO Length of the perianth tube enclosing an ovary cm LIP Length of the inner perianth cm WIP Width of the inner perianch cm LOP Length of the outer perianth cm WOP Width of the outer perianth cm Ww Width of the widest leaves cm RESULTS Correlograms for each character are presented in Figures 2-3. Moran's / was significant in 26 of 72 cases (36%). The overall correlogram for width of the lowest bracts (WLB), length of the inner perianth (LIP), width of inner perianth (WIP), length of outer perianch (LOP), width of the outer perianth (WOP), and width of the widest leaves (W WL) was significant (P < 0.01). As presented in Figures 2—3, plant (scape) height (PSH), length of inflorescence minus flowers (LIF), and length of the perianth tube enclos- ing an ovary (LPO) show depression correlograms (Sokal & Oden 1978b). On the other hand, number of bracts per scape (NBS), number of flower per scape (NFS), WIP, and WOP represent clinal variation with two, WIP and WOP, significantly autocorrelated (Figs. 2-3). In addition, it is apparent that each floral size character shows a similar pattern of geographic varia- tion. For example, LIP and LOP, WLP and WOP (Fig. 3) revealed significant positive autocorrelation (localities connected for that distance class had simi- lar values) for distance classes | to 3, followed by a steady decline until significant negative autocorrelation (localities for that distance class have dissimilar values) are recorded for distance class 5. Although LIP and LOP show increase in autocorrelation between distance classes 5 and 6, significant negative autocorrelations are also observed on these distance classes (Fig. 3). As seen in Figures 2—3, length of the lowest bract (LLB) has a crazy quilt correlogram pattern with no significant autocorrelations, whereas width of the lowest bracts (WLB) and WWL display intrusion patterns. DISCUSSION There were two types of relationships between means and spatial autocorrelations observed. The first pattern of significant heterogeneity of means with significant autocorrelation was observed for WLB, LIP, WIP, O10 7 0150 | ee | e. | i . ] ~ : 0100 4 \ 0.100 x PSH | | \ LIF | e ! 0.080 + 0.050 aN | . nea mane yh \ | 0000 \ e . wn \ | “” = a a eee ———/ “co 0.000 \ | g -0.050 e / E : 7 S -0.100 \ f \ o- \ f s 0.050 e oe | s \ / , \ (4 “0150 4 \ “0100 \ / ° | ‘\ | ‘ ‘ 0 200 \ \ | ; \ f 01580 | \ if | 0.250 arf \ 250 | . | -0 200 | Pie Ey ero (kd at ea . sa ea aad an a Gs | -0 300 | pe ye ee 0 1 3 4 5 6 0 ! 2 4 5 6 Distance Distance 0100 4— = . = 0.100 = = — Se ) o | NBS | ‘ NFS 0.050 0.050 z s | UA | ~ \ 4 \. Elgg . eo oe : a 0.000 \ wm 0.00) 4 e : | rz baw rot \ i — 2 4 ee een mee S 0050 | eo S 0050 \ | \ 0.100 =| \ -0.100 & . on | : | ~ | < UL i na Oe 0 150 cen ns SR ee a eee 0 \ 2 3 4 $ 6 0 | 2 4 Distance pia 0.100 7 = > ae 02= _ > I \ . 7% a A LLB / \ WB 0.050 x % / \ 01 oe PS | f \ \ a? f ~ 4 \ / \ \ / ~ \ f \ \ / ‘ 000 | / \ Pf \ | , 00 - x / * | s ‘ / \ } = ee eS ee ee a” 1 a \ é \ a 4 \ / ct =“ 5 \ . 30.0505 \ / \ S01 ay, \ = { f \ i) WJ \ = \ / \ = \ “0.100 \ f 0.2 + \ ‘ ; \ \ | \ / 0150 | e 03 \ | ‘ > 20.200 ==} qa a toot AA prereset | oO 3 4 > 0 ! 4 3 4 5 6 Distance Distance Fic. 2. Correlograms for PSH, LIF, NBS, NFS, LLB, and WLB. Significant autocorrelation coefficients (P< 0.05) are indicated by asterisks. LOP, WOP, and WWL. The second pattern of significant heterogeniety of means with no significant autocorrelation is found for the other six charac- ters. Sokal and Oden (1978b) suggested biological explanations for each of the two types of patterns. The first could resule from migration, founder effects, and selective agents. The second type of relationship between means and aucocorrelations could result from drift, weak migration and selection, frequent local extinctions followed by the establishment of new patches by founders, or from selection in isolated, patchy environments. Except for LLB, all characters show short distance positive autocorrelations, indicat- ing that migration could be operating over the distances. Certainly in some KANG, ET AL., Hemerocallis hakuunensis Moran's | “0.100 4 / 0.150 4 oe / 1 / Distance Moran's | | ea ae la aes ] -0.80 Jor Pare ore pore ore pe rer 0 | 2 4 3 Distance 03 wa ve | 02 ae . LIP 01 a \ seal | n = 0° | cd S01 2 * | 02 : . » \ AA 0.3 \ we | 04 +> a ee (age ai (SLL (LL 4 1 at 5 6 Distance 03 ——-———— ———- — 7@ 0.2 LOP - a Ol hdl | et \ | nn 2 00 x : = Ol = La 02 \ | \ < 03 \ J 4 £08 pa qo aes ( ' 2 3 4 5 6 Distance 06 = —- See eee — 04 . A WWL 024 . I! ne) | \ we im \ . so 00 \ 8 sisi PME. Tee ae : a = ‘ r : 0.2 2 1 f 0.4 | Ww | 0.6 jy saLLa SELES GL | 0 1 2 3 4 5 6 Distance Fic. 3. Correlograms for LPO, LIP, WIP, LOP, WOP, and WWL. Significant autocorrelation coefficients (P < 0.05) are indicated by asterisks. of the populations examined there are small patches whose diameter is less than interpopulational distances. The width of the lowest bracts and the longest leaves yielded a relatively long distance positive autocorrelation, which indicates these characters occur with circular or symmetrical gradi- ents. It is of interest to note that the width of inner (W/IP) and outer (WOP) perianth show a typical monotonic decline from significant positive autocorrelation at 67 to 102 Km toa significant negative autocorrelogram from 221 to 357 Km. Monotonicity is expected when separation by dis- tance and gene flow between neighbor populations is the main factor. 674 Sipa 17(4) The results of this study support the suggestions by Chung and Kang (1994) that there is a gradual transition of the floral morphology of H. hakuunensis from larger flowers in the central Korean Peninsula to smaller flowers in the southern Korean Peninsula. A similar pattern of geoclinal variation of floral morphology is previously reported in H. dumortieri com- plex in japan (Noguchi 1986), who noted that the flower tube enclosing the ovary and the length of the internal as well as external perianths exhib- ited a geoclinal variation. For example, northern populations possessed much shorter, somewhat thicker flower tubes and smaller perianths in compari- son with those from southern populations. In general, vegetative characters have much showed phenoplasticity than do reproductive characters in response to environmental and developmen- tal factors (Stebbins 1950). It might be more reasonable to discuss the biological meanings of the results using reproductive characters. Accord- ing co the criteria of Sokal and Oden (1978b), patterns such as those seen in several reproductive characters of H. hakuunensis might result from the combinations of gene flow via pollen and seed dispersal, founder effects, genetic drift, selective forces in patchy, isolated environments. More re- cently, we have analyzed the spatial distribution of genotypes using spatial ations (20 x 20—m) of H. bakuunensis. — autocorrelation of 28 alleles in popu A genetic similarity was shared among individuals within 4.5 m distance, whereas an overall genetic dissimilarity among individuals beyond the dis- tance, indicating that individuals in populations of H. hakuunensis occur on a gradient (Chung unpubl. data). Based on allozyme data obtained from 19 populations of H. hakuunensis, Kang and Chung (1997b) further suggested that gene movement and genetic drift may be primary factors for shaping population genetic structure among populations of H. bakaunensis, ACKNOWLEDGMENTS We thank Sun Gi Chung, Duk Seo Gu, Ou Kun Chin, Seok Jun Yun, Jun Gi Hong, and Gap Su Oh for their company on field trips. Special thanks go to J.G. Conran and D.J. Boyler for comments on the manu- script. We are also grateful to the Department of National Monuments of Culture Property Preservation Bureau of Korea for permission to collect samples from Hallasan, Hanryohaesang, Sobaekan, Sokrisan, Tasohaesang National Parks and Hong and Cheju islands. This research was supported in part by a grant from the Korea Science and Engineering Foundation (96-0500-006-2) to MGC. REFERENCES CuHuNnG, M.G. 1995. Spatial genetic structure among ees aa of Exrya japonica and Ef. emarginata (Theaceae). Ann. Bor. Fennici 32 KANG, ET AL., Hemerocallis hakuunensis 675 Cuunc, M.G. 1996. Spatial genetic structure among Korean populations of Hosta minor and H. capitata (Liliaceae). Bot. Bull. Acad. Sin. 37:25— CuHu M.G. and S.S. Kana. 1994. ee jometric analysis of dhe Hemerocallis L. (Liliaceae) in Korea. J. Pies Res. 107: 165-17 CLIFF, AD. and J.K. Orv. 1981. sae processes-methods and applications. Pion Lim- ited, London. GrirfitH, D.A. 1988. A oo on spatial autocorrelation. Washington DC: Association of American Geographer JENSEN, R.T. 1986. eee spatial autocorrelation in Quercus e/lipsoidalis. Bull. Torrey Bot. Club. 113:431-439. KANG, : ad M.G. Cuunc. 1994. Hemerocallis hakuunensis (Liliaceae) in Korea. Sida 162 oe = and M.G. CHUNG. 1997a. Hemerocallis taeanensis (Liliaceae): a new species from oe Bot. (in pres KANG, on and M.G. one 1997b. Genetic variation and population structure in Korean endemic species: IV. Hemerocallis hakununensis (Liliaceae). J. Plant Res. 110: 209217 Moran, P.A.P. 1950. Notes on continuous stochastic phenomena. Sona 37:17-23. Nocucut, J. 1986. Geographical and ecological differentiation in the H lumorti complex with special reference to its karyology. J. Sci. Hiroshima Univ., ines Japan, Ser. B. Di. 2, Bot. 20:29-193. Nocucui, J., M. Tasaka, and M. Iwasucut. 1995. The historical differentiation process in Hemer aie pelle Liliaceae) of Japan based on restriction site variations of chlo- roplast DNA. J. Plant Res. 108:41-45. Sakal, A.K. ve N.L. Open. 1983. ot pattern of sex expression in silver maple (Acer saccharum L.): Moristia’s index and spatial autocorrelation. Amer. Naturalist 122:489- O8. SoKAL, R.R., T.J. CRovELLO, and R. UNNaSCH. eae oe raphic variation of vegetative characters of Populus deltoides. Syst. Bot. 11:418— SokaL, R.R. and N.L. a Spatial ec dice in biology. 1. Methodology. Biol. J. Tina, Soc. igo oe SoKAL, R.R. and N.L. ODEN. ea Spatial autocorrelation in biology. 2. Some biological implications and four applications of evolutionary and ecological interest. Biol. J. Linn. soc. 10:229—2 StrepBins, G.L. 1950. Variation and evolution in plants. Columbia Unity. Press, New York. ow BOOK NOTICES DuNMIRE, WILLIAM W. and Gait D. TigERNEY. 1997. Wild Plants and Na- tive Peoples of the Four Corners. (ISBN 0-89013-319-0, pbk). Museum of New Mexico Press, P.O. Box 2087, Santa Re e, New Mexico 87504. $22.50..312 pp. This handsome book is both a traveler's companion and ready reference work. The authors have thoughtfully compiled and beautifully presented a wealth of information on the cul- tural and natural heritage of the Four Corners region of the American Southwest. The first half of the book examines the land, prehistoric and extant cultures, and plant uses by native peoples living in southwestern Colorado, northwestern New Mexico, southeastern Utah, and northeastern Arizona. The second half of this book is devoted to oneaare illustrated (with excellent color photographs) ethnobotanical descriptions of fifty taxa or- ganized according to lifeform—trees, shrubs, grasses and herbs. An annotated table ‘“ 515 useful plant taxa appears after the species’ ethnobotanical descriptions. This table provides a marvelous synthesis of culture-specific plant uses summarizing information from a long list of historic and recent literature. The bibliography is extensive and the index detailed. If you plan to travel in the Southwest or live there and plan to explore the region’s natural and cultural riches, buy this book to help you do it.—Brace Benz. BOARD ON SCIECE AND TECHNOLOGY FOR INTERNATIONAL DEVELOPMENT, NA- TIONAL RESEARCH COUNCIL. 1996. Lost Crops of Africa. Vol. Grains. (0-309-04990-3, pbk). National Academy Press, Washing- ton, D.C. $24.95. 383 pp. USAID has done yet another service by sponsoring BOSTID’s compilation of information on useful African biodiversity. This ce provides an extremely useful treatment of six important African gr rains—Africi in Rice, Finger Millet, Fonio, Pearl Millet, Sorghum and Tef , six short treatments discussing very locally distributed cultivated species and numer- ous other wild grains barheree i. ree ins for food. Taxonomic and evolutionary relation- ships of cultivated grains and wild relatives is provided and comparative nutritional value of each grain is compared to maize, the grain that has come to dominate a majority of cultivated acreage in Africa. If you teach a course in Economic Botany and don’t mention these grains, do yourself and your students a favor by incorporating the information pro- vided in this book.—Brace Benz. Sipa 17(4): 676. 1997 NEW SPECIES OF RHAMNACEAE FROM YUNNAN, CHINA FAN GUO-SHENG and DENG LI-LAN Forest Plant Laboratory Southwest Forestry College Kunming, 650224 Yunnan, CHINA ABSTRACT Two new species of Rhamnaceae—Sageretia gongshanensis G.S. Fan and L.L. Deng from Gaoligongshan Mountain, Yunnan, China and Rhamnus datiensis G.S. Fan & L.L. Deng from Dali City, Yunnan, China are described and illustrated. RESUMEN Se describen e ilustran dos especies nuevas de Rhamnaceae—Sageretia gongshanensts GS. Fan and L.L. Deng de la Montana Gaoligongshan, Yunnan, China y Rhamnus daliensis GS. Fan & L.L. Deng de Dali, Yunnan, China. Key worps: Rhamnus, Sageretia, Rhamnaceae, Yunnan, China, new species pea gongshanensis G.S. Fan & L.L. Deng, sp. nov. (Fig. 1). Tyee: CHIN . YUNNAN: © wesenschas Mountain, 17 Apr 1979, He Zhi-Ming 79-0185 van Xian (County), 6 Oct 1988, > (HOLOTYPE: KUN); ParatyPe: CHINA. Yunnan: Baos Investigative team 7625 (SWEC) Species haec S. /axifloram Hand.-Mazz. Valde similis praesertim habitu, a qua differc foliis tenuis chartaceis vel submembranaceis, venis lateratibus non impressis supra, elevatis infra, venis principalis infra tomentosis fulvidis vel brunneolis. Woody vines, 8 m long; young shoots with gray-brown tomentum, old shoot glabrous. Leaves thin papery or submembranous, subopposite or al- ternate, ovate to elliptical, 4-9 cm x 2.5—5.0 cm, adaxial side densely tomentose when young, glabrescent and dull in old age except for yellow- brown tomentum near veins, the apex acuminate, the base rounded to sub- cordate, veins 5—7 pairs, not impressed abaxially, margins serrulate; peti- oles 5-9 mm long, dense brown tomentose. Flowers unknown. Inflorescence axillary, spicate, densely brown tomentose, 7-10 cm long. Fruit a drupe, obovate, seeds 2—3. This new species is similar to Sageretia laxiflorum Hand.-Mazz., and is easily recognized by its thin papery or submembranous leaves, lateral veins beneath, the midrib and lateral veins yel- en not impressed above and raisec low-brown tomentose beneath. Stipa 17(4): 677-680. 1997 678 Sipa 17(4) Fic. | Sageretia gongshanensis G.S. Fan & L.L. Deng sp. nov. 1. Shoot with leaf, 2. Inflores- cence with fruits, 3. Fruit. 4. Seeds, 5. Amplified lower surface of leaf. Drawn by H.P. Wang. FAN AND DrENG, Rhamnaceae from Yunnan, China 679 \\ | = Fic. 2 Rhamnus daliensis G.S. Fan & L.L. Deng sp. nov. 1. Shoot with fruits and leave, 2. Fruit, 3. Amplied lower surface of leaf. Drawn by H.P. Wang. 680 SIDA 17(4) Rhamnus daliensis ri - Fan & L.L. Deng, sp. nov. (Fig. Sees YUNNAN: Dali City, in evergreen needle- leat | forest, 2000 m alt., 21 Jul 1956, Hsw Yune-chun AIS6 cee PE: SWEC), Cy A Rhannas globosus athnis, sed differt foliis majoribus (4-9 em x 2—2.5 cm), ellipticis vel oblongis, raro obovatis, non rotundatis, venis lateralibus S—7-Jugatis. Shrubs; young shoots grey-brown, opposite or subopposite; young shoots pubescent. Leaves papery, opposite or subopposicte, rare alternate, elliptic to oblong, rarely obovate, not rounded, 4—9 cm long, 2—3.5 cm broad, apex obtuse to acute, widely cuneate or slightly rounded, the base s slightly oblique, margins finely serrulate, the midrib pubescent or glabrous and light yellow above vane dry, densely pubescent and yellow beneath when dry; lateral veins 5—7 pairs, slightly impressed above, raised beneath; stipe 4—8 mm long, lence y pubescent; stipules lanceolate, persistent, densely pubescent. Flowers unisexual, clustered in leaf axils. Fruit subglobose, black, about 4 mm diam., persistent calyx at base, stem of fruic 5—8 mm long, glabrous to rarely short pubescent, seeds 2. This new species has a close affinity to Rhamnus elobosus Bung., but it is easily recognized by its elliptic to oblong, rarely obovate, non rounded, 9 cm long, 2—2.5 cm leaves, and lateral veins of 5—7 pairs. — ACKNOWLEDGMENTS This research was supported by the Academic and Technical Leader of Ministry of Forestry in China, Academic and Technical Leader of Yunnan Province, and Natural Science of Yunnan Province Funds. LEAF-PRINT ANALYSES: AN ECOLOGICALLY FRIENDLY METHODOLOGY FOR PLANT IDENTIFICATION R.J. FERRY SR., R. FOROUGHBAKHCH, L.A. HAUAD, S. CONTRERAS, J.V. STAR, M.H. BADH, and H. GAMEZ Facultad de Ciencias Biolégicas-Division de Estudios de Postgrado Universidad Auténoma de Nuevo Leon, A.P. F-2 San Nicolas de los Garza, N. L., MEXICO 66450 ABSTRACT Because the taxonomy of Stanbopea (Orchidaceae) has been established exclusively on analyses of the flower, field identification is complicated by the fugacious habit. As an aid to species confirmation, a system is offered that ts ecologically friendly, economical, and statistically based. Clinical use of this system suggests it may also have value in determi ing the degree to which natural and man-made hybrids are related to each species-parent. RESUMEN Debido a que la taxonomia de Stanhopea (Orchidaceae) ha sido estab en base al andlisis de la flor, la identificaci6n en el -omplhc lecida exclusivamente | debido a su fugacidad. se ofrece un sistema que es respetuoso ecolégicamente, econdmico, y con base estadistica. El uso clinico de este sistema sugiere que posiblemente tenga valor en la determinacion del grado en el que los hibridos natu- rales y los obtenidos por el hombre estan relacionados con cada parental. Como ayuda para la identificacién de esta especie, s INTRODUCTION The genus Stanhopea was named in honor of Sir Philip Henry, the 4th Earl of Stanhope (1791-1855), president of the London Medico-Botanical Society from 1829-1837. Estimates of its size are numerous with old pub- lished estimates often repeated without reference to recent sources: Hawkes (1965: 8—25 or more); Hamer (1 974: ca, 20); Arditti (1992: approximately 50); Dressler (1993: 55); and Bechtel et al. (1992: ca. 25). Jenny (1993) offers “about 52 species, two varieties (subspecies) and six natural hybrids.” A search of the literature coupled with clinical research suggests that com- bining the Dressler and Jenny estimates offers the most accurate estimate for the size of Stanbopea. The genus is known from Mexico, throughout Central America, east- ward across northern South America, and south-southwest into Bolivia, Ecuador, and Peru. Its northernmost invasion is reported by Kennedy (1974) from western Mexico, at a latitude farther north than San Antonio, Texas Sipa 17(4): 681-690. 1997 682 Stipa 17(4) while its souchernmost reach is from the area of Sao Paulo, Brazil (Pabst & Dungs 1977). oral keys have been offered for regional areas, with most concentrating on Mexico. A key by Williams (1951) was followed by publications by Ames and Correll (1952, 1953), and a key to the Mexican members by Dodson (1963). Kennedy (1975), working with Dodson’s more recent tax onomy (1975), published a hierarchy of the genus Stanhopea in Mexico, and more recently Williams and Whitten (1988) offer a key to the Stanhopea species of Panama. However, a comprehensive well illustrated treatment of the genus has yet to be offered and, in fact, the genus has compratively ignored because its flowers are generally not long lasting. Curtis (1910) cites Stanhope flowers as large and very attractive, but notes that “they are short-lived and cannot be used in floral decorations, hence cannot be con- sidered first class.” Stanhopeas have thus been relegated to the domain of orchid aficionados with the desire and space to maintain them, and as an occasional study topic by orchid botanists. lal QO Species identification is necessary for government support of conserva- tion. Lawmakers bluntly want to know the specific identity of the organ- ism targeted for government conservation support. In addition, compared with other national priorities, funds for conservation are in short supply in the United States and are even less available in most of the countries situ- ated in tropical areas where much of the world’s orchid flora is abundant. hus, working against their preservation are explicit and implicit costs attendant to identifying orchids in their habitats. New problems arise as field collections are to be made of material to be used in the identification process. ee often take a dim view of removing plants or plant parts, and usly—any plant chat’s collected, pressed and dried will never again set seed in its natural habitat. “The removal of even a “window” of lamina, as outlined by Cutler (1978) and utilized by Stern and Morris (1992) damages the plant (albeit minutely), opens the door to local political restrictions regarding the removal of plant material from the habitat, and requires detailed laboratory dissection and staining procedures. In general, as procedures become more complex, equip- ment costs increase and the likelihood of artifact introduction likewise in- creases. Adding to the problem is chat some of these orchids flower for only very short periods during the year. Stavhopea flowers last only a few days so one needs to be at exactly the right place at the right time or to be able to identify members of this genus when they're noc in flower. However, most orchid taxonomy is based on analyses of the flower. Indeed, Curry et a (1988) state “the taxonomy of Stanhopea species rests exclusively (italics ai) on the morphology of the flower, changes which have apparently been influenced by the pollinators.” _ Ferry, ET AL., Leaf-print analyses for plant identification 683 Thus, although problems remain with trying to correctly identify plants, this approach attempts to be bounded by parameters which clearly meet local economic, political, and botanical benchmarks. Strict requirements have been self-imposed for this study, and the identification system must meet all of the following tests: 1. It must require minimal material in che field and only that equipment in-house or easily obtainable by the laboratory of a foreign university. The field work and laboratory work should be able to be carried out by any properly trained secondary science student or university undergraduate student of average ability. 3. The identification protocol should be “ecologically friendly,” that is, it should not dam- age the plant being tested, nor should it require any part of the plant to be removed from its habitat. Briefly, ic should be possible to obtain a print-sample from the plant, tag both the print-sample and the plant for field identification purposes, carrying out the identification protocol with minimal disturbance of the plant in its habitat. 4, The system must be simple and the methodology inherently economical. An identification protocol requiring extensive, expensive, detailed procedures, material, and equipment ts patently undesirable in countries where economic pressures are particularly acute and gov- ernment, private agencies, and individuals are all hard-pressed to fund conservation-ori- ented work. 5. Identification confirmations must be objective, not subjective. Identification predic- tions should be supported by statistical methodology not only by the personal opinion of individual A or B. 6. Lastly, and probably the greatest “acid test” is that the system must reasonably work in the “real-world” and constructively contribute co the body of botanical knowledge. What is sought is not merely a simplistic method of helping provide correct plant identifications, but one that contributes to a greater understanding of species and genera, and assists in their conservation and a more enlightened awareness and appreciation of their economic and ecological value by the general populace. The genus S¢ mmboped Was selected for several reasons. In nature, mem flower only for a few days, resulting in field identifications being a matter as one of the genera suggested bers of seeing the plant at the right time, and by Dr. Carl Withner—it appeared that a sufficient number of study speci- mens could be secured on a limited budget. The genus offers a particularly interesting challenge because, despite being represented throughout the tropics of the New World, it will not normally be encountered when in flower, thus presenting a practical group for the investigation of an identification confirmation system using means other than floral analyses. The use of fingerprinting by law enforcement agencies depends, in part, on having a sufficiently extensive file of known prints against which an “unknown” may be checked against the three general groups of arches, loops, and whorls (U.S. Dept. Justice 1984). Among other advantages, this system is non-invasive; doing no damage to the individual being printed. Although the print match may be attempted by mating prints from an object directly with those of an individual, the system can be effective by comparing prints from an object with those from an extensive file of known 684 Sipa 17(4) subjects. With this background from law enforcement work, particular note was taken of “leaf fingerprinting” used by students at the Universidad Autonoma de Nuevo Leén, the specific methodology of which was said by the professor to be unpublished work by him. However, just prior to re- turning the galley proofs of this manuscript, a publication by Petroski referencing one by E.M. Stoddard, was received from Dr. John Beckner of the Marie Selby Orchid Identification Center at Sarasota, Florida. Petroski (1965), outlines a similar method of leafprinting of orchid leaves using cellulose acetate (clear fingernail polish), with che resulting dried cellulose acetate film removed by forceps and dry-mounted on a microscope slide. This work's basic fingerprinting technique does not greatly differ from that of Petroski (1965) and Stoddard (1965), although the statistical analyses of the cell measurements developed by one of our number (Ferry) offers a new approach, objective in nature, to the identification of species. In addition, Scoddard’s work with alfalfa, chrysanthemums, and marigolds infers the usefulness of this system to other plant families. METHODS AND MATERIALS Clean white styrofoam “popcorn” is dissolved in xylol until the liquid is about the consistancy of warm syrup. This is applied to a clean leaf surface over an area of +2 X 5 cm, drying in two or three minutes. A short strip of clear transparent tape is pressed evenly and firmly over the film, but not with enough pressure to damage the leaf. The tape is peeled from the leaf, gently pressed onto a glass slide, and examined with a compound micro- scope. If it is desired to retain the slide permanently, a thin glass coverslip may be applied with its longitudinal edges taped to ensure prince flac. — rolding the leat The slide should be marked as to which leaf surface, adaxial or abaxial, was printed. Using a felt writer or other marking pen, a small “H” or an ” (haz: Spanish for adaxial, or envéz for the abaxial or underside of the leaf) is normally marked. The letter is followed by six digits to indicate the date, always as day-month-year (e.g. HO20496/7 = Haz; 02 April, 1996/ the seventh specimen done on that date). This writing is small and done where it can be removed when the slide’s permanent label is placed. In the {Bz a small plant tag gently tied to the plane with the same set of numbers (H&E020496/7) penciled on both sides or imprinted with a stylus. It is imperative that the location be clearly stated on either the envelope or a card within the envelope for relocating the plant at a later date! In the laboratory, the slide is photographed at 80X magnification. The microscope used in this research is a Microscoptics compound microscope with a trinocular head on which is mounted a Nikon HEM photo system. held, the slide is now placed in a slide box or small envelope, anc i eRRY, ET AL., Leaf-print analyses for plane identification 685 An indexing lens inserted in the field lens assembly of the photosystem prints index marks on each photomicrograph enabling measurements to be taken from the print. A Reichert-Jung micrometer slide of 2 mm divided into units of 0.01 mm is used to establish the lens correction factor for each magnification capability of the microscope. Black and white or color film may be used, and processed privately or commercially. In this study, Kodacolor film was used to make 10 X 15 cm (4 x 6 inches) prints. For color transparencies, Kodak Tungsten film has been the film of choice. From the photographs, 25 each of adaxial cells, abaxial cells, guard cells (both as a unit) and subsidiary cells are measured using a set of calipers to measure average lengths and widths. The number of trichomes (ad axially and abaxially) shown on each photomicrograph ts noted on the specimen’s data sheet, as are the number of stomata. A Macintosh SE/30 computer with 5 mb of RAM and 80 mb of internal memory was initially employed, augmented later by a Power Tower Pro 225 computer with 128 mb ram and 2 gb of internal memory. A Microsoft Excel spreadsheet combined each raw measurement with the lens correc- tion factor, entering measurements on the data sheet in microns (Lt) and combining them to present the total, mean, and standard deviation for each set of measurements. The statistical treatment used was an analysis of variance (ANOVA) with p = .05. The number of cells per square millime- ter was computed and printed on the specimen’s data sheet. Totals from each individual data sheet are linked to provide a combined worksheet from which data totals are analyzed, and individual and groups of Gaussian curves can be obtained on printouts. RESULTS Data have been collected from fifty Stanbopea specimens, 32 of which are different species (multiple samples of some), and two man-made primary hybrids. Data have also been collected from plants of Goventa utriculata (Sw.) Lindley, Govenza superba (Llave & Lexara) Lindley ex Loddiges, Malaxis corymbosa (S. Watson) Kuntze, and Malaxis macrostachya (Lexara) Kuntze in the Sierra Madre range southwest of Monterrey, Mexico. Subsequent check- ing indicates no damage done to any field or greenhouse plant from which leaf-prints have been taken. A Stanhopea plant received as an “unknown” on 29 April, 1996 was num- bered C26 and data taken from it were compared with that from known plants. Comparisons of the mean of its adaxial cell areas with known spe- cies (r = .05) indicated it as S. t#grina. On 20 June, 1996 another unknown arrived and was tagged C33. The data inferred that it coo was S. tigrina. On O08 July plant C26 flowered, confirming the prediction made on the basis of the statistical data from the leaf print. This provided the first example of 686 Sipa 17(4) 5.0000 () LOOO.00 Fic. 1. Gaussian curves of adaxial cell areas of §. figrina specimens C26, C33, C59, and CO7 the successful prediction of the plant's identity by use of this methodology prior co seeing it in flower. A few days lacer, despite the destruction of plane C33 bya resident macaw, analysis of the remains of a not-fully-opened flower confirmed that its predicted identification had also been correct. Gaussian curves are presented for plants C26 and C33 and the two confirmed S. tigrina plants (CO7 and C59) in Figure 1. Although the normal curve for specimen C26 was more leprokurtic (higher “crested”) than the others, the adaxial means of all specimens did not significantly differ (ANOVA, p =.05). Amplified data for the four Stanhopea tigrina specimens is shown in Table 1. Stanhopea Chocolate Chips = 8. tigrina X panamensis by D. Pulley, 199] (Fisher 1994). Data from specimen C56, Chocolate Chips ‘Linde’ was com- pared with that from CO7 S. tigrina, C59 S. tigrina ‘Glory of Mexico,’ and C58 S. panamensis, all of which were supplied by Dr. Douglas Pulley of Los Gatos, California. While the exact S. f/grina parent of this hybrid was un- known by this worker, all plants had been received from the same grower- hybridizer, and che assumption was that one of the two 8. fgrina plants was one parent and the S$, pavamensiy plant the other. Therefore both S. tigrina plants are included in the chart of the Gaussian curves and Table 2 gives a summary of the data for the four specimens. DISCUSSION ‘Oz, axial cell area means of the hybrid and parents (Table 2) infers that $. Chocolate Chips ‘Lindt’ is vegetatively more closely allied Analysis of the ac with the S. “7grina parent than with S. panamensis. This is borne out pictori- ally by the position and shape of the curves (Figure 2), and abaxial epidermal cells indicate a similar relationship. However, correlation attempts using — stomata guard and subsidiary cells have been inconsistant in and between TABL idch; SSarea: area, subsidiary cells; SSI/w: 1. Statistical data: Four specimens of S. tigrina (CO7, C26, C33, & € are means derived from data collections. Ad: adaxial; StcdDev: standard een Ab: abaxial; Aad/Aab subsidiary cells length/width. Specimen C59 is S, ae ‘Glory of Mexico.’ irea, adaxial/area, abaxial; GCI/w: guard cells Note that areas and numbers of cells length/ wi C# Ad Area Ad#Cells AdStdDev = AbArea AbStdDev — Aad/Aab GCArea GCl/w SSarea SSI /w Q7 4259 234.79 852.13 2857.66 700.88 1.49 19972 1.42 2126.08 1.50 26 1220 236.94 805.00 2856.17 734.26 1.48 1128.68 1.25 2159.94 1.30 Te) 4234 236.21 869.12 2816.88 815.48 1.50 1537.7 1.34 1483.57 1.25 59 4245 235.5 836.57 2917.18 639.82 1.46 1817.58 1.48 3745.29 1.25 Tate 2. Statistical data: CO7 S. tigrina, C59 S. tigrina ‘Glory of Mexico,’ . Chocolate Chips ‘Lindt,’ & C58 S. ehameha Ad: adaxial; Std Dev: standard deviation; Ab: abaxial; Aad/Aab: area, adaxial/area, abaxial; GCI/w: guard calls a width; SSarea: area, subsidiary cells; SSI/w: subsidiary cells lengch/width. C# Ad Area Ad#Cells — AdStdDev AbArea AbStdDev = Aad/Aab GCArea GCliw SSarea SSI/w O7 1259 234.79 852.13 $57.66 700.88 1.49 919.92 1.42 2126.0 1.50 39 4245 2357 836.57 2917.18 639.82 1.46 1817.58 1.48 are 1.25 56 1095 244.19 850.31 2828.01 622.69 1.45 1566.14 1.30 3477.2 1.39 38 724 268.50 822.46 2657.20 392.69 1.40 1541.51 1.32 1484.26 1.58 “TY LA SAUNA ra uonvoyiuapr quvyzd roy sasdpeue dursid-yeoT L89 O88 Stipa 17(4) 5.0000 1000.00 6000.00 7 Fic 2. Gaussian curves of adaxial cell areas of C58 S. panamensis, C56 $. Chocolate Chips ‘Lindt,’ and S. trerina specimens C59, and C07. species as well as between species and their hybrids. More trials linking leaf cell data with proven floral qualities are indicated. If parent-offspring curves of adaxial cell areas or areas of other external organs can be corre- lated with specific Horal qualities sought by the hybridizer, these results could be of economic value as a predictor of floral characteristics while still in the seedling stage and could afford the grower improved quality control over seedling crops. With regard to taxonomic avenues, it is hypothesized that the statistical relationship established by DNA sequencing will not significantly differ from that arrived at by this methodology. If this proves to be the case with botanical specimens, a vast array of possibilities may be applicable cto other life forms. The identification of these unknowns offers encouragement for the greater use of this leaf-print-statistical analysis methodology for con firming plant identities in nature, and preliminary field studies lend support to the use of this methodology. Adapted for field use, population surveys appear to be possible without disrupting the ecosystem, while affording humans the option of hand-pollinating specifically identified field plants, thus assist- ing in reviving a low plant population without disturbing its members. In the field, this system could be of use in confirming the identity of both species and hybrids and provide the number of each in specific areas. Grant's work (1981) wich the genus Gilia in California provides strong evidence that—with sufficient time and the invasion and adaptation of a species into new habitats a species can vary sufficiently over its range to provide new fixed gene combinations resulting in a new species. Grant’s basic data was originally published in 1971 with the 1981 edition provid- ing refinements, corrections, additional data, and preliminary results of unpublished clinical work with specimens of S. saccata and S. radijosa ap- Ferry, ET AL., Leaf-print analyses for plant identification 689 pear to provide confirmation for his work. In light of the work of Dodson (1963) and of Kennedy (1975), it is suggested that field identification work using this statistical methodology over the range of the Stavhopea radtosa- saccata complex would provide useful data for clarification of this specta- tion phenomenon and assist in establishing specific points of variation at precise locations over the geographical range of the two species. In conclusion, this is an objective approach to plant identification confirmation relying on data capable of statistical verification. In outlining the six requirements listed earlier in this paper, the attempt has been to detail a system workable within the realities of governmental attitudes, and the conservation desires of the scientist; yet one capable of being easily understood and supported politically and economically by government agencies and local people within the countries concerned, — ACKNOWLEDGMENTS We wish to thank Eric Olsen for his computer expertise. His help greatly assisted data calculations and provided for the display of Gaussian curves. Thanks are also due Dr. Douglas Pulley for providing specific research speci- mens as well as constructive comments during the course of this work. We also thank Dr. Dwight T. Kincaid for reviewing this manuscript and pro- viding pertinent changes to it. REFERENCES Ames, O., and D.S. CorreLt. 1952. Orchids of Guatemala. Publ. Field Mus. Nat. Hisc., Bor. ser. 26:1—395 ~ and D.S. Corrett. 1953. Orchids of Guatemala. Publ. Field Mus. Nat. Hist., Bot. ser. 26:1—727 Arpirti, J. 1992. Fiandsmentals of Orchid Biology. John Wiley & Sons, New York. nia L, H., P. Crips, and E. Launerr. 1992. The manual of cultivated orcuid species. MIT Press, Cambridge, M Curry, K.J., WL. Stern, and L.M. McDowet.. 1988. Osmophore development in Stanhopea I anfracta and S. pulla. (Orchidaceae). Lindleyana 3:212—220. Curtis ee 1910. Orchids for everyone. J.M.Dent & Sons, Ltd., London. Cuter, D.F. 1978. Applied plant anatomy. Longman, London and New York. Dopson, me H. oan The Mexican Stanhopeas. Amer. Orch. Soc. Bull. 32:115-129, 1975. Clarification of some nomenclature in the genus Stanhopea ~ {Om nidacese. oe 1:46-55. Dressler, R. 1993. Phylogeny and classification of the orchid family. Dioscorides Press, Portland, OR. Fisner, J.R. 1994. Bishop's interim list of orchid hybrids registered during 1991-1993. 241 Hillside Circle SW, Vienna, Virginia. Grant, V. 1981. Plant speciation. (2nd ed.) Columbia University Press, New York. Hamer, FE. 1974. Las orduideas de El Salvador. Ministerio de Educacion, San Salvador, E] Salvador HAWKES, A. D. 1965. Encyclopedia of cultivated orchids. Faber and Faber Ltd., London. 690 Sipa 17(4) Jenny, R, ae The genus Stanbopea. Amer. Orch. Soc. Bull. 62:1270—1277. KENNEDY, 1974. Stanhopea fregeana Rchb. f. discovered north of San Antonio, Texas. Onhia oe 38:89, - 1975. The Stanhopeas of Mexico. Orchid Digest 39:178-18 Pansr, G.F.J. and Fo Dunes. 1977. Orchidaceae Brasiliensis. Band II. Brive Verlag Hildescheim. PETROSKI, ‘ J. 1965. Colonal identification by means of leaf epidermal characters. Orchidata 5-191_193 STERN, WL aad M.W. Morris. 1992. V Veg tative anatomy of Stanbiped | (Orchidaceae) with ial reference to pseudobulb water- -storage cells. Lindleyana 7:34—53 SroppaKp, E. M. 1965. Identifying plants by leaf 9) epidermal character: rs. Connecticut Agric. 1984. The science of fingerprints, (Federal Bureau of Investiga- tion.). ULS. — — Olfc., . The coe oe Pies fae: Ceiba 2:1—321. Wittiams, N.H., ian WOM. Warren. 1988. Stanbopea panamensis, a NEW § tral Panama (Orchidaceae). Lindleyana 3:9-1 1. 2 < > Ks ioe pas ~ Y species from Cen- SUGGESTED MODIFICATIONS OF THE SALISBURY STOMATA INDEX DEVISED FROM A STUDY OF STANHOPEA (ORCHIDACEAE) R.J. FERRY SR., R. FOROUGHBAKHCH, Teh HAUAD, H. GAMEZ, J.V. STAR, 8. CONTRERAS, and M.H. BADII Facultad de Ciencias Biolégicas Universidad Antonoma de Nuevo Leon, A.P. F-2 San Nicolas de los Garza, N.L., MEXICO 66450 ABSTRACT Calculations of the stomata index have resulted in wide variances in indices both single and between plants. Modification of the formula is suggested, including a metl of more accurately estimating stomata and subsidiary cells, with ae — of be checked by the use of an identity involving adaxial and abaxial leaf surface RESUMEN -os calculos del indice estomadtico dieron variaciones amplias, tanto en un indivic como entre diferentes plantas. Se sugiere la modificacion de la formula, para incluir método de cdlculo mds seguro de estimacion de estomas y células subsidiarias, con t precisién capaz de ser comprobada por el uso de una identidad que abarque las supertic del haz y el envés de las hoyas. INTRODUCTION When initiating our investigation of the role of stomata in water loss leaves of members of the genus Sfan/opea (Orchidaceae) we approached 1 problem by initially attempting to determine the stomatal density 11 given area of leaf surface. However, results were inconclusive despite tempts to ameliorate variations by recording stomatal frequency in ter of the stomata to the proportion of epidermal cells as suggested by St (1965). Sinclair (1990) expressed preference for the use of the scomata dex (SI) of Salisbury (1928) over the mere determination of stomata d. sity. This index is arrived at by adding the number of epidermal cells to number of stomata, dividing this into the number of stomata, and mu plying the result by 100. Stace (1965) as well as Metcalf and Chalk (1979), working with dico ledonous material, found this index to be quite variable both in sin plants and between plants. The rationale given for this variability was t although a leaf developing under water stress retains the total number: epidermal cells, the cells are smaller, with the result that stomata densit ‘ncreased in the stressed leaf. Conversely, Rowson (1943 a, b, 1946), ci IDA 17(4): 691-696. 1997 69? Sipa 17(4) in Metcalf and Chalk (1979), working with the genus Cassia (Fabaceae), concluded that species of the same genus may be differentiated by means of the stomatal index. Rowson also found that the stomatal index: (a) did not vary significantly at different positions upon the leaf surface; (b) was inde- pendent of leaf size and habitat; and (c) was the same for different varieties within a species. Working with monocotyledons, Singh and Singh (1974) report an SI range of 2.0—17.0 for seven species of epiphytic orchids, and Boros (1980) reports 2.5—30.3 for 1 species of terrestrial orchids. Rasmussen (1987) mentions Stanhope only casually, stacing that a spe- cies (S, f7gria) was included ina summary by Sctrasburger (1866-67). Simi- larly, reviewing water relations in orchids, Sinclair notes the genus once as a reference by Link (1849) who saw spirally thickened idioblasts in roots of Stanhopea ebornea (Link's spelling = ebuvrnea). Other than these casual refer- ences, data is scant Congerniag Stanhopea stomatal complexes. During normal development, members of this genus receive copious water. Thus, they should exhibit stomata indices not greatly different from one another. However, use of the Salisbury formula yields inconclusive data when calculating a baseline stomata index for individual species (see ST eels: Table 1). With wide ranges reported by previous workers and the minimal date regarding this genus, a closer look at the formula was indicated. The stomata index formula assumes that the leaf surface displays epider- mal cells and stomata within a given area, and given the material with which Salisbury worked, this assumption is both obvious and quite correct. How- ever, the literature is unclear regarding whether certain epidermal cells (tri- chomes and their flanking cells) are excluded, included as a unit, or counted as individual entities. Additionally, the literature is unclear whether sto- matal subsidiary cells are counted separately or included as an entity of the stomatal complex. Thus, applying this as a general formula may not allow for other organs on the leaf surface, and results may be correct or merely the product of a mathematical error. In Stanhope plants trichomes are present on both adaxial and abaxial leaf surfaces, with stomata on y on the abaxial surface with the stomata are flanked by paracytic subsidiary cells. Thus, in order to arrive at a more accurate measure of the stomata index, the leaf surface space taken by these epidermal organs should be considered. How- ever, counting the number of stomata within a given area leads to esti- mates when several of these organs overlap the borders of the selected area, and thus inaccuracies can be incroduced in the number of stomata actually represented in that area of the leaf surface. Additionally, the basic formula does not clearly consider variances in the sizes of subsidiary cells. — ae METHODS AND MATERIALS Photomicrographs of the leaf surface were taken and a random sample (N = 25) of adaxial cells, abaxial cells, stomata guard cells, and individual Ferry, eT AL., Modifications of the Salisbury Stomata Index 693 subsidiary cells was measured and the mean of each is shown in the table. Although the number of cells varies, the adaxial leaf area is equal to its abaxial area. Hence this identity can be expressed, using areas instead of numbers of cells, as shown in formula (1): (1) Adces + Aderichomes = Abceris + Aberichomes + Stomaca + Subsidiary Cells The number of trichomes in these Stanhopea plants number only one or two in a standard area, resulting in a negligible difference in the adaxial and abaxial areas. Thus their effect has been assumed as equal and they are dropped from these calculations. However, where trichomes are numerous or adaxial and abaxial quantities differ greatly, their numbers and areas should be included in calculations. With this modification, the identity 1s expressed as in formula (2). (2) Adcetis area = ADeetls area + StOMAtAgrea + Subsidiary Cellsarea Using the random sample of 25 epidermal cells, another modification 1s necessary to arrive at an accurate number of stomata complexes. As each set of Stanhopea guard cells is supported by two paracytic subsidiary cells, the identity becomes as shown in formula (3). (3) 25 (Adcetis area) = 25 (AD celts area) + & [Stomatagrea + 2 (Subsidiary Cellsarea) I. Entering numerical values (e.g. CHOI, S. saccata) gives: (3a) 25 (3097.40) = 25 (2219.28) + X [1476.94 + 2 (2024.48). Thus the number of stomata (X) within the specified area = 3.973. Inserting this value into the modified Salisbury formula gives the following: Sie Ss. #HStLareaSt+ 2(areaSS)} 3.97 3[1476.94+4 2(2024.48)} _ 21.954 E4+S E+ #StareaSt+ 2areaS$)} 25(2219.28)+3.973{1476.94+42(2024 48) 77436.4 rh and Ly area ee Merely counting the number of stomata complexes within a specified area introduced inaccuracies due to variances in guard cell sizes, as well as stomata overlapping area boundary lines, and exclusion of the areas of sub- sidiary cells. These counts produced false stomata indices as shown in the column headed SI Cell, and thus showed greater differences between plants of the same species as well as of different species of the same genus. Using the number of abaxial cells per square millimeter (e.g. COL S. saccata = 45().6), and the number of stomata (3.973) in the classic formula gives the index as shown in formula (4): (4) st=_ 5 _ «100 =__3-9? x100 = 0.874 E+S 3 450.6 + 3.973 694 Stipa 17(4) RESULTS ABLE i adaxial cell surface area; Ab Cells: abaxial cells p/sq mm; # Scoma: mean number of stoma p/sq mm, GC Area: mean area of guard cells; SS Area: mean subsidiary cell area; $I Cell: index per Salisbury I. Summary of 30 Stawhopea species. Ad Area: mean adaxial cell surface area: Ab Area: mean formula; SI Area: index per modified formula C# Species Ad Area Ab Area Ab Cells #Stoma GC Area SS Area SI Cell SI Area OL saccata 3097.40 2219.28 450.60 3.973 1476.94 2024.48 574 284 O2 saccata 3040.90 2219.09 450.64 3.715 1264.14 2133.04 B17) 270 O38 mmsienis 4769.92 3247.80 307.90 6.567 1993.45 1900.39) 2.088 319 O4 hernandezii 2782.83 2096.11 477.07 1.202 1291.47 1397.28 873 247 OS ocuwlata (123.24 2574.44 388.43 9.18 1476.80 IS81.22) 3.055 376 06 oculata 4173.52 2535.24 394.44 8.50 1554.22 1632.02 1] 93 O7 tHerind 1259.11 2857.66 349.94 6.774 919.92 2126.08 1.899 329 O8 eraveolens 3380.38 2372.18 121.55 3.698 1467.43 2859.7 870 298 09 eraveolens 3288.62 2309.55 132.98 5.93 1518.74 1400.99 1.35] Z LO martiana 3744.41 2700.86 370.25 5.837 1215.20 1627.29 1.552 288 L 1 sntermedita 2694.32 2060.25 185.38 2.888 1318.93 2085.25 59] 23), 12 wardi 1603.10 3546.56 281.96 1.796 1302.88 2100.33 1.673.230 13 wardti 1594.98 3526.38 283.58 6.004 1568.32 1440.43 2.073 233 14 ward 1660.02 3570.86 280.04 3.092. 1343.93 2001.75 1.786 234 LS nigrovtolacea 3129.27 2263.86 41.72 1.982 1137.87 1602.38) 1.115 277 16 snsients (780.59 3251.88 307.51 7.176 1287.34 2019.38 2.290 319 17 costaricensis 3605.70 = 2648.90 377.52 1412 1427.67 1997.20 1.155 205 18 pozoi 1S91.37 2991.96 334.23 10.542 1520.17 1492.10 3.058 388 19 shuttleworthii 4377.18 2808.95 356.01 7.304 1334.66 2016.36 2.010 358 20 fricornis 4532.35 3555.51 281.25 5.077 1181.80 1813.5 773.216 21 rerwchenbachiana 5623.69 3766.63 265.49 11.819 1337 21 1295.46 4.262 330 22 ¢ Me 1498.03 3726.32 268.36 3.157 1419.09 2346.20 1.163 172 23 ecornut 3837.05 2756.76 362.75 1.838 1450.68 2065.54 1.316 282 24 pentschiana 1368.36 2899.4] 14.90 5.508 1360.17 2653.5 1.572 336 25 grandiflora (816.54 3714.49 269,22 8.525 1222.8] 1004.59 3.069 229 26 trerin 220.15 2856.17 350.12 6.258 1128.68 2159.94 | ) 323 27 candida 1856.62 2790.16 358.40 11.420 1355.76. 1584.00. 3.088 425 28 connata 4713.00 3213.80 311.16 6.833 1477.35 2003.84 2,146 318 29 haselowanta 1215.97 3556.07 281.21 2.295 1471.04 2858.26 S10 157 30 rmpressa 4307.50 3119.53 320.56 10.26 1072.09 1228.61 3.101 .276 DISCUSSION Although the area method involves more calculations, it provides a more accurate indication of the stomata and subsidiary cells within a given leaf surface area. While the number of cells varies on both the adaxial and abaxial leaf surfaces, obviously the areas of the leaf surfaces should be equal as should the sums of the areas of the organs on each surface. The literature is silent concerning subsidiary cells in calculations of the stomata index, but these cells occupy significant areas of the abaxial leaf surface in Stanhopea as well as other monocotyledons. In Stahopea, stomata and subsidiary cells occur only on the abaxial leaf surface, with the guard cells of all species surveyed accompanied by subsidiary cells of varying areas. However, this does not FERRY, ET AL., Modifications of the Salisbury Stomata Index 695 hold true for all other plants or even all members of the Orchidaceae, and particular attention should be paid to modification of the general formula in order to properly reflect the particular physiological condition of the plants being analyzed. Previous workers have reported wide ranges of the stomata index in and between plants of the same species. However, in this survey of 50% of this genus the stomata index is not significantly different for plants of the same species, nor is a wide range observed between different species. It is sug- gested that calculations of the stomata index include subsidiary cells and any other types included in the count, as well as noting them as such. It is also suggested that the stomata index formula be modified to consider the areas of all components rather than being calculated by merely a count of unspecified epidermal cells and only the guard cells of stomata. on) REFERENCES Ayensu, E.S. and N.H. Wiuiams. 1972. Leaf anatomy of Palumbina and Odontoglossum sub cenu: Lee Amer. Orchid Soc. Bull. 41:687—696. Borsos, O. 1980. Anatomy of wild orchids in Hungary. I. Tissue structure of f leaf and floral axis. Acta Agronomica Acad. Sci. Hung. 29:369-389. Dressier, R.L. 1981. The orchids: natural history and classification. Harvard University Press, Cambridge, M Ss 1D: Phylogeny and classification of the orchid family. Dioscorides Press, epene: OR KENNEDY, G.C. 1974. Stanhopea fregeana Rchb. f. discovered north of San Antonio, Texas. Orv {Dige 38:89, 95. Link, H. 1849. Bermerkungen iiber den bau der orchideen. Zweite Abh. 117-127 + plates. Pl ae eee Abhandlungen der Kénigliche Akademie der Wissenschaften zu Berlin Metca.rr, C.R. and L. CHALK. 1979. Anatomy of the dicotyledons. Vol. I. 2nd. ed. Clarendon Press, Oxford. Passt, G.FJ. and BF. Dunes. 1977. Orchidaceae Brasiliensis. Band I]. Hildescheim: Briike- Verlag. see sseN, H. 1987. Orchid stomata structure, differentiation, function, and phylogeny. . Ardicti, - Orchid biology: reviews and perspectives, IV. Cornell Univ. Press, a NY. Pp. 105-138. Rowson, J.M. ee , 1946. In: Metcalfe, C.R. and eee 1979. Anatomy of the dicotyledons. Vol. I. 2nd. ed. Clarendon Press, Oxford. P. 1 SaLisBuRY, E.J. 1928. On the causes and cree significance a stomatal frequency, with > special reference to the woodland flora. Phil. rans., Ser. B, 216:1—65 SINCLAIR, R. 1990. Water relations in ae AL). ten ed. Ori biology: reviews and perspectives, V. Timber Press Inc al OR. Pp. 63-11 SINGH, V. and H. Sincu. 1974. cere of stomatal comq ees in some Orchidaceae. Curr. Sci. 43:490-491. Stace, C.A. 19 ie Cuticular studies as an aid co plant taxonomy. Bull. Brit. Mus. (Nat. Hist.) Bot. STRASBURGER, : 1866- 67. Ein Beitragzur Encwicklungsgeschichte der Spaltéffnungen. Jahrb. Wiss. Bot. 5:297—342 BOOK REVIEW BERLIN, ELots ANN and Brent BERLIN. 1996. Medical Ethnobiology of the Highland Maya of Chiapas, Mexico. (0-691—03741-8, cloth). Princeton University Press, Princeton, N.J. $85.00. 557 pp. — The Tzeleal and Tzotzil Maya in the highlands of Chiapas, Mexico have been in the fore- front of paradigm development as the interdisci ay field of ethnobiology matures. Contributions to understanding the principles of classifi nand taxonomy made through the systematic study of these indigenous peoples has ey accomplished more than all systematic lists of useful plants and animals published since Castetter defined the field. The authors and collaborators document a system of traditional knowledge based on years of explicit empirical experimentation about the effects of herbal remedies on body func- tion. The authors provide an ethnomedical characterization of gastrointestinal conditions recognized by these highland Maya and describe existing evidence on the phar spent activity of thirty-four medicinal plant species used to treat gastrointestinal diseases y the highland Maya. The authors claim that existing evidence on the pharmacological oe of plane species used in the treatment of gastrointestinal ailments of the Highland Maya supports the notion that these indigenous people have a scientific basis for traditional medicine possessing well-developed therapeutic efficac ithors have gone to great lengchs to As ciareat the cultural understanding of illness including cause, treatment and prognosis and is surely che most thoroughly re- searched ethnomedical characterization of gastrointestinal illness to date. The thorough- ness of ethnomedical research on each of three major gastrointestinal conditions—diar- rheas, abdominal pains and worms—is complemented by description of the documented biological activities of the principal medicinal plants used to treat each gastrointestinal affliction. Details of known biological activities of thirty-eight species comprising the “basic medicine kit” (el cuadro basico) is supplemented with an evaluation of an additional fifty-seven species used in admixtures with the basic medicine kit. Evaluation of a subset of the plant species used in admixcures further support to the idea chat these indigenous people possess a sophisticated understanding of complementary or intensifying nature of biological activities of the admixtures used in treating gastrointestinal conditions among the highland Maya. This book marks the maturation of ethnobiological ing uiry in which attention 1s fo- cused as much on demonstrating the scientific basis of indigenous health care as on docu- menting efficacy of traditional treatment of illness. In the context of growing interest in self-determination among indigenous groups and sovereign rights over knowledge about use of biological diversity, this books goes further than any other in demonstrating how academic research can also satisfy local community interests and needs. This information- packed volume should be on the bookshelf of all who would consider biological prospect- ing in forests possessed by indigenous societies because it provides a benchmark method that can help to empower local communities to assume responsibility in the conservation of biological and cultural heritage.—Brace Benz. Sipa 17(4): 696, 1997 A SYNOPSIS OF THE GENUS GENTLEA (MYRSINACEAE) AND A KEY TO THE GENERA OF MYRSINACEAE IN MESOAMERICA JON M. RICKETSON Mussouri Botanical Garden -O. Box 299 St. Louis, MO 63166-0299, U.S.A. pricketson@ lehmann. mobot ore JOIN), PIPOLY Ii Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4060, U.S.A. ppipoly @brit, org ABSTRACT An updated description of the genus Gevt/ea and its taxonomic synonyms is provided, along with a key and a nomenclatural summary for its nine species. Nine binomials are newly relegated to synonymy, and three taxa are excluded. Heberdenia Banks, once thought to occur in the Neotropics, is shown to be eels a known only from the Canary Islands, necessitating the new combination G / I:flora (A. DC.) Ricketson & Pipoly. To facilirate identification of members of the genus es a key to the genera of Mesoamerican Myrsinaceae is also provided. RESUMEN Se aporta un resumen nomenclatural para las nueve especies del género G tlea. Se incluye una descripcién actualizada del género con una clave para separar las especies. Se comprobé que el género Heberdenia es monotipico, y restringido a las Islas Canarias, resulcando la nueva combinacion Gentlea enduliflora (A. DC.) Ricketson & Pipoly. Para facilitar la identificacién del género, se ofrece una clave para separar los géneros mesoamericanos de la familia Myrsinaceae. También se ofrece una clave para todos los taxa del género Gentlea, se enumeran sin6énimos tanto nomenclaturales como taxonémicos. Se relegan nueve binomios a la sinonimia, y se excluyen tres taxa del género INTRODUCTION The neotropical genus Gentlea Lundell, as here circumscribed, comprises nine species, ranging from Mexico to Peru. The genus occurs primarily in upper cloud and elfin forest formations, and is defined by long exerted stamens with minute dorsifixed cordate anthers, an elongated style, and terminal inflorescences that are short-pedunculate or subsessile, and when paniculate, generally broader than long. The genus has been confused with Stipa 17(4): 697-707. 1997 698 Sipa 17(4) Cybianthus and with Ardisia, but given the confusion surrounding generic delimitation in the family, we present a key to all Mesoamerican genera below. KEY TO GENERA OF MYRSINACEAE IN MESOAMERICA Calyx closed in bud, irregularly opening into 2-8 lobes within the same inflorescence Geissanthus 1. Calyx open in bud, ope ning into (3—)4—5(—6) lobes, the number of calyx lobes per flower consistent within one inflorescence. 2. Calyx and corolla lobe valvate in bud. 5. Flowers bisexual; inflorescences paniculate, longer than the petioles; branchlets tomentose with stellate or dendroid trichomes: calyx and corolla tomentose and glandular-papillose; anthers bright yellow; style elongate. Parathesis 3. Flowers unisexual; inflorescences vlomerulate, umbellate or rarely in reduced corymbs, much shorter than the petioles; branchlets glabrous or when tomentose, then with simple trichomes; calyx and corolla gla- brous or the corolla sometimes pense papilos: along the margin within; gin rs cream to > white; s style obsolete or reduced. Ces tecteaasle te ace ae Myrsine alyx and wiaeuncak ail | ] , IMbricate (alchough some times inconspicu- mr or contorted in bud. 1. Filaments connate basally to form a staminal tube adnate to the corolla tube 5. Inflorescence lateral (axillary); corolla tube gli andular-granulose at least at the junction of the tube and lobes . Cybianthus Inflorescence terminal; corolla tube glanc le ar- ‘uvenilent at Dase. es Synardisia 1. Filaments free from each other and free from or variously adnate to the corolla cube 6. Filaments adnate to the corolla; anthers less than 3 times longer ae wide. . Filaments exserted, more than 6 times longer than the anthers; ee of elfin and cloud fores Gentlea 7. Filaments included, shorter ne or as long as the anthers; planes of montane to lowland forest Stylogyne 6. Filaments free from corolla; joes at least 3 times longer than wide 8. Petals connate 1/4 their length; ovules uniseriate, the placenta- tion free-centra ieee .. Ctenardisia 8. Petals connate up to 1/5 their length; ovules pluriseriate, the placentation basa —. Ardisia In preparing a treatment of the genus for Flora Mesoamer wana, it became 5 necessary to assemble complete synonymies and ee oe references for Gentlea. Because of the somewhat abbreviated format of the Flora, the present synopsis is intended to provide a complete nomenclator for Genslea, which has been poorly understood. RICKETSON AND Prpoty, Gentlea in Mesoamerica 699 TAXONOMIC TREATMENT Gentlea Lundell, Wrightia 3:100. 1964. Type Species: Gentlea venosissima (Ruiz & Pavon) Lundell. eee Swartz subgen. Wa/leniopsis Mez in Engl., Pflanzenr. IV. 236 (Heft 9):77. 1902 Dioecious shrubs or small trees. Leaves alternate, petiolate. Inflorescences terminal, paniculate with corymbose branches, broader than long, short- pedunculate or subsessile, rarely umbellate or a reduced to a solitary flower; floral bracts thin, caducous. F/owers 4(—6)-merous, unisexual, pedicellate; sepals inconspicuously imbricate, connate basally or nearly free; petals con- nate 1/4—1/2 their length into tube, the lobes imbricate in bud, spreading and sometimes appearing valvate at anthesis; stamens exserted, the fila- ments long and slender, more than 6 times longer than the anthers, the anthers minute, cordate, dorsifixed, epunctate or inconspicuously glandu- lar-punctate dorsally; pistil ovoid or subglobose, the style slender, shorter or sometimes subequaling the corolla, the stigma punctiform, minute; ovules few to many, in 2 or more series. rit subglobose, one-seeded, exocarp thin, often drying costate. A small distinctive genus of nine species ranging from Jalisco, Mexico to Venezuela, Colombia and Peru. Members of the genus occur in upper cloud and elfin forest formations. Gertlea can be confused with Ardisia in fruit, except that the inflorescence of those with panicles is wider than long (rare in Ardisia), and the fruit is either inconspicuously or conspicuously costate (smooth in Ard/s/a). KEY TO SPECIES OF GENTLEA . Inflorescence een or rarely a solitary flower 1. G. penduliflora ; Eup paniculat 2. Leaf blades ‘lliptic or cso a lepidoce below, the margin slightly to deeply crenulate in the distal 1/2. 2. G. vatteri 2. Leaf blades oblanceolate, obovate or oblong, glabrous or with scattered trichomes below, of various shapes except lepidote scales, the margin en- tire to crenulate or serrate with teeth uniform throughout. ae Inflorescence rachis and pedicels glabrous. 4. Leaf margins entire 3. G. standleyi 4 Leaf margins crenulate to serrulate. 5. Leaf margins crenulate; petals translucent-lepidote abaxially. 4. G. crenulata . Leaf margins serrulate; petals glabrous abaxially. .............. 5. G. tacanensis 3, Inflorescence rachis and pedicels glabrate, with a few scattered furfu- raceous scales or simple, glandular trichomes, or tomentose with simple or dendroid trichomes 6. Inflorescence eich ae — tomentose, with a dense mat of simple or dendroid hair 7. Leaf blades face 0.5—1.5(-1.8) cm wide, inconspicuously 700 Stpa 17(4) pellucid-punctate, che aes and tertiary veins inconspicuous; sepals 0.8—1.2 mm lon 6. G. lancifolia . Leaf blades stiffly econ to subcoriaceous, (1 1.5—)2.0—6.0 cm wide, prominently black punctate eee i one and tertiary venation prominently raised; sepals 1.5—2 mm BS . G. venosissima penne: rachis and peduncle glabrate, with a few ae erect, scattered, simple or sessile glandular trichomes 8. a. with appressed glandular trichomes; sepals 1.2—1.8 mm long; petals 5—6 mm long; leaf blades 2—6(—8.5) cm long, 1-2 (—3.5) cm wide. 8. G. austinsmithit 8. Pedicels with eon erect, simple or glandular hairs; sepals 2—3 mm long; petals 3.5—4.2 mm long; leaf Gide: (7.5—)10—23 cm long, 2.5—)4.5-6.5 cm oo. 9. G. micranthera 1. Gentlea penduliflora (A. DC.) Pipoly & esa comb. nov. Myrsine eee les A. DC., Trans. Linn. Soc. London 17:1 10, 4. Heberdenia penduliflora ) Mez in Gaal, Pflanzenr. IV. 236 (Heft 9): ce 1902. Type: Calques Fi. en t. 739. 1874 (HoLoryPE: G, F! as photo- f neg. no. 30749) Upon review of the genus Gentlea, it became obvious that Heberdenia penduliflora could not be separated from it except for the inflorescence type. Further examination of the type of Heberdenia excelsa revealed that there were significant differences between that taxon and H. penduliflora. There- fore, transfer of Heberdenia penduliflora to Gentlea leaves Heberdenia a mono- typic genus known primarily from the Canary Islands. Heberdenia, as here circumscribed, is characterized by having included stamens with filaments adnate to the corolla 1/2 their length and dorsifixed, reflexed anthers: greenish petals thick and “leathery,” with few black punctations, and the adaxial surface densely covered with very short papillae. While Heberdenia excelsa superficially shares leaf and inflorescence characters with Gentlea penduliflora, the latter difters from che former in having exerted stamens with minute, erect, dorsifixed anthers and finally, membranaceous petals, with conspicuous reddish or black punctations, pink-white in color, and glabrous throughout. Despite the fact that Gentlea penduliflora differs from the rest of the spe- cies in the genus by its inflorescence of umbellate flowers or rarely a soli- tary flower, versus a paniculate inflorescence in other members of the ge- nus, but its exserted stamens with minute, dorsifixed anthers, elongated style, and costate fruit indicate it is well placed in Gevtlea. There are vari- ous examples of species in other genera with markedly reduced inflores- cence structures, such as Myrs/ne laetevirens (Mez) Arechay., the ea Myrsine with an abbreviated raceme, Sty/ogyne membranacea Pipoly, he only Stylogyne with two-flowered corymbs, Cybianthus fabiolae Pipoly, the only Cybianthus with one flower or a two-flowered raceme appearing umbelliform, and RICKETSON AND Pipoty, Gentlea in Mesoamerica 7O1 Crenardisia stenobotrya (Standley) Pipoly & Lundell, the only Crenardisia whose panicles bear flowers in umbels. [In attempting to locate the type of Gentlea penduliflora, study of the re- sults from the Royal Botanical Expedition to Mexico led by Don Martin de Sessé and José Mariano Mocifio between 1787 to 1804 was necessary. Re- sults from that expedition are well chronicled by Barkley (1945), Rickect (1947), Wilson (1962), Arias Divito (1968) and McVaugh (1977, 1980, 1982). The expedition collected specimens that are housed at MA and their duplicates were later distributed by Pavén. However the most outstanding contribution of the expedition was the commissioning of 2,000 watercolored drawings of Mexican plants and animals. Work on the collections and draw- ings was slowed after the return of Sessé, the principal investigator, to a Spain in political turmoil, and his subsequent death in 1809. Mocino gave Augustin Pyramus de Candolle a number of duplicate drawings and lent him his manuscripts and the original drawings in 1813 when he was forced to take asylum in France. Approximately 1,000 color “copies” were made of the original drawings before they were returned with Mocino to Spain. Unfortunately, the original drawings disappeared after the death of Mocino in 1819 and were not rediscovered until the 1970s when they were ac- quired by the Hunt Institute for Botanical Documentation in Pittsburgh, Pennsylvania. A significant number of new genera and species are based on the collections that were released by MA and the copies made by A.P. de Candolle between 1803 and 1874. Alphonse de Candolle (1874) distrib- uted 10 sets of tracings of the color copies of the original made by his father in 1817. Finally, two posthumous works of Sessé and Mocino, Plantae Novae Hispaniae and Flora Mexicana, were published between 1887 and 1894, nearly a century after the expedition occured. The above information is used to help illustrate the difficulty in track- ing down the type of Gentlea penduliflora. Alphonse de Candolle first de- scribed this taxon in 1834 based on “Icon. Mexic. ined.,” the Sessé and Mocino as In his treatment of the Myrsinaceae for the DC. Prodromus (1844), Alphonse lists only “Ex tc. ined. n. 739. fl. mexic.” It seems fairly clear that cies de Candolle based this name solely on the color copy in Geneva, since it appears that no actual collection 1s housed presently at G from that period and it is unlikely that he studied the Sessé and Mocifio Herbarium in Spain. Therefore, this is a holotype and no lectotypification is needed. The first planc collections at G do not appear until the 1840s. In addition, since the originals disappeared for over 150 —_— years and Alphonse de Candolle was only 10 years old when the originals were returned to Spain, it is unlikely that chey could have been used to describe the taxon 17 years later. Comparison of the original drawing at 702 Stipa 17(4) Hunt and the copy in Geneva are very similar with only slight, non-essen- tial differences, owing to differences in the portions painted. Thus, we se- lect the original color copy in Geneva (t. 739) as the lectotype. This draw- ing was photographed by the Field Museum of Natural History (Neg. No. ly distributed in many herbaria. Unfortunately, om 49) and photo's are wide the original drawing now housed in the Torner Collection at the Hunt Institute for Botanical Documentation at the Carnegie Mellon University in Pittsburgh, Pennsylvania, becomes nomenclaturally irrelevant. Distribution and ecology.—Gentlea penduliflora is infrequent in pine-oak forests on steep slopes and along montane watercourses, 1,400—2,900 m elevation. It is endemic to Mexico, in the states of Queretaro, Hidalgo, — Puebla, Veracruz, Oaxaca and Chiapas. 2. Gentlea vatteri (Stand. & Steyerm.) Lundell, — 3:102. 1964. Ardisia vattert Standl. & Steyerm., Publ. Field Mus. Nac 4 Bote pe 23220, 947. Ty GUATEMALA. Hurnuer : Cerro Lae oe Mimanhuitz and ae Cerro de los Cuchumatanes, |,500—2,600 m, 14 Jul 1942 (fl), J.-A. Steyermark 48558 (HoLoryPE: F!; isorype: US!). Distribution and ecology.—Gentlea vatteri is locally common in the wet cloud forests of Guatemala and El Salvador, at 1,500—3,300 m elevation. 3. Gentlea standleyi Lundell, Wrightia 4:69. 1968. Ardisia i Lundell) Lundell, Phycologia 61:67. 1986. Type: COSTA RICA. Herepta: Cerros de Zurqut, NE of San Isidro, 2,000—2,400 m (fr), 03 Mar 1926, PC. - Standley & J. ht S0613 (HOLOTYPE: US!; isorype: LL-TEX!). Ardisia minor Standl., J. _ Acad. Sei, ee 22.1927, nom. illeg. non King & Gamble, J. Astat. Soc. Bengal, Pr. at. Hist. 74:146. 1906. Syn. nov. Gentlea minor (Standl.) Lundell, W esas 3: 104. 1964, nom. ae Gentlea costaricensis Lundel , Wrightia 6O:115. L980. : COSTA RICA. Hrrepia: Cerro de Zurqui, NE of oe Isidro, ca. A » 2,200 m (fr), 03 a 1926, PC, caer & J. Valerio SO57 1 (HoLroryee: US!). Distribution and ecology.—Gentlea standleyi is endemic to Costa Rica, grow- ing 1n premontane wet forests, at 1,100—2,400 m elevation. The type of Ardisia minor was collected from the same locality, on the same day as was the type of Gentlea standleyi, and differs only slightly in appearance. Lundell failed to compare his Gentlea costaricensis to Gentlea standley?. 4. Gentlea crenulata Lundell, Wrightia 6:96. 1979. eae Lundell Phytologia 61:63. 1986, non Ardisia dae ita C. Lodd., Bot. Cab. 1817, Ardisia crenilata Vent., Choix Pl., 5 . 1803. Type: GUATEM vi en Me RAPAZ: Nino Perdido, bordering Rio San a 8 ieee N, in high foresct, on top of hill, May 1977 (fl, fr), C.L. Lundell & E. Contreras 21004 (HoLorypr: LL-TEX!; tsorypts: CAS!, LL-TEX! -2 sneets,). Because it is endemic to Guatemala and known Distribution and ecology. RICKETSON AND Prpoty, Gentlea in Mesoamerica 703 from the type collection, there is a dearth of information about its ecology. 5. Gentlea tacanensis (Lundell) Lundell, 3:105. 1964. Ardisia tacanensis Lundell, Contr. Univ. Michigan Herb. ¢ 1940. Type: MEXICO. Cutapas: side of Volcan de Tacanad, 2,800 m, 30 a ee (fl), E. Matuda S-226 (HOLO- , US!). type: MICH!; tsorypes: A!, F!, LL-TEX!, NY! Distribution and ecology.— Wet montane evergreen cloud forests, 1,400— 3,000 m elevation. Endemic to the area around Volcan Tacana on the Chiapas, México-San Marcos, Guatemala border and Volcan Tajumulco in Guatemala. 6. Gentlea lancifolia ee Phytologia 58:273. 1985. Ardisia dances Lundell, Phycologia 61 1986, non Ardisia inet he Merr., Philipp. J. Sci. 13:113 LOLS. Type: iG IntTIBUCA: between Calaveras and El ein pear lera Opalaca, 1,800 m, 12 Mar 1970 (fr), A "Molina R. & A.R. Molina 25547 (HOLO- type: F!; rsorype: US!). Distribution and ecology—Montane rain forest to elfin forest and cloud forests, 800—1,800 m elevation. Honduras and Nicaragua 7. Gentlea venosissima (Ruiz & oe ee ll, Wrightia 3:103. 1964. Caballeria venosissima Ruiz & Pavon, _ Fl. Peruv. Chil. 282. 1798. Mlyrsine venostssima (Rutz & Pavon) Spreng., coe st. Veg. 7 :664. 1825. Ardisia venosissima (Ruiz & Pavoén) J.F. Macbr., Fieldiana, Bot. ‘e 186. 1959. Type: PERU. HuANnuco: Mufia, MA, without elevation, 1878-1888, (fl, fr), H. Rutz & J. Pavin 5/37 (HOLotyPe: n.v., F Neg. 29508!). Ardisia propingua Kunth in H.B.K., Nov. Gen. Sp. 7:213 propingua (Kunth) Lundell, Wrightia 4:166. 1971. Type: VENE ripam fluminis Orinoci, without elevation or date ae rane & Bonpland s.n. (AO- LoTYPE: P-BON!; tsoryPe: P-BON, me as ee Nec. 38723!). 844. Tinus Ireviflora (A. DC.) Kuntze, Revis. — ay eee A. DC., Prodr. 8:122. {uANUcO: Muna, without elevation or date ee 1825. Syn. nov. Gentlea ZUELA. Crescit juxta 1891. Tye PERU. 7 pre tper G-BOIS, n.v. et as photo-F Neg. 852 4: isorypes: F!, MA, n.v.). Ardisia robinsonit Mez in Engl. ey oeee IV. 236 (Heft 9):77. 1902. Wy PE: VENEZU- n Tovar}, A. MIRANDA-ARAGUA: ee en Caracas and Colonia Tovar [prop 6,000 ft [1,829 m}, 26 Mar 1857 (fl), Fendler 2357 (HOLOTYPE: GH) Ardisia meiantha Donn. Smith, Bot. Gaz. 44 ie 15. 1907. Type: COSTA ‘A RICA, CARTAGO: Cartago, 1,500 19 i. 1906 (fl), WR. Maxon 44 (HoLtoryre: NY!; vicinity of ISOTYPES: ? SHEETS! ). St ph cies ria cages ene 7 Donn. Sm., Gi iz. 46: 113. 4 VeRAPAZ: Coban, ee. ) Lundell es 3:78. 1 Type: GUATE 1,350 m, Jun 1907 (fl), H. von aha 11814 (HOLOTYPI LL-TEX!, NY!, US!) Biase meridensis Steyermark, Fieldiana, Bot. 28:45: : fig. 95. 1953. Type: VENEZU- Ay: ae 1,820 m, 6 May 1944 (1), : Use. IsoTyPEs: C!, GH!, pA: between Mucuchachi and Canagua, ee VEN, n.v.) . Syn. nov. Ardisia jitotolana Lundell, MerRI JA. Steyermark 56325 (HOLOTYPE: F!; IsoT) Gore. tenuis Lundell, Wrightia 7:24. 1981 704 Stipa 17(A) Phytologia 61:65. 1986, non Ardisia tennis Lundell, Wrightia 4:149. 1970. Tyet MEXICO. Cutapas: Municipio de Rayén, in the Selva Negra 10 km above Rayén Meycalapa along road to Jitotol, 1,700 m, 12 Dec 1971 (fl), D.E. Breedlove 23144 (HOLOTYPE: Fly isoryprs: DS!, LL-TEX!, MO!, NY). Distribution and ecology.—Montane rain forests, evergreen cloud forests, premontane wet forests, 400—2,700 m. The most widely distributed spe- cies of the genus, Gentlea venosissina ranges from Oaxaca, Mexico south- ward through Mesoamerica to Venezuela, Colombia and Peru. It is expected, but has not as yet been collected, in Ecuador. In many broadly ranging species of Myrsinaceae, most of the taxonomic overdescription for this species is attributable to the use of combinations of quantitative features to define taxa and differences in the expression of quali- tative ones. Populations corresponding to the type of Ardisia propingua Kunth are notable only for their slightly thinner leaves and less prominent punc- tations. The type of Ardisia breviflora is notable only for the slightly more persistent tomentum of the apical buds and more highly branched inflores- cences. The type of Ardisia robinsonii represents populations from Colonia Tovar in Venezuela, with sepal apices slightly more acute, and petal punc- tations brown instead of black. The types of Ardisia meiantha and Stylogyne phaenostemona are notable only for their more subsessile inflorescences and acuminate leaf apices. Ardisia meridensis is quantitatively and qualitatively identical to the type of Gentlea venosissima. Populations corresponding to the type of Gentlea tenuis are notable only in their very chin and inconspicu- ously punctate sepals and petals. Another common problem is that the poe) species have been treated regionally, thus names are rarely compared over ¢ range as large as Mesoamerica. _ 8. Gentlea austinsmithii (Lundell) Lundell, Wrightia 4:68. 1968. dete austinsmithy Lundell, Contr. Univ. Michigan eth, 7:36. 1942. Typr: COSTA R LAJUELA: Region of Zarcero, Palmira, ca. 1,920 m, 9 Dec 1937 (fl), Austin er A673 (HOLOTYPE: MICH! isoryprs: F!, MO!). Gentlea molinae Lundell, Wrightia 4: —— 1970. Syn. nov. Ardista motinae — Lundell, Phytologia 61:65. 1986. oo RAS. La Paz: Montana Verde o Cordillera Guajiquiro, 1,900 m, 23 e ur eee oe A. Molina R. G ALR. Molina 24. 389 (HOLOTYPE: LL-TEX!; isorypes: F!, LL-TEX!, Gentlea parviflora ey Wrightia 5:89. 1975. Syn. nov. Ardisia parvissima Lundell, rytologia 61:66. 1986, non Ardisia parviflora Valboc, Syst. list Trees, Bombay ed. 2, 204. 1902. ve GUATEMALA. Baja Verapaz: Union Barrios, in high forest, top of hill, E of km 154, 11 Apr 1975 (fl), C.L. Lundell & E. Contreras 19212 as rypPe: LL-TEX!; isorypes: CAS!, LL-TEX!). ~ Distribution and ecology.—G entlea austinsmithii occurs in wet premontane, montane, elfin and cloud forests, and along watercourses in oak forests from Guatemala to Panama at 400—2,500 m elevation. The type of Gentle molinae Lundell, described in fruit, is notable only RICKETSON AND Pipoty, Gentlea in Mesoamerica 705 for its slightly longer petioles and more prominent pitting on the abaxial surface. Populations represented by the type of Gentlea parviflora difter from the So of populations of G. austinsmithir because of their more membranaceous leaves and slightly smaller flowers. 9. Gentlea micranthera(Donn.Sm.) Lundell, Wrightia 4:69. 1968. Parathesis aniciela Donn.Sm., Bot. Gaz. 18:205. 1893, non Ardisia micranthera Picard in : e, Fl. Gen. ae Chine 3:817. 1930. Ardista micrantha Donn.Sm., Bot. Gaz. ee “1889, nom. illeg., non Ardisia micrantha Kunth in H.B.K., Nov. Gen. Sp. 3:246. 1818. eae micrantha (Donn.Sm.) eee Weigh 3:107. 1964, nom. illeg. Ardisia staminosa Lundell, W eee 3:78. 1963 EMAL LTA VERAPAZ: mountain forest near Coban, 4,600 ft {1, 402 i ae 1888 (fl), H. von Tiirckheim 1365 (noLtotyre: US!; tsorypes: F!, G!, GH!, LL-TEX!, NY!) Gentlea mevanghii (Lundell) Lundell, Wrightia 3:106. 1964. Syn. nov. Ardisia arin Lundell, Wrightia 3:77. 1963. Type: MEXICO, JALisco: steep mountains S of Talpa de Allende, in the headwaters of a west branch of Rfo de Talpa, somes above a rapid clear stream, |,200-1,700 m, 18-19 Oct 1960 (fl), R. McVaugb 20396 (HOLOTYPE: MICH!; tsorype: LL-TEX!, MEXU!). Gentlea cuneifolia Lundell, Wrightia 7:2: 45. 1983. Syn. nov. Ardisia se a Lundell, Phytologia 61:63. 1986. Type: MEXICO. Veracruz: Municipio de San Andrés Tos cerca del aguaje en el lado S del Volcan San Martin ae ee ae 15 Feb 1972 (fr), J.H. Beaman & C. Alvarez del Castillo 5751 (HoLotTyPE: LL-TEX!; isoTyPEs: F!, MEXU!). Gentlea auriculata Lundell, Phytologia 58:273. 1985. Syn. nov. Ardista eee mises 1, Phy ye a 66. 1986, non Ardisia auriculata Donn.Sm Tyre: GU ALA. Zacapa: Sierra de Las Minas, mmidale a upper south- fac ing Lie of ate Gemelos, 2,100—3,200 m, 26 Jan 1942 (fl, fr), J.A. Steyermark 7 (HOLO » Fl; isorypes: LL-TEX!, US!). a pailete Ta |, Phytologia 58:274. 1985. Syn. nov. Ardisia morazanensis Lundell ologia 61: 1986, non Ardisia maculata Poit. ex A. DC., Trans. Linn. Sac, London L7il ne 1834. TYPE: a FRANCISCO ee on mountain La Tigra, SW of San Juancito, 1,800—2,10 , 02 Feb 1966 (fl), A. Molina R., L.O. US!). WL az. 2 Williams, W.C. Burger & B. Wallenta a pe: Fl; isorypes: NY!, Distribution and ecology. ete ON TaNS montane rain forests, evergreen cloud forests, primary forests, 900—3,300 m. A widely distributed species, from Jalisco, Veracruz and Chiapas, “Mexico to Costa Rica. The type of Gentlea mcvanghii represents populations with yellowish- green (instead of green) flowers and very short inflorescences. Populations corresponding to the type of Gentlea cuneifolia are notable only for their more cuneate leaf bases and inflorescence branches more densely glandular- puberulent. The type of Gentlea maculata is notable only for its slightly longer flowers. EXCLUDED NAMES Ardisia mexicana Lundell, Wrightia 3 3:77. 1963. Gentlea mexicana (Lundell) Lundell, Wrightia 5:44. 1974. Type: MEXICO. Jaisco: crest of ridge facing the 706 Sipa 17(4) Pacific, LO mi S of Autlan, ca. 5,700 fe {1,737 m}, 20 Aug 1949 (fl), R.L. Wilbur & C.R. Wilbur 2460 (HoLotyPeE: LL!; isoryer: MICH!) This species is a member of Ardisia subgenus Icacorea (Aubl.) Mez, on account of its furfuraceous-lepidote leaves and ciliolate calyx margins. How- ever, we have as yet to determine if this species has an earlier name in Ardisia. Gentlea latisepala Lundell, Wrightia 5:4. 1972. Tver: GUATEMALA. Baja Verapaz: Chilasco, on Coneauain Road, without a ion, 29 Jul 1971 (fr), E. Contreras 10927 (HOLOTYPE: LL-TEX!). This species belongs to Ardisia Subgenus Icacorea (Aubl.) Mez, and spe- cifically, to che Ardisia costaricensis Lundell complex of species. However, in che absence of flowers, we have been unable to determine its exact system- atic position. Gentlea stevensii Lundell, Wrightia 6:97. 1979. Ardisia stevensii (Lundell) Lundell, Phytologia 61:67. 1986.11 l'ypr: NICARAGUA. CHonra es: ca. 2.8 km above N of C uapa, 400-500 m, 21 Jan 1978 (fr), WD. Stevens 6053 ee PE: LL-TEXN!: ISOTYPE: MO!), This species is a synonym of Ardisia densiflora Krug & Urban. Upon closer examination of the specimens, their sessile fruits and prominently rugose calyx lobes leave no doubt as to the true identity of this taxon. ACKNOWLEDGMENTS We thank the Missouri Botanical Garden and the Flora Mesoamericana Project for funding that allowed J. Ricketson (MO) to travel to Fort Worth, where he was joined by J. Pipoly (BRIT) on visits to the C.L. Lundell Herbarium (LL-TEX) housed at the University of Texas at Austin. We are grateful to the staff of TEX, especially Tom Wendc, Carol Todzia and Billie Lee Turner, for their cooperation and hospitality. We are also grateful to the assistance given to us by Gerrit and Jeany Davidse, Linda Oestry and Catherine Mayo (MO), Barney Lipscomb, Lindsay Woodruff, Stacy Miles and Jim Rivers (BRIT). Our most sincere gratitude is also given to Rogers McVaugh, who generously gave us information regarding the Sessé and Mocino plates. We gratefully acknowledge the assistance of James T. White, Curator of Are and Principal Research Scholar at the Hunt Institute for Botanical Documentation in Pittsburgh, who generously provided us witl a copy and slide of the original plate of Heberdenia penduliflora. Charlotte Taylor, Gerrit Davidse, and Barney Lipscomb reviewed the manuscript and provided helpful suggestions. REFERENCES Arias Diviro, J.C. 1968. Las Expediciones Cientificas Espafiolas Durante el Siglo XVIII. 27 pp.,59 plates. Real Jardin Botanico, Madrid. RICKETSON AND Pipoty, Gentlea in Mesoamerica 707 Bark ty, FA. 1945. Spanish Botanical ace to pou Spain from 1787 to 1804: The Anacardiaceae of Sessé and Mocifio. Biologist 28:9-2 CaNDOLLE, A. De., J.M. Mocino, and J.M. br Se I 874, Calques des dessins de la flore du Mexique, de Mocifio et Sessé qui ont servi de types d’espéces dans le Systema ou le Prodromus, 10 pp., 1171 plates. Geneva LuNDELL, C.L. 1964. Studies of the American Myrsinaceae—II. Wrightia 3:97-114 McVaucu, R. 1977. Botanical results of the Sessé and Mocifio Cee (1787-180 3): I. Summary of Excursions and Travels. Contr. Univ. Michigan Herb. 11:97— _ «21: 2980. Botanical results of the Sessé and Mocino Espen Been 1803): Il. The Poe Florae Mexicanae. Contr. Univ. Michigan Herb. 14:99-140 «1:98 2. The isa paintings of the Sessé and ey Expedition: A newly available resource. Taxon 31:691-692, Rickert, H.W. 1947. The Royal Botanical Expedition to New Spain. Chron. Bor. 1 1:1- 86, plates 44-52. Wison, IH. 1962. Scientific cues of Spanish exploration in New Spain during the late eighteenth century [i-x}, 330 pp. Ph.D. thesis, Univ. of Southern C alifornia BOOK REVIEW CiesscH, Bersy, drawings by Caror D. Barner. 1997. A Book of Salvias— Sages for Every Garden. (ISBN 0-88192-369-9, hbk.) Timber Press Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, (800) 327-5680, (503) 227-2878, Fax (503) 227-3070, hetp// www.timber-press.com/ $29.95. 269 pp, 87 color photos, 9 color paint- ings, 40 b&w line drawings done at 1.5 of actual size 6" X 9". Betsy Clebsch ts a noted amateur bocanist and horticulturise in che San Francisco Bay area who has designed and maintained gardens in Virginia, Texas, and California. “Gardening has been a thread pulled tight thi oughout my Life. a steady interest that binds the pages that follow,” says the auchor. She became intrigued with salvias when she began her second garden, a California country retreat. U nattende d tor several weeks at a time required the plants to be of resilienc and drought tolerant stock, planes like many of the salvias. “Writing about the culture of salvias would not have been possible had | not had a large garden in which to grow, observe. and enjoy the plants...and nursery people and botanists who visited the garden gave invaluable — ition as to a plant’s source and who had grou Wonderfully diverse in fragrance, bloom, habit and color, Sa/vias are the largest genus in ae mint family, Lamiaceae. There are over 900 species worldwide and well over half occurring in the Americas with predictions by Ms. Clebsch of new and unusual salvias soon to be available in the US from the Far East (China, Japan and India). Through twenty centuries salvias have been used and cultivated, but due to the large lem in ase of species and diverse native locations, botanists have been unable to creat t ered a fine Qu a single work. From a horticulcurise poine of view, this book should be consic start with lots of practical cultivation information. For each of the 101 alphabetically li salvias in the book, there is given: common and botanical name with Latin seme inet synonym (when a name change has caused confu- sion), cultivars and hybrids, native habitat, elevation range, temperature tolerance, and practices are well defined including tips on measurements (metric and imperial). Cultura pruning. Listed in the back are extremely helpful topics: Where to See Salvias (US, British Isles, France and Australia), Where to Buy Salvias, Flowering Guide by Season, Cold and Shade Tolerance Guides, Salvias with Especially Handsome Foliage, Salvias for Contain- ers, Color Design: itions of Unusual Flower or Foliage Color, and Geographical Origin of oa n Certain Sa/via Spec 300k of Salvias otters a wealth of information with lots of color photography and ac and new ideas for every gardener.—Lenny Heagy. Sipa 17(4): 708. 1997 NEW SPECIES OF PSYCHOTRIA SUBG. HETEROPSYCHOTRIA (RUBIACEAE: PSYCHOTRIEAE) FROM SOUTH AMERICA CHARLOTTE M. TAYLOR Missouri Botanical Garden P.O. Box 299 St. Louts. MO 63166-0299, U.S.A. ABSTRACT he new species Psychotria canligera, P. fusiformis, P. hypochlorina, and P. paeonia are de- scribed and illustrated. These are all shrubs or small trees found in wet forests 1n Colom- bia, Wenezuela, Ecuador, and Peru RESUMEN Se describen e ilustran las especies nuevas Psychotria cauligera, P. fusiformis, P. hypochlorina, y P. paeonia. Todas son arbustos 0 arbolitos que crecen en bosques htimedos en Colombia, Venezuela, Ecuador y Pert. Among recent collections from western South America, the following undescribed species of Psychotria L. Subg. Heteropsychotria Steyerm. were eee The characteristics of this ae its relationships, and its rec- ognition have been documented recently by Steyermark (1972, | 974) and Taylor (1994, 199Ga, 1996b). a cauligera C.M. Taylor, sp. nov. (Fig. | A—D). Typr: ECUADOR. Pastaza: Veracruz, ca. 900 m, 18 Feb 1956, E. Aspland 19454 (HOLOTYPE: 9). Haec species a congeneris stipulis sat grandis ovatis vel ellipticis atque inflorescentils (ut videctur caulogenis, vere terminalibus in ramulis vestustioribus) irregulariter ramosis, rhachidibus irregulariter flexuosis raceimformibus, floribus in glomerulos sessiles bracteis violaceis bene evolutis involucratos dispositis distinguitur. Shrubs flowering at 0.25 m tall, to 2.5 m tall; stems quadrate becoming terete, glabrous or pilosulous to villosulous becoming gl labrescent with age. Leaves paired; blades elliptic to obovate, 12-23 cm long, 8-13.5 cm wide, at apex acute to usually acuminate with tips to | cm long, at base acute to obtuse, drying papyraceous to chartaceous, adaxially glabrous, abaxially glabrous or hirtellous becoming glabrescent with age; secondary veins | 6- 20 pairs, spreading, broadly curved, extending to near margins or some- times looping to interconnect in distal part of blade, with (O—)-1 weak intersecondary veins usually present between pairs of secondary veins, adaxially the costa and remaining venation plane, abaxially the costa prominulous to prominent, the secondary veins prominulous, and the mi- Sipa 17(4): 709-718. 1997 710 Sipa 17(4) Fic. 1. A-D, Psychotria cauligera C.M. Taylor. A, Apex of leaf-bearing stem. B, Inflorescence borne on older stem. C, Fruiting glomerule, with fruic partially visible. D, Corolla. E-G, Psychotria fusiformis C.M. Taylor. E, Flowering branch. F, Flower bud. G, Flower at anthe- sis, partially dissected. A, B, E to 5-cm scale; C, D, F, G to 1-cm scale. A, B, based on Olleaard 98491:C, D, based on Asplund 19454; E, F, G, based on Lugo 426. Tavtor, New Rubiaceae species from Colombia 711 nor venation plane to a little thickened; margins thinly to distinctly carti- laginous; petioles 1.5—5 cm long, glabrous or hirtellous becoming glabres- cent with age; stipules glabrous or hirtellous becoming glabrescent with age, persistent at least with leaves, interpetiolar, ovate to elliptic, 12-30 mm long, obtuse to rounded, entire to ciliolate. lflorescences terminal but overtopped by well developed stems from axils and appearing cau —_— igerous, deflexed to pendulous, sessile or supsessil e, corymbiform to rounded, 4—15 x 6-15 cm excluding corollas, with 3-6 pairs of developed secondary axes, with flowers sessile in glomerules of 3-7. bracts entire to ciliolate, elliptic to ovate, 4-6 mm long, glabrous to puberulous, obtuse to rounded or trun- cate, purple to lilac; flowers with calyx limb glabrous, ca. | mm long, di- vided nearly to base, lobes deltoid to ovate, acute to shortly acuminate, ciliolate; coro//a tubular-funnelform, white, straight at base, externally gla- brous, internally glabrous except for a sparsely pilose ring ca. 2/3 of length of tube above base, tube ca. 8 mm long, ca. | mm diam. near middle, lobes triangular, 2—2.5 mm long, acute, not thickened adaxially; anthers in ee styled form ca. 1.5 mm long, partially exserted; s¢7gmas in short- styled form ca. 3.5 mm long and positioned near middle of tube; disk ca. 0.5 mm high, annular. Infructescences similar to inflorescences (plants frequently flowering and fruiting concurrently); frit ellipsoid, ca. 5 x 2.5 mm, somewhat flaccened laterally, glabrous to usually puberulous, bitte: pyrenes with 3-5 low rounded longitudinal angles or ridges. Paratypes. ECUADOR. Morona- Santiago: Along narrow flood plain of Rio Itzintza, Cordillera Cutuctt, ca. 02°40'S, 78°W, Camp 1222 (NY, S); ridge eee Rios Itzintza and Chupiasa, Cordill era Cutucti, ca. 02°40'S, 78°W, Camp 1281 (NY, US), eastern slope and crest of main Cordillera Cutuct, Jorgensen CuJ-376 (NY); Cordillera de Cutuct, 25 SE of Logrofio, Madison & Coleman 2624 (NY); — . Cutucti, western slopes, along — ny =) > a eal from Logrofio to Yaupi, in the general region, 02°46'S, 78°00 W, Magons ét al. 320 (US). Napo: S slope of C eee de Cusco new ond Cotunda-Coca, 3— ae turn-off of Baeza-Archidona road, 01°52’S, SW, Stein 50Gr (MO). ieee Puyo, ca. 2 km E of village, Asplund 19313 (S); 3. — N of Puyo, Fagerlind & Wibom 1074A (S); Tentiente H. Ortiz, 18 km from Puyo on can to ae O1°23'S, 77°57 W Olleaard & Balslev 9224 (AAU, MO, NY). Zamora-Chinchipe: Rio Na angaritza, Colina Salada, ca. 2 km E of Destacamento Shaime, 04°22'S, 78° vie W, ee 98491 (AAU). Canton Nangaritza, Rio Nangaritza, Shaime, 04°20’S, 78°40 W, Palacios 6631 (MO, Distribution and Habitat.—In wet forest at 900—1,600 m, central to south- ern Ecuador. Collected in flower and fruit apparently usually concurrently, June, November, December. This species is distinguished by its relatively large, ovate to elliptic stipules and inflorescences that appear cauligerous; these are ter minal from older growth and irregularly branched, with the axes irregularly flexuous and racemiform; the flowers are borne in sessile glomerules and enclosed by Sipa 17(4) well developed red-purple bracts. The specific epithet refers to the position of the inflorescences. The inflorescences appear to be perennial, though field observations of this feature are lacking. This species is placed in Psychotria rather than Palicowrea based on its corollas chat are straight at the base and pubescent internally only above the middle of the cube; within this group, it belongs to Subg. Hereropsychotria. However, Psychotria cauligera is overall most similar in general aspect to Paltcourea grandistipula (Standl. ex Steyerm.) C.M. Taylor, which can be distinguished by its capitate or subcapitate inflorescences that are borne at and tubular corollas chat are swollen at young stem apices, with the leaves, the base and pubescence just above this basal swelling. oan fusiformis C.M. Taylor, sp. nov. (Fig. LE-G). Typr: ECUADOR. AZA: Colonia 24 de Mayo, side road to road Puyo-Puerto penC, ca. 18-20 km en Puyo, 13 Sep 1968, H. Lugo 8S. 426 (HoLoTypE: MO-4278937: tsorypr: GB), species a congeneris Horibus pedicellatis patentibus atque corollarum fusiformium Hac bale appendices lineares gerentibus dist se cinas Shrubs and trees lowering at 3 m tall, to 12 m tall; stems quadrate to terete, glabrous. Leaves paired; blades elliptic to elliptic-oblong, 5-14 cm long, 2—5.5 cm wide, at apex acuminate with tips 5-13 mm long, at base cuneate to obtuse, drying papyraceous, adaxially and abaxially glabrous; secondary veins 7—1 1 pairs, extending nearly to or sometimes uniting weakly with margins, with |—3 pairs of intersecondary veins usually present be- tween pairs of secondary veins, adaxially costa and secondary veins plane to a little thickened, abaxially costa prominulous and secondary veins plane or a little thickened; petioles 5-25 mm long, glabrous; stipules glabrous, interpetiolar and also sometimes shortly united intrapetiolarly, incerpetiolar portion ovate, 4—5 mm long, bilobed for ca. 1/4 of their length, the lobes ligulate to somewhat elliptic, often overlapping laterally, obtuse to rounded, entire. Inflorescences terminal, apparently deflexed, with peduncles 3—10.5 cm long; panicles pyramidal, 5-12 x 5—12 cm, with 5-8 sale of developed secondary axes, with flowers pedicellate in dichasial cymules of 2—3 or soli- tary; bracts glabrous, nee acute, ciliolate, green, those subtending secondary axes 2-6 mm long, those subtending flowers 0.8—-1 mm long; pedicels in terminal flowers of cymules and solitary flowers ca. 0.5 mm long, pedicels in lateral Howers of cymules 3-5 mm long; flowers with hy- panchium obconic, 1-1.2 mm long, glabrous; ca/yx limb 0.5—1 mm long, glabrous, truncate to sinuate; coro//a in bud tubular to somewhat fusiform and externally densely puberulous to shortly, stoutly papillose-pubescent, at anthesis infundibuliform, yellow, externally sparsely pubescent to gla- brescent, internally glabrous, tube ca. 8 mm long, ca. | mm diam. at base and ca. 3 mm diam. at middle, lobes 5, triangular, 1.5—2 mm long, at apex —_— Taytor, New Rubiaceae species from Colombia 713 acute with a linear, glabrous appendage 0.2—-0.5 mm long; anthers ca. 2 mm long, positioned ca. 2/3 of length of tube above base; s/y/e and stigma not seen; disk 0.3—0.5 mm high, smooth, annular. Infructescences and fruits not seen. Paratypes. ECUADOR. Morona-Santiago: Puerto Santana, ca. 5—6 km from Shell- Mera, H. Lugo S. 448 (GB, MO). Napo: 9 km rio abajo de Puerto Misahualli y 2 km al S de la cuenca del Rio Chinguipino, 01°05'S, 77°36'W, Neill et al. 6053 (MO). Pastaza road Puyo-Macas (under construction), 31 km from Puyo, 01°37’S, 77°50 W, Olleaard & Balslev 9044 (AAU, F, US). Distribution and Habitat.—Wet forest at 430—-1,100 m, east-central Ec- uador; collected in ower March, August, September. This new species is distinguished by its interpeciolar stipules that are shortly bilobed with the lobes rounded to obtuse, spreading pedicellate flowers, and corollas that are densely puberulous to stoutly papillose-pu- bescent externally in bud and bear a short linear appendage at the apex of each lobe. The specific epithet refers to the distinctive shape of the corolla in bud. It belongs to Subg. Heteropsychotria, vegetatively and in many de- tails of the Howers, this species is similar and probably related to Psychotria caerulea Ruiz & Pav. and P. nantensis Standley, both of which differ from P. fusiforme in having sessile flowers that are partially enclosed by their larger floral bracts. Psychotria hypochlorina C.M. Taylor, sp. nov. (Fig. 2A—B). Type: PERU. Loreto: Prov. Maynas, Saboya, Rio Pintuyacu, 03°50'S, 74°10 W, 150 m, 19 Apr 1986, R. Vdsquez, N. Arévalo, GN. Jaramillo 7420 (HOLOTYPE: MO-5079650; ISOTYPE: GB). Haec species a Psychotria costanensts Steyerm. et P. cuatrecasasit (Standley ex Steyerm.) C.M. Taylor inflorescentia subcapicata vel breviter semel-ramosa atque foliorum venatione minore abaxialiter invisibili distinguitur. Shrubs and trees flowering at 3 m tall, to 8 m tall; stems quadrate be- coming terete, glabrous. Leaves paired; b/ades elliptic to elliptic-oblong, 8— 30 cm long, 2.8—14 cm wide, at apex acute to usually acuminate with tips 8-12 mm long, at base acute to cuneate or obtuse, drying chartaceous, adaxially and abaxially glabrous, when dry discolorous, adaxially dark, abaxially paler and yellowed; secondary veins | 1-14 pairs, extending nearly to margins and then reticulating extensively, with 1-3 intersecondary veins present between pairs of secondary veins, adaxially venation plane or costa a little thickened proximally, abaxially costa prominulous and remaining venation plane or a little thickened; petioles 1—-5.5 cm long, glabrous; stipules glabrous, interpetiolar and also shortly united intrapetiolarly, interpetiolar portion bilobed, between the lobes truncate to a little concave, 3-4 mm long, lobes triangular to narrowly so, 1-2 mm long, acute, entire. Inflorescences terminal, apparently erect, glabrous, subcapitate or branched once, with Stipa 17(4) A-B, tie at ae C.M. Taylor. A, Flowering branch. B, Flower bud, chotria ane M. ‘Taylor. C, F low vering branch. D, Fruit. E, limb and hypanthium. > 5-cm scale; B, D, E, F lon line & Stahl 26747; ), based on Fic. 2 partie a dissected, € Corolla, partially sees, P, Calyx o l-cm scale. A, B, based on Croat 20513: C. basec Schunke 5796; E, B based on Smith 2438. peduncles 1.2-5.5 cm long and stout, heads or panicles hemispherical to é i pyramidal, 1-2 x 1—3 cm excluding fruits, unbranched or sometimes with | pair of developed, stout secondary axes, with flowers sessile in glomerules of ca. 5—15; bracts glabrous, triangular to ovate, acute to acuminate, ciliolate, Taytor, New Rubiaceae species from Colombia 715 apparently green, those subtending secondary axes and glomerules 5-8 mm long, those subtending flowers 3—5 mm long ; flowers with hypanthium obconic, ca. | mm long, glabrous; calyx limb 12 mm long, glabrous, truncate or sometimes 5-denticulate; coro//a in bud infundibuliform, greenish white to lilac, externally glabrous, lobes 5, triangular. I nfructescences similar to inflorescences or with axes expanded; frvts ellipsoid, 15-16 x 10-11 mm, glabrous, becoming yellow-orange or orange then red then black, crowned by the persistent calyx; pyrenes 2, with 4—5 longitudinal ridges. PARATYPES. PERU. Loreto. Prov. Maynas: Rio I Mepe at Bateada de Isla Inayuga, Croat 20513 (MO); via Nauta-Iquitos, 3 km de Nauta, 04°30'S, 73°32'W, Diaz G Jaramillo 1260 (MO); Rio Nanay, Caserio Mishana, 30 km SW of Iquitos, ee 4230 (MO), 4368 (MO, NY); Mishana, Rio Nanay cae ay fee Iquitos and Santa Marfa de Nanay, near Sc eln , Gentry et al. 3656 QO); carretera Nauta- pa Km 4.5, 04°29'S, 'W, Grandez & Ruiz 2149 oe eve nn 03°59'S, 73°35 W, Vdsquez & Criollo 1975 (MO), Vasquez et al. 5340 (MO). Prov. Requena: without cies Pereira et al. 106 (MQ); distrito Sapuena, Poblado Jenaro Herrera, CIJH-HAP, Acevedo & Daly 1602 (NY), Jenaro Herrera, Rio Ucayali, 73°45'W, ( 04°55’S, Gentry et al. 56277 (MO), north sac of fe Chacra Canamé, east side of river, Mathias G Taylor 5540 (MO), Requena, Chacra Canam, J. Schunke Vigo 6248 (MO), south of Requena, J. Schunke Vigo ve 3 (MO); alrededores de Requena, 05° > S; 73°50'W, Vasquez & Jaramillo 4729 (MO). Prov. Ucayali: Canchahuayo (Rio Ucayali), 07°05'S, 75°10 'W, Vasquez et al. 6967A (MO) Distribution and Habitat.—Wet forest at 125-500 mm, northeastern to east-central Peru, frequently on sandy soils; collected in Hower November, in fruit February—September and November—December. This new species is distinguished by its leaves with the lower (abaxial) surface drying markedly paler than the upper (adaxial) surface and the ve- nation hardly visible, acne to shortly branched inflorescences, trun- cate relatively well developed calyx limb, and relatively large fruits. The specific epithet refers to the unusual drying color of the abaxial leaf surface, which is distinctive and is also found in a few other, apparently related species of Psychotria Subg. Heteropsychotria. These species also share similar leaf venation characters, inflorescence morphology, and relatively large fruits. Steyermark treated the species of this group from northeastern South America in Section Psexdocephaelis Steyerm., Series Appunianae Steyerm. These have been frequently confused, but can be separated according to the key below. KEY TO THE SPECIES OF PSYCHOTRIA SUBG. HETEROPSYCHOTRIA SECT. PSEUDOCEPHAELIS, SERIES APPUNIANAE 1. Inflorescences umbelliform, with secondary axes 3—5, all produced from firs node and equal in length to primary axis ....... P.c ionephoraides (Rusby) C .M. Taylor ay _ Inflorescences capitate or pyramidal, a secondary axes 1—2 pairs anc shorter than the primary axis, so the structure is pyramidal. 116 Sipa 17(4) 2. Stipules 11-15 mm long, with lobes 1/2 or more of this length ........... P. tepuiensis (Steyerm.) Steyerm. 2. supe les 3—8 mm long, with lobes less than 1 2 this length. . Inflorescences with eae nee axes Clearly ue the primary axis with at least one internode > | cm long. . Reciculate minor venation ae on lower (abaxial) sal f surface igo} . transiens Wernham . Secondary and intersecondary veins evident on lower (: tee leaf surface but reticulate minor venation noc visible. 5. Leaves with usually | intersecondary vein present between pairs of secondary veins; growing at 100—-1,200 m P. cordobensis C.M. Taylor 5. Leaves with | or 3 dence veins a net between pairs of secondary veins; growing at 1,500—2,05 ervisei (Standley) C.M. Taylor 3. Inflorescences capitate or with secondary axes developed, the primary axis with internodes all < 1 cm long. . Inflorescences subcapitate to once branched, with bracts subtend- ing secondary axes or outermost bracts of head triangul ar tO Ovate, acute, 5-8 mm long; peduncles 1.2—5.5 cm long; leaves with re- ticulate minor venation not ev ident On lower surface Pp ian C.M. Taylor . Inflorescences capitate, unbranched, with outermost bracts ovate suborbicular or oblong, acute to obtuse to rounded, 5—14 mm con le aves with reticulate minor venation evident on lower surface - Involucral bracts 5-8 mm long; peduncle 2.5—9 cm long; sdiouiles 3—4 mm long; notheastern to central South America P. costanensis Steyerm. Bae ral bracts 10-14 mm long; peduncles 1-5 cm long; stipules QO mm long; western South America cuatrecasasui (Stand. ex Steyerm.) C.M. Taylor (perhaps noc distinct from P. costanensis, see Taylor 1994 Psychotria paeonia C.M. Taylor, sp. nov. (Fig. 2C-F). Typr: PERU. Amazonas: alrededor de Yucui Entsa, 6 horas de Pongo por el Camino de Kusu, | nie 2 OOO Mar 1973, R. Kayap 550 (uoLorypr: MO-507965 1; isorype: G ec species a Psychotria adenophora Steyerm. stipulis 8-15 mm nen atque ili cetera bracteis externis 10-15 mm longis distinguitur. Shrubs and subshrubs flowering at 0.7 m tall, to 2.5 m tall; stems quad- rate becoming terete, glabrous. Leaves paired; blades elliptic to somewhat lanceolate, 11-19 cm long, 3.5—8 cm wide, at apex acute to sligl atly acumi- nate, at base acute to cuneate, drying papyraceous, a glabrous, abaxially puberulous and green or purple; secondary veins 9-12 pairs, ex- tending nearly to margins and often uniting into a looping < vein, with 1-3 weak intersecondary veins usually present between pairs of secondary veins, adaxially costa and secondary veins plane to thickened abaxially costa prominulous and secondary veins a little thickened to Taytor, New Rubiaceae species from Colombia 717 prominulous; petioles 5-15 mm long, glabrous; st/pu/es glabrous, interpeciolar, lanceolate to ovate, 1O—15 mm long, bilobed for ca. 1/8—1/4 of their length, the lobes triangular, acute, entire. Iflorescences terminal, perhaps deflexed, capitate to eee, with peduncles 3-8.5 cm long, heads hemisphert- cal to oblate, 1-2.5 X 2—3.5 cm, with flowers sessile in glomerules of 5— 15; bracts glabrous, ae to rounded, entire, green becoming purple, those enclosing the inflorescence elliptic to suborbicular, 10-15 mm long, those subtending flowers elliptic to narrowly so or obovate to oblanceolate, 6-10 mm long; flowers with hypanchium obconic, ca. | mm long, glabrous; calyx limb glabrous, 0.8-1 mm long, lobed to base, lobes oblong to ellip- tic, obtuse to rounded; coro//a funnelform, straight to a little swollen at base, straight co a little constricted near middle, and then spreading, white, externally glabrous, internally glabrous except for a zone of sparse pubes- cence from near middle to the base of the anthers, tube 7-8 mm long, |- 1.5 mm in diameter near base and 3—4 mm in diameter at mouth, lobes 5, 1.5—2 mm long, triangular; anthers in apparent long-styled form 1.5—2 mm long, positioned at or just below middle of tube; s/gmas in apparent long-styled form linear, ca. 1 mm long, positioned at mouch of corolla tube; disk annular, ca. 0.5 mm high. [vfrvctescences similar to inflorescence or becoming a little more expanded, with bracts purple; /rvz ellipsoid, 5 2.5-3 mm, purple; pyrenes 2, with 3—5 low, rounded, longitudinal ridges. Pararyprs. COLOMBIA. Amazonas-Vaupés: Rfo Apaporis, Cachivera de Jirijirimo y alrededores, Schultes & Cabrera 12898 (COL). Antioquia: a - San Carlos, Corregimiento Alto de Samana, Vereda Ne eaioies. Fonnegra el a 3039 (HUA, MO), a sio de Amalfi, Vereda Peldar, Mina La Viborita, 06°55°N, 75°04 a : onnegra et al. 4516 (HUA, MO). ECUADOR. Morona-Santiago: 6 km E of ae on road (under eeeras to La Unién, Harling & Andersson 24481 (GB); end of road in construction Limon-La Unién, ca. 10 km from Limon, Harling & Stahl pia (GB), 26747 (GB, S$); Shuinia Naic, ca. 8 km SE of Misi6n Bomboiza, 03°30'S, VW, Holin- Ae et al, 4381 (AAU,S), 4382 (AAU, MO, §); about 1/2 hour by car ae unfinished road E of El Limon, van der Werff & Palacios 10425 (MO). Zamora-Chinchipe: El Pangui Canton, along road between Zamora and Gualaquiza, 70. 2 km N of bridge over Rio Zamora in Zamora, between Los Encuentros and El Pangui, 03°42’S, 78°25'S, Croat 72733 (MO); 3 km E Paquisha, Harling G sea ie 2 ee seep ene Canton. P che utza, Camino al Hito, Cordillera del Céndor, 04°07'S, 37'S. Palacios et al. 8321 (MO), 8372 (MO); hill about 2 km downstream oe oe Shaime along Rio Nangaritza, van der Werffet al. 13055 (MO). PERU. Amazonas. Proy. Condorcanqui: near Yuwi Inci Creek, 6 hours’ walk from the Pongo Mori, Rio Comaina, Ber i 958 (MO); Quebrada de Basusinuk, tributary of Huampami, Kayap 90 (MO), distrito El Cenepa, Region Norori ental del Maranon, Rio Cenepa, comunidad Tucino, 04°33'S, 78° 10'S, Vasquez et al. 18519 (MQ), 18544 (MO). Cuzco. Prov. Paucartambo: Cosnipata Valley, Rio Tono First, on road N of Patria, 13°07°S, 71°12'W, Wachter et al. 217 (F, MO). Huanuco. Prov. Leoncio Prado: E of Tingo Maria, J. Schunke Vigo 5796 (MO), districo Rupa Rupa, E de Tingo Marta, cerca al Cerro Quemado, J. Schunke Vigo 9951 (MQ). Pasco. Prov. Oxapamp : Pichis Valley, San Matias Ridges, 10-12 km SW of Puerco Bermudez, above Santa aa de Chivis, trail to = 718 SipA 17(4) Loma Linda, 10°20'S OO'S, Foster et al, 8987 (MO): W of Cordillera San Matias above Santa Rosa de C | : N. Smith et al. 2438 (MO). VENEZUELA. Mérida: Distrito Tovar, La Llorona, on road to Amparo, 08°22’N, 72°45'W, van der Werff & Ortiz 5752 (MQ). Distribution and Habitat.—Wet forest at 250—2,000 m, south-central Peru (Cuzco) to northwestern Colombia (Antioquia) and western Venezu- ela (Mérida); collected in flower February—April, September, November, and December, in fruit February, April, June, July, October, and November. This species is distinguished by its lanceolate to ovate, interpetiolar stipules that are LO—15 mm long and shortly bilobed, capitate to subcapitate inflorescences with obtuse to rounded bracts that are green when the flowers start blooming and become purple as the fruits develop, funnelform white corollas with tubes 7-8 mm long, and rather narrow fruits. The specific epithet refers to the general aspect of the inflorescences and infructescences, which are reminiscent of Howers of Paeonia (Ranunculaceae). Within Subg. Hereropsychotria, this new species is similar to Psychotria adenophora Steyerm. and P. ostreophora (Standley) C.M. Taylor (syn., P lucentifolia (Blake) Steyerm.): both of these latter species can be distinguished by cheir stipules with cruncate sheaths 0.5—1 mm long and slender lobes 5-12 mm long. ACKNOWLEDGMENTS I thank R.E. Gereau for preparation of the Latin diagnoses, D. Lorence and J. Pipoly for helpful reviews, and my collaborators in western South America for providing access to specimens. REFERENCES STEYERMARK, J.A. 1972. Psychotria. In: B. Maguire & Collaborators, Flora of the Guayana Highlands. Mem. New York Boc. Gard. 23:406—717. —________.. 1974. Psychorria. In: T. Lasser, ed. Flora de Venezuela 9:1111—1683. Instituto Botanico, Direcci6n de Recursos Naturales Renovables, Ministerio de Agricultura y “ria. Caracas, Venezuela. TAYLOR, C, a 1994, Ta axonomic Notes on Psychotria (Rubiaceae) in western South America. Novon 4:303—306, 199G6a. Taxonomic Notes on the tribe Psychotric ae (Rubiaceae) in Panama, 214 western Colombia, and Ecuador. Novon 6: 210— —______.. 1996b. Overview of the Psychotrieae ioalbaceney in the Neotropics. Opera Bor. Belg. 7:261—270. THE VEGETATION OF CUCPHUONG NATIONAL PARK, VIETNAM NGUYEN NGHIA THIN Department of Botany Vietnam National University of Hanot VIETNAM ABSTRACT Cucphuong National Park, eon about 120 km south from Hanoi, northern Viet- nam, is the first National Park of nam. It is occupies approximately 22,220 hectares. The flora is rich containing 1,924 species in 990 genera and 229 families. It is characterised by the following: Fabaceae, Euphorbiaceae, Rubiaceae, Asteraceae, Lauraceae, Moraceae, Anacardiaceae, Combretaceae and Juglandaceae. The dominant genera of the fora are of Shorea, Ter minalia. Dracontonelum, Cinnamomum, Hirettera, Ananancarya, Tetrameles, The lite- forms of the flora are diverse and shown the following: SB = 60.44 Ph + 9.77 Ch + 11.57 H+ 7 - c + "a oth in which Ph = 13.37 MM + 13.32 Mi + 10.08 Na + : 42 Lp + 11.37 Ep + 0.67 Pp + 0.33 Hp RESUMEN — El Parque Nacional de Cucphuong, situado unos 120 km al sur de Hanoi, Vietnam de Norte, es el primer Parque Nacional de Vietnam. Ocupa aproximadamente 22,220 hectareas. La flora es rica, ya que contiene 1,924 especies pertenecientes a 990 géneros y 229 familias. Se caracteriza por las siguientes: Fabaceae, Euphorbiaceae, Rubiaceae, NER: Lauraceae, intes de la flora son Shorea, Terminalia. Dracontomelum, Cinnamomum, Hiretiera, pen Tetrameles, Las formas vitales de la flora son diversas y muestran lo siguiente: = 60.44 Ph + 9.7 oe 11.57 H + 7.82 Cr. + 10.29 Th, donde Ph 3.37 MM + 13.32 Mi + 10.08 Na + Lp + 11.37 Ep + 0.67 Pp + 0.33 Hp. 5 A Moraceae, Anacardiaceae, Combretaceae y Juglandaceae. Lo & = hou INTRODUCTION Cucphuong is situated about 120 km south of Hanoi on the boundary intersection of three provinces Hoabinh, Ninhbinh and Thanbnes, with longtitudes from 105°24'E to 105°44'E and latitudes from 20°14'N to 20°24'N (Fig. 1). It was classified for the first time as a primary forest area in 1960 and declared the first National Park of Vietnam in 1962 by the Vietnam Government. It is occupies approximately 22,220 hectares and includes a part of a limestone belt running from northwest Vietnam to the seashore in Ninhbinh Province. Geographical and topographical conditions are complex and thus the flora and fauna are also of great diversity. Topography Cucphuong is situated on karst limestone mountains with many caves, Stipa 17(4): 719-759. 1997 720 Sipa 17(4) . Cucphuong National Park. Kia. including Hang Dang or Dong Nguoi Xua (the cave of Primitive man) in which human archeological remains dated to 7,000 years ago have been found. In some caves, the mountains serve as natural walls co protect its diverse flora and fauna. The highest point is 650 m. One large valley is oriented in a northwest-southeast direction. Many other small valleys, separated from one another, contain many interesting features of natural h story. Climate Cucphuong has a monsoon, tropical climate with an average annual tem- perature during the year of 24.7°C, with a minimum 16.6°C, and a maxi- mum 39°C. The average rainfall is 2,157 mm, with an average humidity of 90%. Soil The Cucphuong soil is derived from limestone, with schist, argillite and quartz. There are different kinds of soil varying from the foot to the top of the limestone mountains including Rensin red greiss-yellow, red-yellow, black, margaliste-ferralite yellow. On the non-limestone mountains, there are several types of soil, such as ferralite yellow and ferralite yellow-red. In general, the soil of Cucphuong is of a light texture, and is generally moist because of its rather favourable character of imbibition. THIN, Cucphuong National Park, Vietnam ~I NO Streams and water regimes Song Buoi River is the only river passing through the northwest part of Cucphuong, and many streamlets occur in the summer. The streamlets dry up in the winter due to the shortage of rainfall in Cucphuong. DIVERSITY OF FLORA On the basis of collected specimens—which are deposited in the Cucphuong herbarium (CP) and on the basis of local floras (Anonymous 1971; Aubreville 1960-1994; Do Tat Loi 1991, Chan et al. 1995; Lecomte 1907-1951; Thin 1987, 1992; Pocs 1965; Trung 1978; Vu Nguyen 1987)—1,924 species in 990 genera and 229 families of higher plants have been identified and arranged according to Takhtajan’s system (Takhtajan 1966, 1969). The major groups (7 divisions) are listed in Table 1. All representatives of higher plant divisions are present in the Cucphuong forest. As expected, the most diverse and profuse division is Angiospermae. Gymnospermae are poorly represented with three species; Podocarpus fleuryi (Podocarpaceae) is a relict of the Tertiary era. Its pollen record shows that it grew in this area in the Tetriary period (Colani 1920; Trinh Danh 1985, 1986). It is fairly common on the slopes of the limestone hills. Equisetophyta is represented only by one species of Eqgassetuim. Bryophyta have been well studied is the most complete for any park in Vietnam, with 125 species in 70 genera of 31 families (Pocs 1965b). This group also shows the relatively primary character of the forest area. Polypodiophyta is rather diverse, with 127 species in 56 genera of 27 families. Some relict genera are also present - such as Claes (Fig. 2), Angiopreris, Archangiopteris and Cibotinm. Table 2 lists the most common genera and the number of species for the Caephnone flora. The Angiospermae is the most diverse, especially the class Dicotyledones (Table 1). The numbers of species in selected families are noted: Fabaceae (107 sp.), Euphorbiaceae (101), Rubiaceae (78), Asteraceae (64), Lauraceae (54), Bea Moraceae, Anacardiaceae, Combretaceae, and Juglandaceae, fol- lowing. Some families have a small number of species but may be quite dominant, such as Sterculiaceae, and Dipterocarpaceae. Species of Heritiera (Sterculiaceae) are seen everywhere in the forest, the leaves often conspicu- ous with silver white undersides. In the Dipterocarpaceae, Shorea chinensis is common. These trees have straight unbranched trunks to 65 m high and 1.5 m diameter. Terminalia myriocarpa (Combretaceae) (Fig. 3) has trunks to 45 m high and 5.5 m diameter. Tetrameles nudiflora (Datiscaceae) (Fig. 4) has a straight trunks, 45 m high and 5.5m diameter. Crnnamomum balansae (Lauraceae, called “Vu huong”) has trunks of fragrant wood, 48 m high and 2.5 m diameter. Dracontomelum duperreanum (Anacardiaceae) is well-known for its huge buttresses. 722 Sipa 17(4) Tabie |. The flora of the Cucphuong National Park is diverse, fora small area with about 0.7% of the total for Vietnam, yet the number of plant species reaches almost 25% of the total species reported for the country. Taxon Number Species Genera Families Bryophyta 125 70 4] Psilotophyca | | l Lycopodiophyca 9 2 2 Equisetophyca | | | Polypodiophyta 127 56 27 Gymnospermac } 5 3 Angiospermae icotyledones 1,28] O74 134 Monocotyledones 377 183 30 There are many lianas growing upwards by attaching themselves to the forest trees including Gnetwm sp. (Gnetaceae). Of the Angiospermae, there are lianas in Sterculiaceae (2 sp.), Fabaceae (3 sp.), and Apocynaceae (1 sp.). Entada phaseoides (Fabaceae) with stems 10-20 m long and 20-25 cm in diameter (Fig. 5), Bawhinia scandens with stems 10 m long and 10—15 cm in diameter, and Acacia pennata (Fabaceae) are very common. There are seven species of Ficus (Moraceae), some of Schefflera (Araliaceae), and Fagraea (Loganiaceae) strangling and twining around neighbouring trees (Table 3), that start life as epiphytes and whose roots grow down, interlacing around their support. Eventually the host dies as it is shaded out, leaving the stran- gler supporting itself, Various species of shade-loving epiphytes include Ps/otam nudum, Lyco- podium cavinatum, Mecodium exsertum, Lomariopsis spectabilis, Lemmaphyllum microphyllum var. obovatum, Leptochilus laciniatus var. simplex, Loxogramme lankokiensis, Vittaria (7 spp.), Ficus pumila and Aglaonema sp. Light-loving epiphytes on branches of high trees include such orchids as Aerides odoratum, Cirrhopetalum siamense. Cymbidium aloifolium, Dendrobinm spp., Luisa teretifolia, Oberonia, Rhynchostylis gigantea, Sarcanthus spp., Thrixspermum centipeda, and Vanda species. Also, there are Anadendron latifolium. Pothos spp., and Rhaphidophora spp. (Araceae); and Drynaria fortune (Polypodiaceae), as well as many species of lichens and mosses. At ground level very few plants occur except those which obtain nutri- ents from decaying plant matter such as Ba/anophora (Balanophoraceae) (3 sp.) and other parasitic species living on the roots of other plants. Some others are semiparasitic pee iving on the branches of other plants as Viscum (Viscaceae) living on Liguidambar formosana; Viscum and Loranthus spp. (Loranthaceae), and on other plants such as Averrhoa carambola, Ficus sp.. and Citrvs spp. A great number of species with lovely Howers and leaf THIN, Cucphuong National Park, Vietnam 723 Fig. 2. Cyathea contaminans (Cyatheaceae). shapes are Paris chinensis, P. hainanensis (Trilliaceae) (Fig. 6), Amorphophallus paconifolia (Fig. 7), A. rivieri, A. tonkinensis, Arisaema balansae (Araceae), Angiopteris repandula (Marattiaceae), Milletia ichthyochtona, Saraca dives (Fabaceae), Firmiana colorata (Sterculiaceae), Aerides odoratum, Dendrobinm lindleyi, D. superbum, Vanda teres, Paphiopedilum appletonianum, P. delenatit, Haemaria discolor (Orchidaceae), Tacca chantrieri (Taccaceae). Some plants are cauliflorus such as Baccaurea ramiflora, Meliantha suavis, Ficus spp., and Dimerocarpus brenniert. STRUCTURE OF FLORA The Cucphuong flora is characterized by the stamps of the tertiary era flora of North Vietnam and South China (Trung 1978). It includes 594 Sipa 17(4) TABLE 2. The most common genera with number of species for che Cucphuong National Park. Genus Spp. Genus Spp. Pteridophytes Blumea 9 Selaginella 7 Callicarpa 9 Tectarta 7 Justicla 9 Colysis 8 Mallotus 9) Lygodinm 8 Antidesma 10 Asplentun 9 Caren 10 Preris 9 Clerodendrum 10 Cyclosorus 10 Beilschmtedia 11 Spermatophytes Dendrobinm 11 Croton 8 Litsea 4 ere Ost 8 Polygonum 12 goa 8 Cyperas 13 Schefflera 8 Desmodium i) Callicarpa 8 Syzygium 14 Crmanonun 9 Ardisia 1S FPambristylis 9 Ficus 18 many species of Fagaceae, Fabaceae, Lau raceae, Juglandaceae, Altingiaceae, Ebenaceae, Annonaceae, Sterculaceae, Meliaceae, Sapindaceae, Magnoliaceae, and Anacardiaceae, and most species of Gymnospermae. Fossil evidence also affirms that earlier plants similar to the present species, such as Cinnamomum batlansae, Heritiera macrophylla, Celtis sinensis, Liquidambar formosana, Saraca dives, and Amoora gigantea occured in this area (Colani 1920; Trinh 1985, 1986). Cucphuong is a place where many different floras overlap (Pocs 1965a, 1965b) because it is the meeting point of mountain ranges from the Himalaya running northwest-southeast, from South China running north- south, and from South Vietnam running north-south. The flora Malesiana elements are characterized by species linked to a hot and humid climate derived from Borneo (Trung 1978). These elements entered Vietnam in the Tertiary Era (Trung 1978, Danh 1986) and include species of Dipterocarpaceae. Although the number of species is low, the number of individuals is relatively great. The Himalayan element of the flora is characterised by temperate spe- cies from Yunnan, Quichau and Himalaya including numbers of Fagaceae, Aceraceae, Oleaceae, Ulmaceae, Hippocastanaceae, Trilliaceae, and Juglandaceae. The Indo-Burmese element of the flora is characterised by deciduous species like Terminalia myriocarpa, Combretum spp., Anogeissus acuminata (Combretaceae), Lagerstroemia cornicilata (Lythraceae), Bombax malabaricnin (Bombacaceae), Tetrameles nudiflora (Datiscaceae), and Sapindus saponaria (Sapindaceae). Tun, Cucphuong National Park, Vietnam 725 Fic. 3. Terminalia myriocarpa (Combretaceae) with trunks to 45 m high and 5.5 m in diameter. Sida 17( 726 he trunks with straig Datiscaceae meles nudiflora Terr 1. ameter. 19 - IG. p di Tuin, Cucphuong National Park, Vietnam 727 Fic. 3. Entada phaseoides (Fabaceae) with stems 10-20 m long and 20—25 cm in diameter. 728 SIDA 17(4) Taser 3. Lianas and strangling epiphytes of the Cucphuong forest. Ficus annilata Ficus pilosa Ficus benjamina Preus religiosa Prous glaberrima Ficus retusa Ficus gihblosa Schefflera glomerulata Prens lacor Magraca obovate Ficus obtusifolia DISTRIBUTION OF THE CUCPHUONG FLORA Plant groups are divided on the basis of types of the topography and the soil. Plants on the summits of limestone mountains On summits from 300-560 m altitude with sharp hard rocks and with black renzin soil, the humidity is low and the evaporation rate is high. The diversity of plants include 65 common species such as Cycas balansae, Cénnamomum validinerve, Neolitsea zeylanica, Ulicium griffithii, Quercus sp.. Platycarya strobilacea, Eriobotrya benghalensis, E. asa E. Sie, pie chinensis, Schefflera pes-avis, Calophyllum balansae, C. bonii, Dracaena cambodiana, Diospyros mun, Mitrephora calcarea. nee sp., Nerospermum tonkinensis, Canthium dicoccum, Sinosideroxylon racemosum, Murraya Roenraii. M. paniculata, Jasminum lanceolarinm. J. longisetum, Th rigonostemon bonianus, Drypetes perreticulatus, Paphiopedilum appletonianum, P. delenatii, and Dendrobinm dentatum. Plants on slopes of limestone mountains ive growing on the slopes include some 200 species, of which there are 14 species of Pteridophytes, two species of Gymnospermae, and 184 ne of Angiospermae. The common tree species include Streblus ilicifolins, Dimerocarpus brenntert, Teonongia tonkinensis, Heritiera macrophylla, H. cucphuongensts, Plerospermum truncolobatum. P. heterophyllum, and Ficus gibbosa. Many species of understory shrubs and herbs have large thin leaves, provid- ing a large light-absorbing surface, notably species of Preridophytes, Euphorbiaceae, Rubiaceae, Urticaceae, and Araceae. Plants in the valleys In valleys between limestone walls the humidity is high. The soil is of macgalis ferralic brown-yellow. The flora here includes 1,219 species be- longing to Bryophyta, Preridophyta and Angiospermae. Many tree species have straight unbranched trunks, or have buttresses or stilt roots providing additional support. Their glabrous leaves have drip tips probably to pre- vent water accumulating after rain. Examples are Tetrameles nudiflora, Shorea chinensis, Terminalia myriocarpa, Anogeissus acuminata, Cinnamomum balansae. Pometia pinnata, Erythrophloeum fordii, Dracontomelum duperreanum, and Vatica harmandiana. Trin, Cucphuong National Park, Vietnam 729 Fic. 6. Paris chinensis (Trilliaceae). SiN i . = , , a " ity 9 Oe Ps ee a BiG. 7. Amorphophallus paeoniifolia (Arace: e). THIN, Cucphuong National Park, Vietnam eon! Plants on non-limestone soil This group has 440 species especially in the following genera: Machilus, Phoebe, Litsea, Cinnamomum (Lauraceae), Aglaia, Amoora, Chisocheton, Dysoxylum, Heynea (Meliaceae), Castanopsis. Lithocarpus (Fagaceae), Elaeocarpus (Elaeocarpaceae), Macaranga, Endospermum, Bischofia, and Sapium (Euphorbiaceae). The distribution of plants in Cucphuong is interesting. The number of species decreases from the valley to the top of the mountains. The greatest number of Preridophyte species occurs chiefly in the valleys and under the canopy of the forest. DIVERSITY OF LIFE-FORMS —" Life-forms are an ecological characteristic of every flora. Spectrum of the eo life-forms is truly a reflective mirror of ecological conditions as well as a primary or secondary character of the flora in the studied region, and also provides information on conservation and suitable use of the plant resources. On the basis of the classification categories of Raunkiaer (1934), the spec- trum of the life-forms of the Cucphuong flora was established including five main groups as follows (Table 4, Fig. 8): 1. Phanerophytes (Ph).—This group includes 1,175 species in 151 fami- lies making up 60.44% of the total flora (Fig. 8). This dominant group includes seven subgroups. 1.1. Mega- and Meso-phaneraphytes (MM).—This includes 260 large woody species taller than 8 m (13.37% of the total Hora) (Fig. 9). The species-rich families are Lauraceae (37 sp.), Euphorbiaceae (22 sp.), and Moraceae (20 sp.). The high proportion of species belongs to the following families: Bignoniaceae 6/6 species, Buseraceae 6/6, Elaeocarpaceae 8/9, Fagaceae 10/ 12. Some large size species include Shorea chinensis (Dipterocarpaceae), Tetrameles nudiflora (Datiscaceae), Terminalia myriocarpa (Combretaceae), Elaeocarpus (Elaeocarpaceae), Dracontomelum duperreanum (Anacardiaceae), and Cinnamomum balansae Tanea 28) 1.2. Microphanerophytes (Mi).—This includes 257 short woody species (2-8 m) (13.32% of se total flora) (Fig. 9). The species-rich families are Euphorbiaceae (15 sp.) and Annonaceae ( 2h.) 1.3. Nanophanerophytes (Na).—This includes 196 species in 46 families (10.08% of the total flora) (Fig. 9). The species-rich families are Rubiaceae (24 sp.), Myrsinaceae (20 sp.), Acanthaceae (19 sp.), Fabaceae (18 sp.), and Euphorbiaceae (17sp.). 1.4. Lianphanerophytes (Lp).—This includes 222 species in 40 families (about 11.42% of the total flora) (Fig. 9). The species-rich families are Fabaceae (33 sp.) and Rubiaceae (18 sp.). This group is characteristic of EW, Stipa 17(4) TabLe 1. Life-forms of Cucphuong flora. Life-form Number of species Rate | Phancrophyces (Ph) 1175 60.44 L.1. Mega-Mesop! phyces (MM) 60) 1347 1.2. Micropl pl {1) 25 13.32 13. ] phytes (Na) 196 10.08 1.4. Liz eee inerophytes (Lp) 222 11.42 I Epiphyte- B ehiaiad da aa 221 11.37 1.6. Elemi- and P phanerophyces (Pp) 13 0.67 L.7.) Herb Shenerasineest Hp) 6 0.33 2 : alae aa (Ch) 190 9.77 ‘ (H) 225 LL L. Ccouschie (Cr) 152 7.82 5. Therophytes (Th) 200 10.29 tropical forest especially woody liannas as Entada, Acacia, Bauhinia, Strych- Lo nos, Nelodinus, and Tetrastigma. 5. Epiphyte-phanerophytes (Ep).—This includes 221 species in 35 fami- lies going up to LI. 37% of he total Hora (Fig. 9). The species-rich families are Orchidaceae (48 sp.), Araceae (10 sp.). 1.6. Hemiparasite- and Parasite-phaner oD tes (Pp). —This include 13 spe- cies in four families and makes up about 0.67% of the total flora (Fig. 9). 1.7. Herb-phanerophytes (Hp).—This includes six species or about 0.33% of the tocal flora (Fig. 9). 2. Chamaephytes (Ch).—This group includes 190 species in 54 families (9.77% of the total flora) (Fig. 8). The species-rich families are Asteraceae (26 sp.), Fabaceae (24 sp.), Euphorbiaceae (14 sp.) and Acanthaceae (13 sp.). 3, Hemicrytophytes (H).—This group includes 225 species in 46 fami- lies or 11.57% of the total flora (Fig. 8). The species-rich families are Poaceae (36 sp.), Orchidaceae (15 sp.), eee (14 sp.) and Cyperaceae (14 sp.). 4. Crytophytes (Cr).—This group includes 152 species in 42 families con- stitucing 7.82% of the total flora (Fig. ine The species-rich families are Orchidaceae (25 sp.), Convallariaceae (20 sp.), Zingiberaceae (14 sp.) and Cyperaceae (11 sp.). 5. Theophytes (Th).—This group is characteristic for grassland patches next to human activities or along ag It consist of 200 species in 41 families (10.29% of the total flora) (Fig. 8). The greatest number of species are chiefly in the Poaceae (35 s cies Cypetaceae (27) and Asteraceae (26); they are light-loving. In short, the spectrum of the life-forms of the Cucphuong flora is shown: SB (Spectrum of Biology) = 60.44 Ph +9.77 Ch+ 11.57 H + 7.82 Cr + THIN, Cucphuong National Park, Vietnam 70 — Ve GIPh oe Och OH OCr 50 OTh Percent of total flora i t t T T T Ph Ch H Cr Th Fic. 8: Spectrum of main life-form groups of Cucphuong flora. oMM 121] OMi pre L 7 CiNa a ied _ 4 ee Olp | 10 aS DEp e OPp oO a a et a bosood OHp + 3 B_}" L re) L a a tl € 6} o 2 {a} oO — oad Ly rt 4) 21 AA vA AVA GD MM Mi Na Lp Ep Pp Hp Fic. 9: Spectrum of subgroups of phanerophytes life-form of Cucphuong flora. 734 SIDA 17(4) 10.29 Th, in which Ph = 13.37 MM +13.32 Mi +10.08 Na +11.42 Lp +11.37 Ep +0.67 Pp + 0.33 Hp. If making comparation with the spectra of life-forms from North Viet- nam (Pocs 1965a) and from Lamson, Luongson, Hoabinh (Le Tran Chan, Phan Ke Loc, Nguyen Nghia Thin, Nong Van Tiep, 1994) where is near the Cucphuong National Park but destroyed seriously by human activities (Figs. 1, 2) it is clear that the Cucphuong flora is much more primary than the North in general and the Lamson flora in particular. This is shown by high race of Phanerophytes group and low of Chamaephytes, Hemicrytophyces and Crytophytes. ACKNOWLEDGMENTS I would like to thank Barney Lipscomb for help during past few years and B.C. Stone and John Pipoly for comments and corrections. APPENDIX CHECKLIST OF SPECIES FOUND IN THE CUCPHUONG NATIONAL PARK Format: Distribution and soil type follows: V—Valleys, SL—Slopes of limestone mountains, SUM—Sumumits of limestone mou Atains, iWon: limestone soil. Life-forms: Ph—Phanerophytes including MM—Mega-or Meso-phanero- phytes, Mi—Microphanerophytes, Nan—Nanophanerophytes, Lp—Liana- phanerophytes, Ep—Epiphyte—phanerophytes, Pp—Parisite or Hemi- parasite-phanerophytes, Hp—Herb-phanerophytes; Ch—Chamephytes; H—Hemicrytophytes; Cr—Cryptophytes; Th—Theophytes. BRYOPHYTA ANTHOCEROPSIDA Cololejennea dankianensis Tixier, V Cololejeunea floccosa (L. & L.) Steph., V ANTHOCROTACEAE Cololejennea formosana Miz., V Anthoceras miyabenus Steph., V Cololejeunea grushutzkiana Pocs, V MARCHANTIOPSIDA Cololejennea baskarliama (Gott.) Bx., V - Cololejeunea karstenti Steph., V FRULLANIACEAE Cololesennea lanciloba Steph., V Frullania grandipula Lindenb, V Cololejennea ninbinhiana Tixier, V Frullania moniliata (Reinw. Blume & Nees) Cololeseunea nymanii Steph., V “es, V Cololejennea tramninhiana Tixier, V Colura acutifolia P -EJEUNEACEAE = EJECNE : ; . , Colura ari ph., Aphanolejennea proboscoidea Goeb., V Lapa taain se Meee . x : 2 - PeEHN¢ ava (OW etlolejennea meyeniana (Steph.) Kach. & Leptolejeuna balansae Steph., V CHM AL 6 de Schust., V C een Bx. V Leptolejenna subacuta Steph., V ololejeninea appressa Bx., : . : : ee Leptolejeuna vitrea (Nees) Steph., V Cololejennea bichtana Tixier, V JOD) ¢ x1e Cololejenned crysanthemi Vixier, V Lopholejenna vienamica Tixier, V THIN, Cucphuong National Park, Vietnam LOPHOCOLEACEAE Heterocyphus argutus (Nees) Schiffn., V MARCHANTIACEAE Marchantia palmata Nees, V LAGIOCHILACEAE Swe peel = — Plagiochila fissifolia § >> Plagiochila fordiana a Plagtochila subplanata ae H.,V RADULAC EAE Radula acuminata Steph., V BRYOPSIDA AMBLYSTEGI E Scriaroniopsis bonianum (Besch.) Tixier, V BARTRAMIACEAE Philonotis revoluta Bosch & Lacout., V BRANCHYTHECIICEAE Cirriphyllium oxyrrbychun (Doz. &. Moll.) M. Fleisch., V se ie ad aciculum (Broth. & Par.) M. Fleisch., Rhy am colebicum (Lacout.) Jaeg., V Rhynchostegium vagans (Harv.) Jaeg., V BRYACEAE Bryum ambiguum Duby., V CALYMPERACEAE Syrrbopodon larmitani M. Fleisch., V DALTONIACEAE C eee tonkinensis (Broth. & Par.) Broth., DICRANACEAE Dicranella enstegia (C. Muell.) Bosch & Lacout., Vv Holomitrinm er eat Mite. uell.) Mit Hornsch. . a. Trematodon longicollis Michx., ENTODONTACEAE Erythrodontium flarascens (Hook.) Lacout., V Erythrodontium julacerm (Hook.) Par., V Tetrachyphylum inflexum (Harv.) Gepn., V FABRONIACEAE Campylodontium flavascochs (Hook.) Leucoloma molle (C ~~ Leucophanes ca a Bosch & Bosch & FISSIDENTAC Fissidens arate a Hook., V Fissidens incertus Ther., 735 Fissidens nobilis Griff., Fissidens sylvatict oF Vv Fissidens ‘allingeri Montg., V FUNARIACEAE Physcomitrinm repandum (Griff.) Mict., V OOKERIACEAE Callicestella papillata Mitt., V Pon Mum nigricaule Doz. & Molk., V HT ypopterigium sia m Thwait. & Mict., V statum Bosch & Lao Vv lypopterigium oon C. Muell., osch & | yut., W Lopodinm eee (Bosch & Tacone! M. Fleisch., V HY PNACEAE ein chamissonis (Hernsch.) Jaeg., V Ectropothecium monumentorum eae ) Jaeg., V Lsopteryginm chlorophyllam Bosch, Lypopterigium, AY 7 1€S€1 Isopterygium lignicola (Mi itt.) Jaeg., “ ne pohliaecarpum ( (Sull. & Leag.) Jaeg. 7 Lax le taxirameum (Mitt.) M. Fleisch., V ta (Doz. & Molk.) Broth., V Vesicularia tonkinensis (Bosch) Broth., V METEORIACEAE Aerobryidium fillamentosum (Hook.) M. Fleisch., Vv Vestcularia reticula Aerobryopsis deflexa Broth. & Par., sea rabryopsis Fe ads (Doz. & ae sch. I ase pseudofloribunda M. Fleisch., V Merteortopsis buchanat (Brid.) M. Fleisch., Vv Mereoriopsis squarrosa (Hook.) M. Fleisch., V MNIACEAE Plagioninm rostratum (Schrad.) Kop. T., V Plagioninm succulentum (Schard. ) Koy 5. T., V NECKERACEAE ee wrightii Mitt., V fis ae M. Fleisch., cai is targionianus (Mitt.) Dix., V » plamila Hom weal igdendron exiguum (Bosch & tacos )M. sch. Homalin i on flabellatum (Dicks.) M. Fleisch., lle aaa microdendron (Bont.) M. eisch., oe on scalpellifolinm (Mitt.) M. eisch., V Neckeropsis moutier’ (Broth, & Par.) M. Fleisch., Neckeropsis lepianeana (Ape ex C. Muell.) Toun., Vv 736 Neckeropsts sempertana (Mont.) M. Fleisch., V Pinnatella alopecuroides (Hook.) M. Fleisch., V Prnnatella ambigua (Lacout.) M. Fleisch. Vv Pinnatella donghamensis Besc Pinnatella kuhliana . acout. Me Fleisch. Pinnatella microptera (C. Muell) M. Fleisc . ORTHOTRICHACEAE Macromitrium gymnostorum Sull & Lesq., V Macromitrium nepalense H. & G.) Schwaegr., V PHYLLOGONIACEAE Hortkawaea nitida Nog., V POTTIACEAE Hycdrogonium psendochenbergt (M. Fleisch.) Chen., Vv Hymenoctomum tee a Je Vv lyophyla involuta (Hook.) Jaeg a eos c an & Their.) Broth. Tric i involutum Broth., Trichostomum pertvolutum Vixier, Weista longidens Care PTEROBRYACEAE Garovaglia crispata Tixier, V Garovaglia ue Besch., V RACOPI aid ree (C. Muell.) Mitt., V SEMATOPHYLL ACEAE ann Lig i Tix ieee (Harv. M. Fleisch., Vv Eire boutanit (Broth. & Far.) Broth., Semarophy lum seine (C.M.)M. ies Vv wxithel, (Brid.) Broth. Taxithelinm pea (Lacout.) oe & Card., finm Mnstva TRACHY PODIACEAE aa blanius (Harv.) M. Fleisch., V (Card.) M. Fleisch., MW orvida Vv THUIDIACEAE Anomodon tonkinensts Besch., V Claopodinm aciculum (B roth. ) Broth., V & Lesg. re ard., Haplocladium ninutifolion Their. Haplocladium sublacenm (Mitt.) Broth: Haplobymeninm psendotriste (C. Muell) V Claopodinm assurgen ene ge as ae tocaea (Sull. & Lesg.) Ren. & Card., Pelekinm an Mite. Thuidium bonianum Be eu Sipa 17(4) Thuidium cymbifolium (Dz. & Molk.) Bosch & “acout., Thuidinm meyentanum Ape.) Bosch & Lacout., Vv Thuidium tamaricellum (Mitt.) Bosch & Lacout., Vv PSILOTOPHYTA PSILOTACEAE Psilotum nudum (1..) SL, Ep Griseb. (P. triguetrum Sw.), LYCOPODIOPHYTA LYCOPODIACEAE Lycopodium carinatum Desy., SUM, Ep Lycopodiian cernuum L. =| SELAGINELLACEAE Selaginella biformis A. Br. ex V,H Selaginella decipiens Warb., Selaginella intermedta ee 7 ing, V, H se Ma pubescens on ) Spring, V,H : "i laginella stamensis Hier 4 . oe tamariscina ae ) Spring, V, H oe wallichii (Hook. & Griev.) Spring, EQUISETOPHYTA EQUISETACEAE Equisetum ramocissimum Dest., NL Cr ssp. debile (Roxb.) Hauke (EF. debile Roxb.) POLYPODIOPHYTA ADIANTACEAE Adiantum caudatum L., SL, H \diant Habellulatun 12 Vv, G ie tenutfolta (ait: )Sw., V, Th ASPLENIACEAE Asplentum antrophyordes Christ, V, H Asplenium belangeri (Bory) Kze, SL, H Asplenium greviller Wall., S Aspleninm wriffithianum Hook., NL, H Aspleninm nidus L., Asplenium pnt Ching, V, Ep Asplenium saxicola ,H Aspleninm ee nee v, H AZOLLAC Azolla an tcata (Roxb.) Nakai, NL, Th BL eras CEAE lechnum ortentale L., NL, Ch pee Alaena palustris (Burm.) Bedd., NL, C — al Tuin, Cucphuong National Park, Vietnam CYAT ie ta Cop., V, N Cyathea aft. contaminans (Wall.) Cop., V, Mi DAVALLIACEAE Arthropteris obliterata (R. Br.) J.Sm., Davallia formosana Hay. A alla divaricata Blume) var. or le Tard.-Bloc & C. Ch. Humata vepons (L£.) Diels, SUM, Ep Nephrolepis cordifolia (L.) Presl, SL, H Nephrolepis falcata (Cav.) G. Ch., V, H DENNSTAEDTIACEAE Lindsaea orbiculata Lam.) Mett. ex. Kuhn, SL, Microlepia hokeriana (Wall.) Fresl, V, H Microlepia marginata (Houtt.) C.Chr., V, H Microlepia speluncae (L.) Moore, V, Microlepia speluncae a _ var. hancet C. hr. & Tard.-Blot Schisoloma ae oo a Sm Schizoloma heterophyllum (Dry.) J. Sm., Srenoloma chusanum Ching, V, Cr DICKS ee er neene (L.) J. Sm., DRYOPTERIDACEAE Ctenitopsis sp.. V, H Cyclopeltis se crenata (Fee) C. Chr., Le Cr Hemigramma decurrens (Hook.) Cop., — Ouercifilix zeylanica (Houtt.) Cop ee Tectaria decurrens (Presl) Cop., V, H Tectaria devexa ae ,H Tectaria lenana ,V,H Tectaria ae te hing, V,H Tectaria simonsii (Bedd.) Ching, V, Tectaria subtriphylla (Hook. & Arn. c op.. NL, H ., SL,H SL, Cr V,Ch Tectaria variolosa (Wall.) C. Chr., V,H GLEICHENIACEAE Dicranopteris linearts (Burm. f.) Underw., NL, Cr (Gleichenia linearis Clarke GRAMMITIDACEAE Prosaptia aff. ue Cop. (Ching) Ching, § HYMENOPHYLLACEAE Crepidomanes bipunctatum soe ( Cop., V, H Trichomanes bipunctatum Poi Mecodium Wall) a V, (Hymenophyllum exsertum Wall.) Microgoninm beccarianum _ esati) Cop., SL, Ep ae IONIANES Me ae — var. intermedia Ep exsertum “ = ~ wo! ) Blume) ) Cop., SL, Ep aoe aur. ee Blume) vhoschia auviculat. 737 Vandenboschia cystosctroides (Christ) Ching, SL, Ep (Trichomanes cystosciroides G. Christ) LOMARIOPSIDACEA Bolbitis ene (Presl) Ching, NL, Ch peed “oe Mett., NL, Ep MARATTIAC para a men de Vriese, V, H Archangiopteris sp., MARSILEACEAE ursilea minuta L., NL, C ia silea cen Li, NL, Cr OPHIC {AE Ln oeerie zeylanica (L.) Hook., NL, Cr PARKERIACEAE Ceratopterts thalictroides Brongn., NL, Th POLYPODIACEAE Colysis bonii Ching, V, Colysis digitata (Bak.) Ching f. a V,Ep f. annamensis C.Chr. & Tare 1. Colysis dissimialata res Vv G . sis pothifolia (D. Don) Pre , Ep Colysis ellliptica a. var. a a C Ns wrightit Ching, V, Colysis spl., V, Ep Calysis § 2: V,Ep Drynaria fortune’ (Kuntze) J.Sm., SL, Ep Bec Hum microphyllam Presl var. obovatum (Harr.) C.Chr., NL, Ep ee a laciniatus Ching var. simplex Ching, {p iyaueames lankokiensis C.Chr., V, Ep ssa i dilatatum (Bedd. ) Siedee, V, Ch 1. hancocki (B =) es _ rosoriim Steere ing, V,C ] Neocheiropteris ae (Christ) Ching, SL, Phymatosorus longissima (Moore) J. Sm., V, Ch Phymatosorus sp., S Psendodrynaria coronans (Wall.) Ching, V, Ep (Aglaomorpha coronans Cop.) Pyrrhosia adnascens (Sw.) © hing, V, Ep Pyrrhosia flocculosa Ching, SL, Ch Pyrrbosia lingua (Thunb.) Farwell, SL, Ch Pyrrbosia tonkinensis (Gies.) Ching, V, Ep PTERIDA Pro agin Kuhn, NL, Ch Preris cadieri € Prerts Pie Bak., NL L,H(P. a aa L. var. dissHtifolia Tard. Slee &C.C Preris enstformis Burm., SL, } Preris fenotii Christ, a ] Preris grevilleana Wall., SL, 738 Sipa 17(4) Prerts khasyana Hier., V,H Diplazium maximum C. Chr., V, Ch Prerts longipes D. Don, V, H Diplazinm pinne tiftcl sinneatinh Moore, V,H Prerts oshimensis Hier., V, H Diplazium sp., Vv. H Preris semtpinnata L., V,H SALVINIACEAE PINOPHYTA Salvinia cucullata Hoftm. eae “4 h CYCADACEAE Salvinia natans \ Cycas balansae Warb., SL, Na GNETACEAE Gietum montanum Met., SL, Lp SCHIZAEACEAE Lygodinm conforme C. Chr., V, Cr Lygodium flexuosum (1...) Sw., NL, € roe wae oe Clie PODOCARPACEAI Podocarpus flearyi Hickel, SL, MM Lyvodium flexuosium ( NL, Cr L ode japonicum (Thunb.) Sw., NL, , Lygodium ean hyum Desv., NL, Cr MAGNOLIOPHYTA Lygodinm polystachyum Wall., NL, Cr Lygodium scandens (L.) Sw., NL, Cr Lygodium subareclatum Christ, NL, Ch ACANT ee a 1 i indl.-Gaidk., V, TE He -YPTE neers EAE Codonacanthus ee (Nees) Nes - Ch “optert Ching, NL, H (Cyclosorns Dicliptera chinensis (L.) Nees, V, poe: T ard.-Blot) Eranthemum nervosum ee Vv, a Abacopterts simplex (Hook.) Ching, NL, Hyerophila angustifolia R. Br. ae Ch . yelosorus triphyllus Vard.-Blot var. simplex Hygrophila phlomoides Ness Hook.) ; a Justicia equitans ®. Ben., V, Ch Abacopterss triphylla (Sw.) ie NL, H Justicia gendarusa R. a NL, Na (Cyclosorus triphyllus Vard.-Blot Justicia panpercila R. Ben., NL, Na Abacopteris urophylla C = g, NL, a (Cyclosoras Justicia procumbens L., NL, Ch sali Tard.-Ble Justicia vagabunda R. Ben., V, Ch ps Wei ote Wie ll., V, Na MAGNOLIOPSIDA Na Peristrophe roxbur vhtana (Schult.) Brem., NL, th (P. tinctoria Nees) i ada ee R. Ben., NL, Na ees, V, Na G ‘yelosorus sophoroides “~ Flor, V. H Cyclosorus truncatus lto, V,H Cyclosorus unitus Ching, V,H Cyclosorus spl., V, H Cyclosorus sp2., V, H Thelypteris singaiaensis Ching, V,H Pp Pic Sp. Pscnderantonn crenata Lindl., V, Na . : ; ee : seuderanthems, mp hate feruy, Rude. V.Na Lhelypteris tonkinensis Ching, V, H Psenderanthemum p = ee Re Bea Wy VITTARIACEAE casi quads ee R. rere V,Na A) ntrophyun ca Mife olin Wm manga V, Ep Rhj 1s Ne ees JV ; a Antrophyum cortacenm Wall. Ep fanaa communis Nees, ge Na Antrophyum sp., V. Ep Ruellia ca lis Roxb., ,H Vittaria elongata Sw., V, Ep var. merrilii ©. Che. Raellia repen & Tard.-Blot var. zosterifolia C. Chr, & Tard.- Rungia eile msis R, 'Be n., VW, Ch 300 Rungia sp., V, Ch Vittaria flexuosa Fee, V, Ep vee ae sp., V, Na Vittarta hainnanensis C. Chr., V, Ep Strobilanthes acrocephalus T. Anders., V, H STAC Strobilanthes brunnescens R. Ben. V. Na WOODSI/ i a : Str obilanthes dalztellii (Smith.) R. Ben. var. Callipteris pe Retz.) J.Sm., V,H ; vaequalis R. .V,Na Diplaz LOpsts p. ,V,H Sesh Doaethes es aK °C larke, V, Na ) 5 ’ 2(N f Diplazium donianum (Mect.) Tard. “Blot, V, Strobilanthes muiltancurus R, ae V,Na Turn, Cucphuong National Park, Vietnam Thunbercia eberhardtit R. Ben D Thunbergia grandiflora Ros. var, “ude R. Ben., V, Lp ACER Acer eae wm Merr., ae MM Acer oblongum Wall., V, Ma Acer tonkinensis Le SL, Mi Acer sp., SL, Mi ACTINIDIACEAE Saurauia tristyla DC., NL, Mi ALANGIACEAE Alangium barbatum (R.Br.) Baill. var. dectpens vrard) Tardieu-Blot, V, Mi (A. decipiens Evrard) Alangium chinensis (Lour.) Rehd., V, MM Alangium kurzti Craib Alangium salvifolium ( (Lf) Wang., SL, Lp AM eo Achyranthes as ,V,H Achyranthes nan BL, V,H Alternanthera sessilts (L.) R. Br. ex Roem., NL, Th ow Amaranthus ae L., NL, Th Celosia argente h Celosia argentea L. var. cristata Mog., NL, Th (Celosia cristata Miq.) Cyathula prostrata (L.) Blume, NL, ¢ Deertngia amaranthoides (Lam.) Merr., V, Lp ANACAR Reins duperreanum Pierre, V, MM Drymicarpus racemosus Hook., V, Mi _ sarmentosa (H. Lec )H Hand.- Maz., V, Lp Phlebochiton sarmentosum FH. ee 1d cucphuong 757s Dat, NE ee Pistasta weinm vanifolia Poiss. ex Franch. var. mplex Wu & Ifang., SUM, Mi kin chinensis Muller, V, Mi (Rhus semtalata r.) bn d7. q “ilavis (Roxb.) Burtt. & Hil VM Spondias lakonensis Pierre, VW, MM Spondias pinnata (L f.) Kurz, V, MM (S. mangifera W1 Toxicodendron rhetsotdes (Craib) Tard., V, Mi Toxicodendron succedanea (L.) Moldenke, V, Mi ANNONACEAE Alphonsea boniana Finet & Gagnep., SL, MM Alphonsea squamosa Finet & ae SL, MM po tonkinensis ae ., SL, Mi Alphonse Annona reticu es ae, Mi Mi Annona squamosa V, Mi 739 Artabotrys vinbensis Ast., NL, Lp Dasymaschalon rostratum vee & Chun. var. glaucum (Merr.& Chun) Ban, NL, Mi (D. elaucum Merr. & Chun) Dasymaschalon sp., , Mi Desmos cambodicus (Pierre) Ast., NL, Mi Desmos cochinchinensts eae NL, Lp Desmos dubius Hook., NL, Lp Desmos latifolinm 2a NL, Lp Goniothalamus tamtrensis Pierre, NL, Na Fissistigma elaucescens (Hance) Merr., NL, Lp Fissistigma laa Hook., NL, Lp Fissistigma maclurei Mert., NL, Lp \Tsiang & Li, NL, Lp = < ~ SF Po wn > eS = = = me J O8 = - 7 a~s = baa Q = = = s o -, Lp ct & Gagnep. NL, Mi Mi Mailinsa ee iene Pie rre Milinsa filipes b, V, Mi Palaiavnsanuine Mai V,MM Polyalthia laui} Mer ’,.MM he ernie V,Lp Uvaria cordata ( Dun.) Wall. ex Alston, V, Lp (U. macrophylla Roxb. ) Uvaria tonkinensis Finet & APIACEAE (UMBELLIFERAE) G ene astatica (L.) Urb., Crtdinm monniert (L.) Cuss. Eryngium foetidum L., NL, . & Gagnep., V, Lp Hydrocotyle javanica The inb., Hydr acotyle ee Lam., APOCYN Alstonia ery L.) R. Br., SL, MM Alyxia divaricata Pitard, SL, Lp aes ws ie Wall., SL, ees ertostylis Picard var. baviensss H Picard, SL, Lp Curbera ss Es Ecdy santhera rosea Hook.. , Lp (P ee Pichon, V, Na (Tabernaemontana bontt Picard) Sonne bovina (Lour.) Markgr., WV, Na (7 T-repoensis Pierre ex Piti ird) . & Flem.) Ervatamta bontt tonkinensis Pitard, ‘ Holarrhena antidysenterica (Roxb Hilepohena mili Craib., NL, Mi Kitabalia macrophylla Pierre) Woodson, V,Mi Kitabalia microphylla (Pitard) Woo odson, SL, Mi Kopsia tonkinensts Pitard, SL, Na Melodinus annamensis Piced: SL, Lp 740 Stipa 17(4) Melodinus ertanthus Pitard, SL, Lp : ynanchum corymbosum Wright., V, Lp Melodinus tonkinensis Pitard, SL, Lp Dischidia acuminata Cost., SL, Ep Plumeria acutifolia Poir., SUM, Mi Ses alboflava Cost., SL, Ep Parsonsita spiralis Wall., SUM, Lp Dischidia sp., SL, Ep Pottsta cantonensis Hook., SL, Lp Dregea sp., V, Ch Pottsta laxiflora (Blume) icine, SL, Lp Gymnema latifolta Wall., SL, Ch Rauwolfia cambodiana Pierre ex Pitard, V, Mi Hoya bonti Cost., ip Rhynchodia capusii Pierre, V, Lp Hoya ea Binime: SL, Ep Spirolobium cambodianum Baill., SUM, Lp aya vitlose a Cost., SL, Ep Stroph. — divaricatus (Lour.) Hook. & Arn., Hoya § ip NL, Mi (8. cdrvergens Grah.) Ait a ee Cost. + SL, Ch Wrrehtia annamensts Dub. & Eberh., NL, Mi Marsdenta tinctoria Br., SL, Ch Wrightia laevis Hook.f., SUM, Mi W. balansae Marsdenta nrceolata Deon, SL, Ch Picard) Myrropteron sp., SL, C Wrightia eae Gamble, SUM, Mi (W. Oxystalma esculentum Br., SL, Ch stellata rd) Secanione bonti Cost., SL. Ch W aa tomentosa (Roxb.) Roem. & Schule., Streptocauton griffithii Hook., NL, Ch SU Telosma cordata (Burm. f.) vee , NL, Ch W babi nen (JJum.) Pitard, SL, Lp Loxocarpus griffithii Decne, NL, Lp AQUIFOLIACEAE Toxocarpus hosseussii Schlechter, NL, Lp Toxocarpius villosus Decne., D Ilex cinerea Champ., V, C 1“ eal I De ’) : lis : oxocarpus vitlosus Decne., var. brevistylis Cost., Lex memecylifolia C ki amp., V,M eee Ilex purpurea Hassk. var. nervosa Loesn., V, MM Ilex rotunda Thunb., V, MM ASTERACEAE ARALIACEAE Adenostemma viscosum Forst. pM Ch Aralia armata (Wall. ex G. Don) Seem., NL, aaa nae ; bs ns cart ‘flora Wall., Th ; a Artemisia vulgaris NL Ps 4 y, Fe Aratlia chinensis L., ; ; . Aster trinervins ask. NL, i h ., Th fi Aralia thomsonit C.B. C lark var. thomsonis, SL, Ch var. petrolulosa Ha Aralia vietnamensts He. SL, Na Brassatopsts clomerulat fa (Blume) Regel., SL, Mi Brassatopsis bainla (Ham.) Seem., SL, Brassatopsis varianalis Shang, V, Na Heteropanax fragran (Roxb.) Seem., SL, MM Macropanax oveophilus Miq., SL. Mi Schefflera globulifera Grushy. & Svat SL, Na Schefflera glomerulata Li, SL yo Schefflera lencantha R. Vig., sk, Ap Blunea pr is DC. NI Ch Seiniiee nieiapolee Hacc, Blames pubreera Merr., NL. Ch (B. chinensis (L. Schefflera octophylla (Lour. a ae arm. < , Mi ae Schefflera petelotit Merr., Schefflera pes-avis R. ae Na Bike S/ i elie te sen ere Shotts donbinenae Re. Vig., SL, Na VoMmOLdena NOdT flor HIN ( ch.-Bip. ,Ch es on canes Cae. V. Mi Chromolaena odoratum (L.) King & Rabincen: Blumea os anne adie ee , Ch (5, tocephala | wn ct Z. Blumea sp., NL, Ch ARISTOLOCHIACEAE C a reves1i (Wall.) Sch.-Bip., V, Ch Aristolochia aut dongnatensis Pierre, V, Lp Cnicus chinensis Maxim., NL, Ch Aristolochia sp\., V, Lp Crreus japonicus Maxim., NL, Ch Aristolochia V, Lp Conyza viscidula Nee NL, Lp Aristolochia sp3., V, Lp Cotula anthemoides Lou L,C ASCLEPIADACEAE Crassocephalum epidloiae (Benth.) S. Moore, Asclepias curassavica L., NL, Ch NL, Th Congronema sp., V, Lp Crepis bonit Gagnep., NL, Th Cryptolepis buchananii Roem.& Sch., V, Lp Crepis japonica Benth., NL. Th Tun, Cucphuong National Park, Vietnam ae Ne ee (Lam.) DC., NL, Th Eclipta alba H Elephantopus hee Cents V,Ch Endydra fluctuans Lour., NL, H NL, linm indicum L., NL, Th m ee L., NL, Th Frangea maderaspatana Poir., NL, Th Gynura procumbens (Lour.) Merr., NL, Cr (G Savmentosa Gynura pseudo te DG. Gynura sp., Hemistepta rat Bung NL, Th Inula lappa D L, Th Lactuca indica h Partheninm cup L., V, Th Pluchea indica L, H Rhynchospermum se re Reinw., NL, Ch De JC Evigeron canadense L NL, Cr 3 n > Senecio scandens D. n, N Vernonia arborea Ham., NL, Vernonia cinerea Less., NL, Th Vernonia divergens Benth., NL, Ch Vernonia patula M Th Vernonia thorelit Gaseep. NL, Ch nsis Gagnep., NL, Ch NL, Ch NL, Ch Vernonia tonkine ie Nanthium strumarium L., Wedelia calendulacea Less., Wedelta urticaefolia DC BALANOPHORACEAE Balanophora fungosa Forst. & Forse. f., VW, Pp (B. pierre: H. Lec.) Bal ene sp., V, Pp BALSAM aes bonti ok f.,V,Ch kf Ch Impatiens aff. verrucifer as V.Ch BASELLACEA Basella rubra L., NL, Ch BEGONIACEAE Begonia sp} Begonia sp2., y. Ch Begonia sp3., V, Ch ,V,Ch Begonia sp4. Be egonid ee V,Ch B CEAE Pe collignonit (Dop) Steen., V, MM oles aes collignonis Dop) Fernandoa sp., V, MM faa ae Soem., V, MM Oroxylon indicum (L.) Vent., VW, MM Radermachera boniana Dep, VM urz) Steen ee jenea (K , Vv, MM Mayodendron igneum Kurz) BIXACEA Bixa 8 L., NL, Mi BOMBACAC Cae ae icum (DC.) Merr., V, MM BORAGINACEAE Bothriospermum tenellum Fisch., V,Th Cordia grandis Roxb., NL Mi Carmone microphylla ( (iain: ) Don, V, Na Ehretia hanceana Hemsl., NL, Lp Mi Heliotropium indicum L., NL, Th Tournefortia sarmentosa on V,Lp BRASSICACEAE Cardamine hirsuta L. var. silvatica Link., NL, H Rorippa indica (L.) Hiern., NL Rorippa masturtilin- in ae (L. Hay ek., NL, Th BUDDLEJ ACEAE Buddleja asiatica Lour., NL, Na BURSERACEAE Bursera tonkinensis Guill., SL, MM C anarium album Raeusch. , SL, MM , SL, MM (Cc. vittatistipulatum Guill.) Canarium tonkinense Engl., Canarium tramdenum Dai & dik, SL, MM Garuca pinnata Roxb., SL, BU XACEAE Buxus wallichiana Bail CAMPANULACEAE Codonopsts celebica (Blume) Thuan, V, Cr Codonopsis javanica (Blume) Hook.f., V, Cr h SUM, Ch Lobelta chinensis Lour., V Pratia begonifolia Lindl., V, Cr CAPPARACEAE Cappe vis viburnifo lia Gagnep .Y¥, Lp Capparis viminea EHOOR V,Na Capparis spl , Lp e aw Lp Capparis sp2 742 Capparts sp3., V, Lp Cleome gynandra L., Cleome pent. a DC. ONL, Th Cleome sp h Crataeva magna mc our.) DC., V, MM Suxvs ovata (Korth.) Hall. f., V, Lp CAPRIFOLIACEAE Lontera dasystyla Rehder, V, Lp Sambucus javanica Reinw. ex Blume, V, Na Blume, V, Mi (VY. Jiburnum lutescens colebrookianum Wall, Viburnum Ss). CARDIOPTERIDACEAE Cardtopterts lobata Wall., V,Ch CARICACEAI Carica papaya L., CARYOPHYLLACEAE Drymarta cordata Willd., V,Th Malachinm aquaticum Fries, NL, Ch va berth Poi V,PhH / Blt Vv, Lp (C ns oa. O. Caen CELASTRACEAE Celastrus geinata Loesw., V, Lp Celastrus hindsti Benth. & Heck .V, Lp Celastrus orbiculata Metrr., V, Lp Celastrus tonkinensis Pitard, V, Lp Celastrus sp., V, Lp Euonymus mitratus Pierre, SUM, Enonynues att. nitidus Rent. SU 7 Mi Exonymus sp., SUM, Mi Glyptopetalum gracilipes Pierre, SUM, Na Glyptopetalum eli Pieard, SL, Na Hippocratea indica Willd., SL, Lp Hippocratea sp 7 CERATOPHYLLACEAE Ceratophyllum demersum L.., CHENOPODIACEAE G oo album L. NL, Cr = Th .NL, Th ,NL Chenopodium ambr er L. CHLORANTHACEAE Chloranthus brachystachys Blume, V, Na Sarcandra glabra (Yhunb.) Nakai, V, Na CLUSIACEAE Callophyllum balansae Pitard, SUM, MM Callophyllam bonit Picard, SUM, MM Calloph i (Lour.) Blume, NL, Mi (Cratoxylon eae Korth.) yer, NL, Mi Cratoxylon formosum Jack.) Sipa 17(4) ssp. Hii (Kurz) Gagnep. (Cratoxylon prunifolium Dyer.) Cratoxylon sp., 7 Mi Garcinia cowa ce a MM Garcinia sp1., Garcinia sp2., SL, ve > Hypericum japonicum Vhanb., NL, Th Hypericum sampsonii Hance, NL, Ch COMBRETACEAE Anogerssus acuminata (DOC.) Guill.& Perr., V, M 20 al ronkinensts Gagnep. ~~ mibretum latifolium Blume, V, Lp (Combretum extensum Roxb.) Combretum punctatum Blume, V, 1 squamosum (G. Don) Exell. squamosum Roxb.) (Combretum a alts indtce 7L.,V,Lp valia myriocarpa a ot V,MM CONNARACE Cnestis pe as Merr. ssp L . palala Vidal Connarus Lees Roxb. var. paniculatus, NL, Vi fom ip f. tonkinensis Vide Rourea minor (Gaerth.) Leenh., NL, monadelpha (Roxb.) Vidal CONVOLVULACEAE Argyreta capitata (Vahl) Choisy, NL, Lp Areyrera obtusifolia Lour., NL, Lp Lp ssp. Argyreta — Boiss., NL, Lp Argyreia § ., Lp Cuscuta Aphis Pearson, NL, Pp Vv, Ch (L.f.) Kuneze, NL, Th (L.f.) Kuntze, NL, Th (H. ee ees lexittia sublobata bicolor N Wig 1) Ipomoea cairica (L.) Sweet, NL, pulchella Rot th) Ipomoea pes-tigridis L., NL, Ipomoea pileata Roxb., NL, lpontoed § sp Lp (Upomoea ez ., Lp Ipomoea sp2., NL Hallier f., Convolvilus parviflorus Vahl) Lapitepon binectariferum (Wall.) Kuntze, NL, Th Lp Jacquemontia as (Burm.f.) NL, Lp ie ener chin crac ) Hallier f., NL, Lp (Ipomoea chryset ) Merremta ae « hos) Bue f., NL, Lp (pomoea petaloidea Choisy Merremia vitifolia (Burm.f. ) Hallier f., NL, Th (pomoea vitifolta Sweet Nexropeltis maingayi ee ex Oost., NL, Ch (Neuropeltis racemosa Wall.) Turn, Cucphuong National Park, Vietnam Operculina turpethim (L.) S.Manso, NL, Lp (Ipomoea turpethum BR. Br.) CRASSULACEAE Bryophyllum calycinum Salisb., SUM, Ch CUCURBITACEAE Gomphogyne bonii Gagnep. Gymnopetalun eae: (L our.) Kurz, V, T Gymnopetalum ch etal ) Kurz, V, Th (Gymnopetalum monoicum Gagnep.) Gynostemma pentaphy ‘lla (Thunb.) Makino, V, T rynostemma pedata Blume) h Gynostenma sp., V, Hodesonia Macrocay pa (Blume) Cogn., i‘ C Mukia maderaspatana (1.) Roem., V, Th (Melothria maderaspatana Cogn.) Momordica cochinchinensis gece ,V,Ch Sobena a ae our., V, Th iis heterophylla Cogn.) Thladiantha cordifolia (Blume) oe V,Ch Thla diantha spiel Clar raib, V, ket ase ees th f tAALANLRA Sto dade + Gagnep, Trichos gera Blume, V, THC: himalensis , NL, H Crotalaria ee Grah., NL, Ch Crotalaria juncea CACY a Crotalaria linia | LE var. Waites NL, Th Desv. Crotalaria verrucosa L., NL, Na Datlber we balansae Prain, v, MM Dalbergia lanceolaria LE., ae MM Dalbsn ‘eld ViMmosa see V,Na Delonix regia , NL, MM (Potnciana regia i 3 oa Bojer ex Derris albo-rubra Hemsley, SL, Na Derris elleptica Benth., SL, Lp Derris i ne Gagnep., = — Derris wallichiana Prain, SL erris ee es Wall., NL, Na Yesmodium elegans (Lour.) Benth., NL, Ch Desmodium gangeticum (Lour.) DC., NL, Ch = Desmodium gyrans DC., ,Ch ~ Desmodium gyroides D De medium vp a ) a NL, Lp De ( (Willd ] i) DC f Desodian m taifliom DC. var. virgatum Mies : ee et Lita DC., NL, Ch Desmodium polycarpum (Lam.) DC., NL, Ch Desmodium pulchellum (1..) Benth., NL, Ch Desmodium triquetrum (L.) DC Dolichos sp., NL, Th Entada A eee (L.) Merr., V, L tonkinensis Gagne Erythrina stricta ne V,MM > Cf — Ch Crotalaria striata DC., NL, Na (ML. mucronata ) .. NL, Ch dium lasiocarpum (P. Beauv. )DC.. NL, Ch Ery th rophloe one fordlt Oliv., V,MM Fleming congesta Roxb., NL, Na Galactia longipes Cains V, Lp (G. mouretis yagnep. Gleditschia pachycarpa Bala, V, MM Indigofera cele — ,V,Na Indigofera aL., rc nde ir vii . NL, Ch Indigofera i Lepedeza sericea ve , 5UM,N 7 leach Se ) De Wit, NL, Mi ee (Willd enth.) . dielsiana Lee ex Diels, SL, Lp M Milletia pachyloba a N Milletic a pulchra Kurz, NL, Na M Pp Mimosa invisa vere ex C olla, V, Ch Mimosa pudica L. Mucuma es sas Ee yea, V, Lp Peltophorum ladle (Mig. ) Kurz var. tonkinensts (Pierre) K. & S.S. Larsen, V, MM (P. tonkinense Gag nep.) Prerolobium microphyllum Maiq. platypterum Gagnep.) Saraca dives Pierre, V, Mi Psophocarpus eee (L.) DC., NL, Th V, Lp (P. ?yeraria thomsonii Benth., p Pueraria montana au) Mere. NL, Lp (P. tonkinensis Gagne ep. ’,Ch Tephrosia att. Rerriz eee &C deg NL, Na Lephrosta purpurea (L.) Pers, Uraria crinita (L.) Desv., NL, Ch Uraria lagopoides DC. Uraria macrosta ee Wall., L, Ch Uraria rufesce a Cc.) cL. NL, H (U. hamosa Wall. Vigna radiata oe aon var. sublobata (Roxb.) rdcourt, NL, Lp (Phaseolus sublobatus ww Roxb. Viena ia (Thunb.) Ohwi & Ohashi, NL, Lp (Phaseolus calcaratus Re Vigna ,N , NL, Lp Viena sp FAGACEAE Castanopsis indica A.DC., V, Castanopsis lecomtet Hicke L& ch. Camus, V, MM Castanopisis symetricupilata Luong, SL,MM 746 Lithocarpus areca Hickel & A.Camus, Uy MM Lithocarpus cornews (Lour.) Rehd., Lithoc ge psendosundaica eae & . i amus, Lith a aft, stenopws Hickel & A.Camus, NL Lip aff. thomsonts Hickel & A. Camus, maa spl., SL, MM sion sp2 5UM, MM Quercus att. Dried Hickel & A. Camus, SUM an SUM, Mi FLACOURTIAC Jase la ene sels Merr., Flacourtta balansae Gagnep., Vv, Mi V,MM Haweantiaa paniculiflorum ow & Ko, V, Mi Homalium sp., V, Mi Quercus sp., io Fla a ratmonchy UL Herit., J MOC ry bis Da tna nenst (Merr.) Sleum., Vv, MM Hydnoc oe kurzti (King) Warb., ¥, MM ssp. australts Sleum. ’.MM V,Na iDsrxetonenn sp., \ Scolopia chinensis Clos., GENTIANACEAE Exacum tetragonum Roxb., C. obovata ACh V, Lp ( FPagraea cetlantea oa Wall.) GESNERIACEAE Boea martini Lévl., SL, Na Boew umbellata Drake, SL, Ch Chirita ona ie SL, H Chirita hamoea , SL, Ch aeaen eee srmensts (Oliv.) Stapt, var. D Didissandra a ot ‘Dr ses V, Didissandra sp., V, Rhynchothecum ae m Blume, V, Ch HAMAMELIDAC Liguidambar eae Hance, NL, MM HERNANDIACEAE Iligera celebica Miq., Dunn.) Ulivgera rhodantha Hance var. rhodantha, V, Lp vl.) Kubiczki (U//igera glandulosa —. Vi. Lp CL. platyandra var. dunniana (Le HIPPOCASTANA Aesculus assamica ¥. aoe ,NL, Mi HY DRANGEACEAE Dichroa hirsuta Gagnep., V, Na Sipa 17(4) HY DROPHYLLACEAE Hydrolea zeylanica Vahl, V, Th ICACINACEAI Gomphandra aes Pierre, V, odes cirrhosa Tare JHU. a or = ysp., V, thapod ides (Oliv.) Sleum., V, Mi st sp., v, Mi ILLIC ie 1M oe | var. cambodianum Finet & Gagnep., SUM JUGLANDA Annanmocary aes asiy Dode, ae MM E ngethardtia sat B. ume, V, M Ph 7 Sieb. & Zuce., SUM, MM ane ares Dode, V, MM LAMIACEAE Ayuga decumbens Thunb., V, Ch Alisomoles indica (1..) ue V,THCA. ovata _ Br.) Gomphostemma leptodon Dann, V,Ch Gomphostenma lucidum Wall. & Benth., S.Y.Hu, NI V,Ch Leonurus artemista (Lour.) ., Ch (L. heterophyllus Sw.) Leucas mollissima Wall. ex hoe. te Ch | Leucas zeylanica (L.) Br., Mentha arvensis L., NL, H Mentha aquatica L.., NL, H Microtoena imsuavis (Hance) Dunn, NL, Th Mosla cavalertei Levl., V, Th Ocimum basilicum L., NL, Th Ocimum sanctum L., NL, Th Orthosiphon thymiflorus (Roth) Slecsiin, V, Ch (O. fomontosus Benth.) Paraphlomis javanica (Blume) Prain, V, Ch (P. rugosa Prain) la fratescens (L.) ,NL, reli ie - . un Pogostemon auricularius (L.) Hassk., : Th Salvia plebeta R. Br., V,T Salvia splendens Ker. Gowl., NL, T Stachys oblongifolia Wall. ex erp 7 I Tencrinm viscidum Blume, V, H (1. stoloniferum Roxb.) LAURACE Actinodaphne pilosa (Lour.) Merr., MM Beilschmiedia att. eae Hayata, V, MM Betlschmiedia fordii Dunn., V, MM Beilschmiedia ferruginea H., Lin, V,MM Bertlschmiedia laevis Allen, V, MM Berlschmiedia obovatifoliosa Lecomte, V, MM Beilschmiedia robusta Allen, V, MM Turn, Cucphuong National Park, Vietnam Beilschmiedia ey aes Nees, V, MM Betlschmiedia spl., Beilschmiedia sp2., V Beilschmiedia Pets Vv, a Betlschmiedia sp4., V, MM lai a i. baviensis (Lecomte) Airy Shaw, V, MM (Nothaphoebe att. baviensis eco cared tonkinensis (Lecomte) Airy Shaw, V, MM (Nothaphoebe tonkinens1s Lec Cassytha ae rmis L., NL, P Pais mum balansae et te, V, MM Cinnamomum i (Buch.- Ham.) Sweet, ,MM(C. I ( btusifolinm Nee nee Pio 71 Lecomte, “ MM Cinnamomitin es Lecomte, Dog MM Cinnamomum camphora Sieb., : Cinnamomum curvifolinm (Lour.) Kosterm., V, MM Dingle as yl Ne me ur.) Koscerm., V, M(C Lara Thw.) Cinnamomum tetragonum A, Chev., V, MM Cinnamomum validinerve Hance, V, MM Cryptocarya Helens Miq., V, MM Cryptocarya lenticella geet. V,MM Cryptocarya coi Blume, V, MM : ryptocavia § ndera balansae icone V,MM ne polyantha Boerl., V, MM Lindera tonkinensts eae Vv, Mi Litsea baviensis Lecomte, v; Na Litsea cubeba Pers., SL, Litsea aan ie )L. Rab, SL, Mi tea hefferi H V,Na i mollifol ta Can. Vv. Mi(L. mollis Hemsl.) Litsea monopetala Roxb., Vv, MM Litsea panamonja (Nees) Heals f., V, Mi Litsea rotundifolia Hemsl., V, Litsea umbellata (Lour.) Mets. - Mi(L. amara Blume) Litsea verticillata Hance, V,MM Litsec is sp2., V a Machilus bombycina King, V, Mi Machilus hainanensis Merr., V, MM Seite nica Nees, V, MM rpa Chun, V, Mi Heaney ee ‘gli Sieb. & Zucc., Machilus sp., V, MM eens caudatum (Nees), e? H.Liu) V,MM Vi, Mi (N. poilan Neocinnamomum delavayi (Lecomte) H.Liu, V, a(N. parvifolinm (Lecomte) H.Liu) Neocinnamomum lecomter H.Liu, V, Mi 2 ~ LOG 747 Neolttsea zeylanica Metrr., V, Phoebe tavogana (Meissn.) one f., V, MM (P. cuneata Blume) LECY THIDACEAE Barringtomia eberhardtii Gagnep., V, MM Barri 22 etomia edaphocarpa G ragnep., ¥, I LEEACEAE Leea guineensis G. Don, V, Na (L. acuminata Wall.) LINACEAE Tirpitzia chinensis (Hemsl.) Hall., SL, Mi ANIACEAE Strychnos wallichiana Steud.ex DC., V, Lp (S gaulthierana Pier LORANTHACEAE Elyt th, aopytt, 2, 4G. Don, V, Pp s Lecomte, V, Pp Loranthus pentandrus L., V, Pp Loranthus ferruginens ae ,V,Pp ] a nthus 4 /pressz, Lf Loranthus seurrulla L., p villus chinensts (DC. . Dae. V,Pp te estipitatus (Stapf.) Dans., V, Pp LYTHRACEAE Ammarid auric ulata Willd. var. arenaria Sochne, Ammartia bacctfera L, Th Dratane ee pee ex DC.) Walp., V, rstroemia corniculata Gagnep., Mi it rotundifolia Kochne., NL, F MAGNOLIACEAE Magnolia eriostepta Dandy, SL, Mi Manglietia conifera D eee he MM Mi helia balansae Dandy, V, MM Michelta eae en SL, Mi MALPIGHIAC Aspidopteria aft. as ifolia J. nl V,Lp Aspidopteris obcordata Hemsl., V, Lp iptage boniana P. Dop, V, a Hiptage bol atensis (L.) Kurz, V, Lp ee sp., V, Lp MALY dls indicum G. Don, NL, Ch ,Na Urena lobata \.., V, Na 748 SipA 17(4) MELASTOMATACEAE Ficus auricula Lour., V, Mi CF roxburghit Wall.) Blastus cochinchinensis Lour., VW, Na Ficus benjamina 1.., ue MM Blastus multiflorus (Cogn.) Guill. V,Na Ficus callosa Willd., _M Melastoma pied D. Don, NL, Na Ficus callophylloides aa V,MM Melastoma normale | on, NL, Ch Ficus capillipes Gagnep., V, Mi Melastoma rem Sims, V, Na (AL. Ficus chaffansjont Lev., v. p decemfidum Roxb.) Eicus cunta Hamilt., V, Mi Melastoma trungit Pocs. & Tiep, V, Na Ficus curtipes Corner., V, MM Osbeckia truncata D. Don, V,H Ficus dumosa King, V, Na Freus fistulosa Reinw. ex Blume, V, MM (f harlandti Benth. Fic is fulva Reinw. ex B “Ki MELIACEAE Aglaia gigantea (Pierre) Pellegr, V, MM Aglata odorata Lour., V, Mi Aglaia tetrapetala Pierre, V, MM Amoora aphanamixis m. & Schult., V, Mi (Avlata aphanamtrt elles er, var. Faces Cc € £ ' ume var. flea, V. Mi : Ficus glandulifera Wall., V, Mi sie Riou hegpapiulle | Amoora att. cucullata Roxb., V, MM teil aged : Ficus hirta Vahl, NL, Na Amoora sp., V, Mi fans bbede 1. VM chi ! ' elo Pj V, ‘icus hispida i MSOCHELON LiODOSIS ee, Pieus lacor Hi sail »VMM (FF snfectoria Roxb.) mn pantculatus Hiern., v. aan C saa tabularis A. Juss., V, M Cipadessa fruticosa Blume, V, Mi ' ica sisi lal Hooke f., V, MM eee Tera. MME Fieus leckensis Drake, V, Na 2 caultflos Ber aeons ; Dynal aff, eee om V,MM Ficus nervosa Heyne, v ; MM ; Ficus obscura Blume, V, Mi Heynta trijuga Roxb., MM ; : Vv, Ficus pilosa Reinw., V, MM Ficus pumila L. ap Lp Fieus racemosa L.., V, MM (EF. glomerata L. var. Views laevis Blume, V, Lp Ficus lamponga Mig., V, Mi (F. balansae Melia azedarach L., V, MM Toona surent (Blume) ee V,MM Walsura bonis Pelle MM MENISPERMACEAE Freus religtosa 1.., NL, MM Cassampelos pareira L., V, Lp Ficus retusa L., V, MM Cyclea tonkinensis Gagnep., V, Lp Pies rostrata Lam., V, Na Cocoulus laurifolius DC., V,C Ficus sagittata Vahl, V, Lp (F ramentacea Roxb.) Diploclisia macrocarpa Miers, V, Lp Ficus stkkimensis Mig., V, Lp Fibraurea recisa Pierre, V, Lp Ficus sum Gagnep., V, Mi Parabaena sagittata Niavs: Vv, ee Ficus superba MiGs, Vv, MM Tinomiscum tonkinensis Gagnep., VW, Lp Piens variegata Blume, V, Mi Stephania hernandifolia Spreng., . Lp Ficus sp1., V, Lp Stephania rotunda Lour., SL, Cr Ficus sp2., V, MM Malatsia scandens (Lour.) Planch., V, Ch (MI MORACEAE Antiaris toxicarta Leschen., V, MM oa heterophyllus Lam., NL, Artocarpus parva Gagnep., V, Artocarpus petelotii Gagnep., V, MM Artocarpus tonkinensis A. Chev., V, MM tortuosa Blume var. scandens Bur, Morus alba L., NL, Mi Streblus asper ane a Mi Streblus tlierfolia (Kurz) Corner, SL, Na ee iicifolta ( (Kurz) Vidal) Streblus taxordes (Heyne) Kurz, SL, Na (Phy ocblan mys tridentata Gagn Laxotrophis macrophylla (Blume) Me igen Streblus toukinensis (Eberh. & Dub. sCannick SL, Mi ollssenetla pe py thera ent. Maclura bocin Nneads iieuns Corner, V, Lp (Cudvanta cochinchinensis Lour., C. favanensis a Mi (Leonongia tonkinensis Stapf.) Dimerocarpus brenniert Ge agnep., V, MM MYRISTICACEAE Frieus altissima Blume, SL, MM Horsfieldia prainit Warbg., V, MM Ficus annilata Blume, SL, MM Horsfieldia tonkinensis Lecomte, V, MM THIN, Cucphuong National Park, Vietnam Knema conferta (King) Warbg., V, MM Knema globularia (Lam.) Warb.., V, K. corticosa Lout.) MM MYRSINACEAE Ardista maclosa Mez Ardisia ramondiaeformis Pitard, V, Na Ardisia silvestri Pitard, V Ardisia tonkinensis A. DC., V, Na Ardista s spl., V, Na Ardisia sp8., V, a Embelia laeta L., N p Embelia sblineyolea Hemel SL, Lp Embelia ribes Burm., NL, Lp Enbelia scandens ides, SUM, Lp Embelia ‘Pl . V, Lp Embelia sp2.,V, Lp Maesa ye Mez Maesa indica fost ) A DC. ,V,Na Maesa japonica (Thunb.) Mez, V, Na Maesa striata Mez var. opaca Pitard, V, Na Masea tenera Mez, V, Na Maesa tomentella Mez, V, Maesa tonkinensis Mez var. eee V,Na var. bonii Pitard pee EAE mum panicilatum Kurz, V, Mi eee guajava L., WV, Mi Rhodomyrtus tomentosa Wight, NL, Syzygium baviense (Gagnep.) Merr. & Perry, V, Mi (Exgenia baviensis Gagnep.) Syzygtum chlorantha (Duthie) Merr. & Perry, Vv, Mi (Exgenia chlorantha Duthie) Syzygium eburneum (Gagnep.) Merr. & Perrr V, Na (Eugenia eburnea Gagnep., ae dhrmisnins Merr, & Perry) Syzygium jambos (1.) Ast., NL, Mi (Eagenza jambos L.) Syz rygium malayanum (Gagnep.) Merr.& Perry, , Mi (Exgenia malayana Gagnep.) Syzyginm mekongense (Gagnep.) Merr. & Perry, Mi (Evgenia mekongense Gagn Syzygium samarangense (Blume) ie Perry, Mi (Syzygium javanica Lam.) xb.) Merr.& Perry, V, var. ternifolta Syz — rernifolinm (Ro Syzygium formosa Wall. oan “p.) 749 Syzygium tonkinense (Gagnep.) Merr. & Perry, M (Eugenia tonkinensis Gagnep.) Syz} ae zimmermannit (Warb.) Merr.& Perry, ™ NS Eugenia gimerani Warb. OLACACEAE Erythropalum scandens Blume, V, Lp ee adenophyllum Wall., es Jasminum ealan Roxb., Jasminum lo Jasminum undilatum Ker-Gawl. Lp ONAGRACEAE Ludwigia adnascens (L.) Hara, NL, H (Jussia0a repens cds.) Ludwigia Heise Roxb., NL, prostrata (Roxb.) L Ludwi igh octovalis Jac acq.) Raven ssp. sessiliflora (Michx.) Raven., NL, Th Uussiaea suffriticosa H CUussiaea OPILIACEAE pe te sylvestris Blume, SL, Na eliantha suavis Pierre, SL, Mi = OXALIDACEAE verrhoa carambola L., NL, sae Bip sensitivum — NL Ixalis corniculat NL, H PASSIFLORACEAE Adania chevatiert Gé ae V,Ch Passiflora foetida L. Passiflora stamica C eu Ch (P. octandra Gagn p.) PEDALI ane ak L., NL, Th PENTHOR E Penthorum or L., NL, H PIPERACEAE Peperomia leptostachya Hook. & Arn., V, Piper betle L., NL, a Piper bonit DC., V,C Piper gymnostachyum Be Ne Ch Piper hainanense eee Piper lappaceum ni yc Piper lolot D Gi Piper longun . , Ch Piper pada aaa. DC., V, Ch Piper retrofractum Vahl, V, Ch > 750 Sipa 17(4) PITTOSPORACEAE his theezans Brongn. NL, Lp Pittosporum tonkinensts Gagnep., V, Mi alyculata Val. var. macrantha Pitard, PLANTAGINACEAE Plantago major L., NL, Cr Vv, cs Ventilago letocarpa Benth., V, Lp Ventilago pauciflora Picard, V, Lp PLUMBAGINACEAE Zizyphus mauritiana ee ,V,Na Plambago zeylanica L., NL, Ch Ziz ie rm hc Mill., V, Mi POLYGALACEAE ne ue Polygala pa ttad m7 our., V, Na RHIZOPHORACEAE Polygala tonkinensis C bod at, V, Na Carallta . ita (Lour.) Merr., VW, MM (C, POLYGONACEAE Metre Rs gopyrum cymosum (Trev.) Meisn., V, Ch Carallia Doe Roxb., V, Mt Pade gonun ee L. “NL. H ROSACEAE Polygonum chinense L., NL, H Duchesnea indica (Andr.) Focke, Polygonum donit Meisn., V,Th Eviobotrya bengalensis (Roxb.) le . Ny, ae Polygonum flaccidum Roxb., V,H Eviobotrya japonica ( es ) Lind1., Polygonum hydvopiper Pragaria indica L.., Polygonum Dasa ileene Ase. .V,Th Persica vulgaris at ; Polygonum minus Huds., NL, H (P. posumbu Photinia benthamiana Hance, V, Mi Wall.) Prunus mume Sieb. & Zuce. a Mi Polygonum odoratum Lour. 2 Th Prunus salicina Lindl., Polygonum ortentale L., V, Prunus le Miq., - an (P. macrophylla Polygonum perfoliatum L., s Sieb. & Zu Poly ygonum plebejum R. Bi: v, f Pygeum spl., SL, a Polygonum tinctorium Lour. , Th Pygeum sp2., SL, MM Rumex wallichii Meisn. a ,Th Pyrus sp., SUM, MM PORTULACACEAE Raphiolepis indica (L : Lindl. ex Ker Portulaca oleracea L., NL, Th Kosa cymosa Trat., NL, Na - eset ipa pines) Rubus alcaefolius ne V, PRIMULACEAE Rubus cochinchinensis ie V,Lp Androsace saxifragifolia Bge.. V, Th Rubus leucanthus Hance, V, Lp Lystmachia candida Lindl., V, Ch Rubus triphylus Thunb., V, Ch Lysimachia spl., WV, Ch - Lysimachia sp2.,V,Ch RUBIACEAE Lysimichia sp3., V, Ch Adina pilitiflora (Lam.) Franch.ex Drake, V, Mi Lysimichia sp4., V.Ch (A. globiflora Salish. var, eles Pit Atdia cochinchinensis Lour., § PROTEACEAE Anthocephalus indicus A; Rich, var. nie Helicta cochinchinensis Lour., a Mi Pierre, V, MM Felicia att. erratica Hook., Canthium be occu (Gaertn.) Tinn. & Binn, Heliciopsts lobata (Merr.) a co Mi > M RANUNCULACEAE Canthium sp., V, an Clematis armandi Fanch., V, Lp Cephalanthus sp., Clematis meyniana Walp ~V,Ch Chasalia curviflora (Wa i ex Roxb.) Thw., V, Clematis snnee a Walp., V,Ch Na Clematis nmb era Gagnep., V, Lp Diplospora sp., V, My Clematis pide € hamp., V, Lp Duperrea pavettaefoli - es V,Na Ranunculus yaponicus Li angsd., V,Ch Cinders coronaria Ham., V, Na Ranunculus scleratus L., V,Th Gardenta florida L., V,H Geophila reniformis G. Don, V, Th REAMNSG pane : Hedyotis capitellata Wall. ex G. Don, V, MM Berchemta lineata DC., NL Padystis disiiiboia HV an i, Ne Chaydaia tonkinensis } Picard, = Mi . Hedyotis multiglomerulata (Pitard) Tran, VW, Na Hymenodyction excelsum Wall. var. elaeinean > V / A Gouanta leptostachya D 7, Lp Rhannus tonkinensts Pits avd. ViNa ne THIN, Cucphuong National Park, Vietnam ss raged Wall. var. flexi/is Pitard, V, Na AVSOnIANA Wall., Vv, Mi Ixora oe Lev., V, Na spl., V, Na ee hee V,Na [ s andamanicus Hook., V, Mi Le ii satan Jack. alli - Wight, V, Mi V,Na DS Lastanthus bee spl Lasianthus sp2., LY, Na Lasianthus , NL, Na Morinda ial L. var. tonkinensis Pitard, V, a 2 © we p Morinda villosa Hook., V, Lp Morinda sp. Mussaenda ie Pierre, V, Lp Mussaenda dehiscens Craib, V, Mi Mussaenda sp|., V, Lp Mussaenda sp2., V, Lp Mycetia longifolia ane ze, V, Na Myrionenron tonkinensis Pitard, V, Na Myrioneiron Zi ,Th Nauclea sp., Neonauclea sesst aii (Roxb.) Merr., V, (Adina sessiliflora Hoo MM Oldenlandia sp4., Onhoalenemarde Pitard, V, Th Ophiorrhiza aft. baviensis | V,Ch Ophiorrhiza att. tristzs Drake ,T Ophiorrhiza spl., Ss is sp2., Paederia scandens Conk ) Merr. var. tomentosa (Blume) Hand.-Mazz., Paederia sp1., V, Paederta sp2., V, Th Pavetta indica L. var. indica Hook., V, Na Psychotria baviensis Pitard, VW, Ch Psychotria rubra eee ) a V,Na va sp4., V, Randia acuminatissima em V,MM Randia spinosa (Thunb.) Poir., V, Na Randia stenantha Deke. SUM, Na Randa — ,V,Mi Randia sp2., V, Mi See an hae teats ae oe Na rpeanaaellinice' er sib, Vv, Tarenna quosensis Pierre & Pitard, V, Na Tarenna vanpruckii Craib, V, Na Tarenna spl., V,Na Tarenna sp2., V, +8 Tarenna sp3., V, Unearia macniihs 2 Wall., V, Lp Unearia sessilifructus Roxb., V, Lp Uncaria epeigs Havil., V, Lp Urophyllum sp., Nantoned alee Pieri, V,Mi Nantoned sp., Wendlandta ee DcC., V, Mi Wendlandia paniculata DC. V,Mi RUTACEAE A cron) hia pedunculat oxb., V ae Atalantia ouilloumini - ingle, Atalantia monoph: orrea, roxburghiana Hook. Atalantia dee a Hook.f., V, Mi Citrus lemon (L.) Burm. f., V, ne , Mi er Mi (A. Citrus maxima (Buen, f.) Merr., V, Mi (C. ( bt) Q z an 5S Q xX wo zs NLA = ~~ Y a Oe Q ~ = Q a ia) a Clausena excavata Burm Mi Clausena harmandiana Pride ex Guill., Vv, Mi / x cS) Clausena laevis Clansena lansium Skeels, v, Mi (Clausena wampi _B 3lanco) Oliv.) nodta jraxinifolia (Benth.) Hook., SL, MM Eadie lepta (Spreng.) Merr., V, MACE. triphylla ) Ewodta dines Benth., V, Mi Lindl., V, i Gh yoosmas aff. oe 7 Tanaka, V, Mi yeosmis pentaphylla Correa, V, Mi Glycosmis petelotii Guill. Glycosmis puberula Linc i ex tal, V, Mi yoosms aff. tr ps Guill., V, Mi r/ycosmis winiti Cra lum falcatum anak, v, ee Mucrome i irs, SLUM Oli lv, MM icromel]y, ie luni alan Craib, v, is Zanthoxylum nitidum DC ‘Lp Zanthoxylum scabrum Guill., V, Lp SABIACEAAE Meliosma harmandiana Pierre, V, Mi Moliosma thorelli Lecomte, V, MM SAPIND Alp ial eee ete a Mi Hophyl dadIk., Adiesbuka racemosis ra on. a Mi 752 Stipa 17(4) Boniodendron parviflorum (Lecomte) Gagnep., V, Physalis augulata L. var. aia Bonatt, V, Th i Solanum biflorum Lour., V, Na Cardiospermum halicacabum L., V, Th Solanum procumbens eee V, Lp (S. hainanense Dimocarpus longan (Lour.) Ste otk NL, Mi Hance) Euphoria fragifora Gagnep., V MM Solanum indicum L.,V,Na Lepisanthes rubiginosa (Roxb.) Leenh., V, Mi Solanum nigrum L., NL, Th (Evioglossum edule Blume) Solanum torvum Swi artz, NL, Na : ile jails Ri ads we MM Solanum verbascifolinn L., V, Mi f itt fe Lour.) Merr., VW, MM Solanum x a Schrad., V, Ch (M. he Blu ‘2 Nepheliim chrysenm Blume, SL, Mi Pavieasia annamensts Pierre, V, Mi Pometia pinnata J. & G. Forst., V, MM Pometia sp., V, MM Sapindus saponarta Lour., V, MM (S. nkorossi STAPHYLEACEA Lurpinia ee Merr., V, Mi Lurpinia nepalensts ae Vi Mi Turpini. 1 a DC. Turpinia spl., V, Mi Tunpinia sp2., V, Mi Gaertn.) Nerospermum donatense Gagnep., V, MM STERCULIACEAE Zollingera sp., SUM, Mi Abroma augusta L., V Bytinerta apera Colebr., V, Lp D cielo: Byttneria echinata Wall., V, Lp ine Simi: (King & Prain) Exell, V i Sy derox) lon Sp. ‘wmantia | inifolia L., V, tile nee L. : NL, Na Helicteres hirsuta Lour., NL, Na Heltcteres isora L. L, SL, Na Sinetenen lon racemosum (Pierre ex Dubard) Aubr., SL, MM (Sideroxylon racemosum Lecomte) SAURURACEAE Helicteres lancoat DC. ,Na Gynnotheca chinensis Decne., V,H Helicteres viscida Blume NI. Oe Houttuynia cordata Thunb., V, Ch Heritiera mi cs Wall. , MM Saururus chinensis Baill., Vy Cr Herttiera sp. nov, § Mit SCROPHULARIACEAE Melochia eae i L.; Bacopa floribunda (R. os Wertts., V, Ch ipa vanes Eee, Prerospermum meralocarpum Tard., Limnophila sessiliflora . NL, H es i &e ee ; ‘ 2 ai Lindernia antipoda (1..) Alston, NL, Th — a a nan veronic hoes Spreng.) / Be Ch Pterospermum has en SUM, MM - Prerospermum lancaefolinm Roxb., led teeneduneal sche wan mies Beane, Prerospermum venustim Craib, SL, MM NL. Ch Sterculia coccinea b., V, MM Mazus sp., NL, Th Prcria fel-terrae Lour., V, Th (Curanga amara Juss.) lorenta finettana Bonati, V,Ch veal ; : Sterculia radicans Gas mep., VW, Na lorenta tonkinensis Bonati, V, Ch eee ‘ ae H Sterculia tonkinensis Cannes. V,Mi Verontca anagalts L.., V SIMAROUBACEAE oT YRACACTAR Adlanthus triphyse ee nnst. a NL, MM Styrax tonkinensis Pierre, SL, MM (Adlanthus us, wrica DC SYMPLOC SAE a tonkinensis (Lecomte) ese V, Mi Symplocos Sie nsis Desv. ex Merr., V, Na (B. mollis Wall. var. tovkinensis Lecomte) Symplocos ee: Feuie V,MM Picrasma sp., V, MM THEACEAE SOLANACEAE Camellia caudata Wall., V, Mi Datura suaveolens ene & Bonpl., V, Th Camellta flava Cubans — V,Mi (D. fastuosa L. alba Car mellia tonkinensis (Pitard) Cohen-Stucort, V, THIN, Cucphuong National Park, Vietnam 122 Eurya trichocarpa Korth., V, Mi Villebrunea frutescens Blume, V, Na ) ; THY MELEACEAE Villebrunea frutescens Blume vat. nivea Gagnep., Wikstroemta indica (L.) C. A. Mey., SUM, Mi : renee VERBENACEAE Colona poilane? Gagnep., V, MM Callicarpa aborea Roxb., V, Mi (4 a ra + ‘ . - & Callicarpa candicans (Burm. ft.) Hochr., V, Na Corchorus acutangulus i , NL, Th (Cc : ‘ . cand Grewia bilamellata Gagnep., SL, Mi ee - ; me 7 - ienn Uenl cae . Callicarpa dichotoma Seamed, Reausch. var. Ula DISH E re ; Grewia tomentosa Juss., SL, Mi aaa via 10, 16 a ‘ i Vv, Cin pais NL, Mi ee albida Blume, Mi (C. longifolia Trinmphetta bartramia L., NL, Na (T. rhombotdea on one ) ete loureiri Hook. & Arn., V, Na ian beaeadana Soa Cab NT ha Callicarpa macrophylla Vahl, V, Na . -— eee Hook. & Arn., V, Na (C (LT. tomentosa Bojer.) ses all. € ULMACEAE bina ok Lindl., NL, Mi Celtis orientalis Thunb., SL, MM arias , Mi Celtis ari Mi te V, Mi erd subaequalis 7 V,MM eee ndrum coleby cs iin ‘Walp., V,Na meni ortentalts (L.) Blume, V, MM Clerodendrum cyrtophyllum Vurcz.., : Na URTICACEAE Clerodendrum sandichaudii P.Dop, Boehmeria holosericea Blame, V, Ch Clerodendrum kaempferi Jacq. Sib. E 7 Hh aes V,Na Boehmoeria nivea (L.) Gaudich., } p) Boehmeria nivea (L.) Gaudich. var. tenacrssima Clerodendrum lecomter P. Dop i 2p) ¢ c DP. (Gaudich.) Miq., V, Na Clerodendrum philippinum a vuer var. sinplex ,Na Boehmeria macrophylla Hornem., V, Mi (Bb. & Tfang., platyphylla Don) Cleradendrum serratum (L.) Moon., V, Na Bochmeria tonkinensis Gagnep., V, Ch Clerodendrum tonkinensts P.Dop, V, Na Bahinie Premna elon Sale f.) eat V,Na Pellionia np boties V,H (Pellionia daveauana P N.E. V¥;iNa Pellionia V.H Premne es as V,Na Pellionia sp2., V, H Stachytarphyta pamaicensis (L.) Vahl, V, Th Verbena officinalis L., V, Ch Prpiloitedla xe b., V, A ; . Pilea alongensis Gagnep., V, Ch Verecmpanelo LS Pilea ee Gagnep.. are Vitex quinata Williams, MM 4 peltata Hance, V,Ch Vitex stylosa P. Dop, SL, Pilea ee (Gaud ich.) Hook., VW, Ch ea petelotit C cee , Vitex sumatrana Maiq. var. “e King, SL, tripinnata (Lour.) Merr., SL, Mi (V. Pouz se hirta Hassk., V, H Pouzolzia indica ae V,Ch aban: Hall.) Pouzolzia pentandra Benn., V, Ch VIOLACEAI Pouzolzia sanguinea Blume) Merr., V, Ch (P. Viola inconspicua Blume, V, Ch viminea Wedd. rocris SP., ma, VISCACEAE : Viscum articilatum Burm. ae oe oe Villebrunea paso Gagnep., Villebrunea integrifolia Gaudich., V, Mi Viscum articulatum Burm. NS) , Pp 754 Sipa 17(4) Viscum orientale Willd., SUM, Pp Calamus prendoscutellaris Conrard, NL, Lp Calamus rudentum Lour., NL, Lp Calamus tetradactylus Hance, NL, Lp Caryota bacsonensis Mag., SL, MM VITACEAE Ampelopsts he ore ulate ee V,Lp Cayratia tenutfolta Gi varyota mitis Lour., V, Na ane saryota monostac bu Bete V,Ch Caryota sympetala Gagnep., V, Mi Licuala bracteata Gagnep. LY, Ch Licnata fatua Becc., V,Ch Licuala tonkinensis Becc., V, Ch Livistona saribus (Lout, Men & Chev., V, MM (L. cochinchinensis Mart. Pinanea baviensis Becc., V, Ch Rhapis micrantha Becc., SL, Ch Cissis mes Planch. var. swbintegra Gagne L, “Pp Cissus su ene Planch., ee Parthenocissus landuk ( Tetrastigma pach) hyllum ( (Hemel ne nc. V,Lp Lp etrastigma retinervium Planch. — Tetrastigna tonkinensts Cie. [ Lp Vv, Lp Tetrastignic : rE oa Vv E Rhapis laosensis Bec SL, Tetrastignia sp , Lp R/ ; ce or aie V,Ch eed ; an Rein vw. Me Cbs aladccd ee is ; CaALACCE wallich Mar LILIOPSIDA W allichia neal , V,Na ee ASPHODELACEAE Sagittarta sagittifolia L., NL, Cr € SM tum laxum R. Br., V, Cr AMARYLLIDACEAE thlorophytum orchidastrum Lindl., V, Th Crinum astaticum L., NL, Cr ASTELIACEAE Crinum enstfoliion Roxb. NL, Cr Cordyline fruticosa (L.) A.Chev., V, Na (C. ARACEAE termimalis Kanth var. ferrea Bak.) Acorus LV AMANEMS Solar id., BURMANIACEAE _ Gr Aglaonema modestium Scho E y ‘ ss : & Ee taro ttex Engl., V, A ( Burmannia coelestis G. Don, V, Cr sramense Engl.) Alocasia macrorrhixe ’.H ep Se tc Amorphophaltus hen (Denst. : Pe Canna edulis 1... SL, Cr var. campanulatus (Roxb.) Sivad., Cr (A. Canna nen Forks NL, Cr campanulatus Rox Canna indica L., NL, Cr Amorphophallus cirrifer Stapt., V. Cr COM ZAE Amorphophallus tonkinensis Engl., V, Cr ‘nan nudiflorum (L.) Wall., V Anadendron latifolinn He D0k.. ,V,Ep netlema medicum (Lour.) Kost., VW, H (A. Arisaema balansae Engl., V, Th sasnine Hance) Colocasta tndica Hassk., V, Cr Commelina benghalensi Homatomena occulta (Lour.) Schorr, V, H Commelina diffusa a : oe os (C. nudiflora Lasta spinosa (L.) Thw., V, Cr L.) Lasta sp., V, Cr C ones paludosa Blume, V, H (C. obliqua Pothos gigantipes S. Buchet, V, Ep Pothos kerrii 8. Buchet, V, Ep Fiisabs eat care issk., V, H Porthos repens (Lour.) Druce, V, Ep Floscopa ~ 1s Lo Pothos scandens L., V, Ep Floscopa sp. Pothos yunnanensis Engl., V, Ep Forrestia Sabra (Hassk.) Hi ae . H Raphidophora bonii Engl., V, Ep allia thyrsiflora (Blume) Endl, Raphidophora ee (Riss, ae V,Ep Rhaeo spathacea (Sm.) Stearn. — (Saige Raphidophora hookeri Schott, V, Ep color Hance) Typhonium divaricatum (L.) Decne., V, Cr Streptolirion sp. Typhonium trilobatum Schott, V, Cr Zebrina pe ae Aca .V,H ARECACEAE (PALMAE) CONVALLARIACEAE Areca catechu L., NL, MM Aspidistra tonkinensis (Gagnep. Wiles & Lang, Arenga pinnata (Wurmb) Merr., SL, Mi V, Cr (Collania a Gagnep.) Calamus platycanthus Warb., NL, Lp Aspidistra typica Baill., V, Cr THIN, Cucphuong National Park, Vietnam pons _ V,Cr Aspidist aa. ee Ce ib, V — Disporum angustifolium Roxb., V, Cr Dispos ‘um calcaratum D. ee var. rubrifolinm ragne r Diane cantonense (Lour.) Merr., V, Ch Liriope spicata Lour., V, Cr Neolourya pierrer Rodr., V, Cr Neolourya wabari Rodr., V, Ophropogon dracaenotdes Houk: fe Ophiopogon japonicus (L.f.) Ke eon Ophiopogon latifolins Rodr., V, C olins Dec a7 = ,V,Cr Peliosanthes pe oyana Pus ex Rodr., V, H Peliosanthes macrophylla ex Bak., V,Cr Peliosanthes serrulata Rodt., Peliosanthes teta Andt., or Polygonatum ean Gagne V,Cr l H aw) Tupistra tonkinensis Baill., SL COSTACEAE Costus spectosus (Koenig) Sm., V, Cr Costus tonkinensis Gagnep., V, Cr CYPERACEAE Carex brunnea Thunb., SL, Th Carex eryptostachys Brongn., SUM, H Carex indica L., SUM, H Carex leucociflora Bunge, SL, Cr SUM, H Carex trongit Khoi, SL, H Carex maubertiana Boot., Cyperus compressius L., V, Cyperus cuspidatus K. B. K., V, Th Cyperus difformis L., V, Th Cyperus esi Vahl var. diffusus, V, Hvar. latifolius L Cyperus ditidas Roxb., V,Th Cyperus distans L.f., V, Th Cyperus haspan L., V,H Cyperus imbricatns Retz., ahl) Cyperus iria L., Cyperus nutans var. elewsinoides (Kunth) aines Cc giiecale ieRE (L.) Vahl, V, H ,Cr V, Th (C. radiatus ..V, Th Eleocharis atropurpurea Shei) Presl, V, Th eleoch Presl) Eleocharia geniculata (L.) Rocka’ & Schule., V, 755 Th (H. carihaca (Rotth.) Blatke, H. capitata r. F ere complanata (Retz.) Fimbr ee dichotoma (..) Vah diphylla Vahl vat. ni V,Th var. Mitte Clar aca miliacea (L. on V,Th Fimbristylis monostachya (L.) Hassk., NL, Th Fimbristylis pauciflora R.Br T Fimbristylis quinquangularis (Vahl) Kunth, NL, Th ee : Th l, Fimbristylis squarrosa Vahl, NL, Th Fimbristylis thomsonti Bocklr., NL, a Fimbristylis umbellata Roctb. NL is (L.) Roxb., NL, Fuirena ciliat os vibiniate am Helen dats congesta D. Don, V, Th Hypolytrum nemorum (aki!) Spreng., WV, Cr (7. latifolium L. C. Rich.) Kylling oi vtfolia ee ,NL,H K. monocephala Rottb., Mapanta macrocephala ae m. & Warb., Mariscus compastus (Retz.) Druce, V, Th (Sphacromariscus microcephalus E.G. 2 amus) Mariscus umbellat tus Vahl, V, Cr , NL, Cr Pycreis Rhynchospora ibee teu ur i (Rhynchospora wallichtana Kunth) yxb., NL, Cr r igi: NL, Th Seleria juncoides R Silap a mucronatus . NL, Cr Scleria radula Hance, NL, Cr Scleria supinus L., NL Scleria tesselata Willd. ni Cr sp ac Nai ae Dioscorea alata Dioscorea eis »xb., - Droscorea wel ae oo ,¥v, Dioscorea nummutaria a. v, a D. pyrifolta Kunth) Dr0scorea pent aphy lla L . V, Cr Dioscorea per ais Prain & Burk., SL, Cr DRACAE Dracaena angustifolta Roxb., V, M racaena cambodiana Pierre & Gagnep., Mi SUM, ERIOCAULACEAE Eviocanlon mntermedium Koern., NL, Cr HYPOXIDACE AE Curculie ) Ge 1 p., V, Cr é Spades ne Wall., V, Cr Hypoxis sp., V, © 756 IRIDACEAE Belamcanda chinensis (1..) JUNCACEAE ee prismatocarpus R. Br., DC., NL, Cr NL, Th MARA CEAE Phryninm ee Willd., V, Cr Phrynium dispermum Gagnep., V, Cr Phrynium thorelti Gagnep. or Stachyphryninm sp., V, Cr MUSACEAEI Musa ttinarans Chessman, V, Musa uranoscopos Lour. (M.cocctnea Andr.), V,H ORCHIDACE Acampe rigida (Smith) P. P. Hunt, SL, Ep Aerides odoratum Lour., § artordes mn V,H | — ~ Anaphora lip Appendicula cornuta Blume, V, Ep ese chinensis Blume, NL, H ndina graminifolia YD. Don, NL, H L, - babs Mum concinum Hook. t., SL, Ep Sete a i scens ie ae SL, H pl h, SUM, Ep rit she)! S1LMESE — ex Donie a. ryllum nigrescens e, eon stenobullon Par, & Rei ichb.f., V, Bulbophyllum taentophyllam Par. & oe ,H Bullpen xylophyllam Par. & Reichb.t., SL, cote iia Lindl., V, H alia Linc ? , V,Cr pee heaeen Lindl., Calanthe triplicata vis m. veratvifolta R.Br Cleisostema aspersum eee f.) Garay, V, Ep Cleisostema striatum (Reichb.f.) Garay, V, Ep Ep en Or ) Ames, V, Cr (C. dk. V, Ep Coelegyne brachyptera Reichb.f., SL, Ep parishi dk.) atrifolia (Reinw.) Blume, V, Ch E LE Dendr en acinaciforme ee ,V,Ep d SUM, Ep I Dendy biun CVHMenAtH) Sw. 7 Dendrobium dentatum Godaue. (D.tene/lum auct.non Lindl, oe lindleyi Steud. (D. ageregatum Roxb.), V, Ep Dendr pe ae LindL., Ep Dendrobium hercoglobum oo {. Guill.), V, Ep Dendrobinm moschation Sw., SL, Ep SL, Ep (D. poilanei Dendrobium nobile Lind|., Stipa 17(4) ea ‘obinm Aid ace ke Ep ook. f., V, Ep endrobinm pod 101K WL. e var. delaccurii Dendrobium superbum i. ichb ragnep. & Guill., V, Ep Jendropogon aducum Hook.t., V,H Dendropogon wardianum Warner, V, H Desmotrichum poilanei Gagnep., SUM, Ep ‘ria arnert Reichb.f., SL, Ep Pp Re 1c chee Sp Evia coronaria (Lindl.) ., SL, Ep Evia globultfera Seident., Eria petelotti Gagnep. bk Erithrodes blumei find: ) Sei let V,Ep Evithrodes chinensts ae Schler., SUM, Ep Galeola javanica Benth., V," Gatleola ee Lour. Eis Reichb.f.), NL, Gastr ] obliqnus (Lindl.) Kuntze, V, Ep Goodyera fumata Thw., V,H Goodyera hispida Lindl., V, H Goodyera procora (Ker-Gawl.) Hook., V, H Goodyera soliosa (Lind|.) Benth. ex Clark, SUM, H Haemaria sp., V, H Hetaria rubens (Lindl.) Benth. ex Hook.f., V, Cr sen deliciosum (Reichb.f.) Sweet., V, Ep K. decumbens (Griff.) Rolfe) ie cordigera Hook.f. var. argentopunctata Aver.) Aver., V, Ep Liparis distans Clark., V, Ep Liparis latilabris Rolte, V, Ep Liparis manit Reichb.t., ip Liparis nervosa (Yhunb.) L idl, Ludista discolor (Ker-Gawl.) ee f., V, Cr (Haemarta discolor (Ker-Gawl.) Lind], Liisia morsei Rolfe, V, Ep Matlasis acuminata D. Don, V, Ep Malleola insectifera (J.J. Sm.) J.J. Sm. & Schlecht., V Neogyna sp., V, Ep Newiiiedio tinea his (Baker) Rolfe, V, Cr (N batansae Baill.) Oberonta ensiformis (J.E. Sm.) a Oberonta longthracteata Lindl., Oberonta myosunus (Foster) Lin, . E Oberonta pachyphylla King & Pantl., V, Ep Paphiopedilum appletontanum Rolfe, SUM, Cr Paphtopedilum delenatii Guill., SUM, Cr elatatheria insectifera (Reichb.f.) Ridl., SL Phajus tankervillece (Banks ex LHer.) Blum, NL, Cr Phalaenopsis gibbosa Sweet Phalaenopsis lobbit eieu: f.) — V,Ep P hala MOPSIS parishii a Sp SL, Ep ,V,Ep Phreatia secunda (Blume) Y find. Tun, Cucphuong National Park, Vietnam Physurus sp. Pogonia io mis Lindl., Renanthera coccinea Lour., SL, Robiquetia suceisa (Lindl.) Seidenf., V Sarcanthus kre ae Guill., V,E Sarcanthus Sp. p Shiranthes sinensis (Pers) Ames (S. L,C NL, Cr Ep australts Lindl.), N Sunipia pa sonti (Sing & Pantl.) P.F. Hunc., SUM, Ep Cone al ah Braid) bri idl.) Sch leccht., V, Ep Th rapermion es Lour., oo Ep Trichotosia curculigoides Lindl., SL, Ep Trichotosia pulvinata (Lindl.) Kraezl., V, Ep Vanda wile Blume, V, E Vanda doritoides Guill., SL, Ep Vanda teres Lindl., § E Vanda viminea Guill., : Vanilla annamica Ciscee. UM, Ep Vrydagzynea albida (Blume) ee >, V,Ep Zeuxine nervosa (Wall.ex Lindl.) ae ex Clark, SL, Ep Zeuxine vietnamica Aver., SL, Ep PANDANACEAE Pandanus odoratissimus L.£. (P. loureiri Gaudich.), , Mi Pandanus tonkinensis Martelli, VW, Na PHORMIACE Dianella ae DC., NL, H POACEAE (GRAMINEAE) Acroceras munroanum (Bal.) Henr., V, H (A. ,N Arthranxon ciliaris P. Beauv., NL, H (A. pga (Thunb.) Makino Arundinella bengatensis Spreng.) Druce, V, H Arundinella nepalensis Trin. (A. hniee Pan N Buiriahloa gaia aa i peta ae = (Ro; Se tap of) Copiinediun cinctum (Steud.) A. Camus, NL, q pias a - ) Desv., V, H jpogon a s (Retz. ) a NL, H oix Sie ae i a var, ma-yuen (Romanet) VT Stapf, Cynodon a tylon Per L, alin pi see ce V,H bilipes (N pies paten mee A. ee Vv, Dace Sides aegy as = ) Willd., ae Th Es egyptiacum Den Ji oe be ns V, PhH 757 Dendrocalamus longifibriatus Gamble, V, PhH Dendrocalamus membranaceus Munro, V, PhH Digitaria timorensis Domes Bal., NL, Th (D. 14) ichinochloa crusgalli (L.) P. Beaiv, NL, Th Echinochloa frumentacea (Roxb.) Link, NL, Th (E. erusgalli (L.) P. Beauv. var. frumentaceum Trin.) Eleusine indica (L.) Gaertn., V, Th Eragrostis geniculata Nees, N Eragrostis interrupta P. Beauv, NL, 1 Eragrostis nigra Nees ex Steud., NL, C Eragrostis pilosa (L.) P. Beauv., NL, Th var. pilosa var. imberbis Franch. a grostls ene Hochst. ex Miq., NL, Retz.) Nees ex Steud., NL, Eragrostis zeylanica Nees & Mey., NL, H (E. elongata auct. non Jacq.) Eulalia phaeotrix (Hack.) Kuntze, var. phaeotrix, JC r Psabelochen granularis (L.) Kuntze, NL. Th (Manisuris granularis (L.) Lf.) calles bis eae ‘a f.) R. Br., ,H a Bal., NL, H (Sacaien aise (Bal.) ak Chase) Imperata cylindnica (L.) P. Beauv., NL, a Lsachne globosa (Thunb.) ey N JH UO. ee — Isachne miliacea Roth, auct. non Doll. baemum Vr hari C.E.C. Fischer, NL, —~ avy ian auct. non L.) lace rugosum Salish. var. segetum (Trin) ack., NL, as hexandra Sw., sie Th Leptochloa panicea (Retz.) Ohwi., V, Th (Leptochloa filiformis auct. non P. - Luv.) Lapaiaen gracile Brongn., NL Microsteetum ctliatum (Trin.) A. C amus, V, Th Microstegium vagans (Nees) Merr., NL, Th Miscanthus flovidilus (Labill.) ee NL Schum. & Lauterb. See ao ) Narenga porphyrocoma (Hance ex Trimen) Bor., NL, H (Saccharum narenga Wall.) Peas dulloa A. Camus, NL, P Neyrandia arundinacea C ) ee var. me se (Buse) Herr., NL, I Kunth var. zo//inger? A. Camus) Oplismenus compositus (L.) P. Beauv. var. compositus, NL, H rele compositus ( (L.) P. Beauv. var. owtant donda) Nguyen, NL, 7358 NL, Ch (P. Panicum bisuleatum Thunb., acroanthum & Paspalum conjugatum Berg., NL, Th Paspalum longifolinm Roxb., ~ -aspalum orbiculare Forst., Paspalum scrobiulatum L., NL, H Paspalum — (L.) Spreng., NL, H ( compressum R. Br. Pennisetum purpurenm K. Schum., V,'Th H (P. valgaris Phragmites communis Trin., V, conatl init Peas ) Kunth, NL, H Polytoca digitata (L.f.) Druce), V, Th (P. heteroclita (Roxb.) Kor ) Pseundochinol ge pol Th Rotthoellia exaltata L.t., V,Th Saccharum Ree es ,V,H H.B.K.) Stapf, NL, Stipa 17(4) Urochloa paspaloides J. Presl ex Presl, NL, Th (Brachiaria ambicua (Vrin.) A. Camus), Urochloa reptans (1...) Stapt, NL, ‘Th PONTEDERIACEAE Ewhornia crassipes (Maret.) Solms, NL, H Monochoria vaginalis (Burm.f.) Presl, NL, Cr SMILACACEAE Smilax bauwhinivides Kunth, V, € Smilax bracteata Presl, V, Cr S. eee ila A. ray) a. lancaefolia Roxb., V, Cr Smilax ovatlifolia Roxb., V, Cr (S. Roxb.) macrophylla Smilax perfoliata Lour., V, Cr Smilax riparia A. DC., V, Cr STEMONACEAE Stemona saxorum Gagnep., Le Cr Stemona tuberosa Lour., V, Cr TACCACEAE lacca plantagine Drenth, V, Th (Schrzocapsa Haman a (Hance) Schizachyrium breaiiane (Sw.) Nees ex Buse, facca mtegrifolia Ker-€ » Cr (LF. laevis NI. Th Roxb.) Selerat ies punctata R. Br., NL, Th Tacca chantrievi Andr., V, Cr (E paxiana Limpr.) Setaria glauca ae i ee auv., V, Th (S. /vtescens TRILLIACEAE (Weig.) P. T. Hubb. Parts chinensts Franch., V, Cr rita pul eae ) Stapf, I Paris hainanensis Merr.. V. Cr tavlad (Schum.) Stapf& CLE. Hubb., | cee ; NL. H ZINGIBERACEAE ex M. B. Mess (8. avrea Hochst. ex A. Braun) Alpinta nutans Rosc., V, Cr q Setarta pers (L.) P. Beauv., V, Th Alpinia chinensis (Retz) Rosc., V, Cr Sinocalamus latiflorus (Munro) Mclure, NL, Alpinia tonkinensis Gi Be HEP ss ‘3 hH | so Munro) Anonum biftorum Jack, V, Ce mann meal wae: iis Amomum ovoidenm Pierre ex Gagnep., V, Cr H ( rath an ) "Stapf var. se eh Amomum thyrsoidenm Gagnep., V, Cr ans Amonum vespertilio Gagnep., V, Cr Songun a aaa yee Hitche., NL, H Curcuma elata Roxb., V, fine (J. Presl) A. Camus) Curcuma stenochila Gi agnep., ViCr on hol us a Cli) . Br., NL. H Kaempferta rotunda L. v le 7 : Stonotaphrum holfery Manro, NL, H Zingiber casswmunar Roxb, »V,Cr 2 i ( nep J 4 Themeda villosa ( (Poi) Dunes jacks NL, Ch Zingiber mon te “Tt V,Cr Thysanolena maxima (Roxb.) Kuntze, NL, H Zingiber ruber Roxb., Cr Zingiber zerumbet (L.) J. ss Sm., V, Cr REFERENCES ANoNyMous. 1971-1974. Flora Hainanica, I-IV Beving. ANONYMOUS i 980. Flora Yunnanica. I-V Beijing. ANONYMOUS 71-1975. Iconographia cormophytorum sinicorum. I-V. Sci. Publ. House. Stay phy Beying Anonymous. 1971. List of Cucphuong plants. Hanoi. AUBREVILLE, A., E. Vipat (eds.). 1960-1994. Flore du Cambodge, du Laos 7 TARDIEU-BLOT, et du Cicada. Paris. Vols. 1— THIN, Cucphuong National Park, Vietnam 759 Cuan, L.T., PHAN Ke Loc, NGuyen Nouia Tuin, and NonG Van Tigp. 1995. Contribution of research for some basic characters of Lamson flora. Collected scientific works on geog- aphy. Sci.-Tech. Publ. House, Hanoi. Pp. 285-297. Cotani, M. 1920. Etude sur les flores Tertiaires de quelques gisements de lignite de VIndochine et du Yunnan. Hanoi-Haiphong. Danu, T. 1985. Fossil of Dipterocarpaceae. Workshop on Dipterocarpaceae in Ho Chi Minh City. Vietnam 1986. Neogenflora in North Vietnam and its significance. Abstract of do & toral thesis. Do, Tar Lot. _ Sci. Techn. Publ. House, Hano}. Lecomte, H. (ed.). 1907-1951. Flore Gednedrale de |’Indo-chine. I-VII Paris. ACKNOWLEDGMENTS Karen Williams prepared the illustration of E. calendulina. Florida Agri- cultural Experiment Station Journal Series No. R- 05882. REFERENCES Mrrrow, A.W. 1987. A monograph of Excrosia (Amaryllidaceae). Syst. Bot. 12:460—492. NOMENCLATURAL NOTES ON NEOTROPICAL CLUSIEAE (CLUSIACEAE) JOHN J. PIPOLY U1 nical Research Institute o 4 Botanical Research Institute of Texas 509 Pecan Street Fort Worth, TX 76102-4060, U.S.A. fpipoly @ brit.org ABSTRACT In order to validate names for various checklists and florulas now in press, new combi- nations for some species previously placed in Oedematopus, Havetia, and Pilosperma are made. The new names Clusia engleriana Pipoly, C. hylaeae Pipoly, and C. colombiana Pipoly are provided for Oedematopus congestiflorus, O. weberbauert, and Havetra laurifolia respectively. Clusia polyandra, Clusia sect. Pilosperma, and Clusia sect. Havetia are new Spa based on Oedematopus polyandrus, and the genera Pilosperma, and Havetia respective RESUMEN Para validar los nombres de varios cataélogos y florulas ahora en prensa, aes oe ies as ubicadas en los géneros Oedematopus, Havetia, y Prlosperma se tran género Clusia. Se proponen los nombres nuevos Clusia engleriana Pipoly, C. hy ee Pipoly, yc. ee Pipoly para los binémenes Qede seas congestiflorus, O. weberbauert y Havetia laurifolia respectivamente. Clusia polyandra, Clusia sect. Pilosperma, y Clusia sect. Havetia son nuevas combinaciones, basadas en Ocedematopus ae ws, y los géneros Pilosperma y Havetia. INTRODUCTION In preparing treatments of the genus Clusia L. for the Flora of the Venezu- clan Guayana, Catalogue of the Vascular Plants of Ecuador, and Flora de Colom- hia, it has become necessary to relegate the genera Havetia, Havetiopsis, Oedematopus, Quapoya, and Pilosperma in synonymy under Clusia, in agree- ment with P. Stevens’ as yet unpublished treatment of the family for the Families and Genera of Vascular Plants. A key to all the sections of Clusia will be presented in the treatment of the species now placed in Oedematopus (Pipoly & Boom, in prep.) along with the descriptions of several new ones, and the formal transfers of the genera Havetiopsis, Oedematopus, and Quapoya to sections under Clusia are in press in the Flora of the Venezuelan Guayana (Berry et al., in press). Additional new names for some of the taxa in Oedematopus, the genus Havetia and the genus Pilosperma are necessary for areas outside those treatments. Sipa 17(4): 765-767. 1997 766 Sipa 1 7(4) NEW NAMES AND COMBINATIONS Clusia engleriana Pipoly, nom. nov. Oedematopns congestiflorus Engl., Bot. ie Syst. 58 (4/Beibl. 130):6. 1923 1 Clusia congestiflora Cuatrecasas, Rev. Acad. Colomb. Ci. Exacc. 8 (29):52, con tee PERU: Sandia, 2,300—2,400 m, 28 Mar 1902 (A), A. Weberbaner 624 (HOLOTYPE: B-destroyed; isoType: F Ww Distribution. Clusia oe is known only from the eastern slopes of the Andes in Peru, 2,300—2,400 m. Etymology. —Clusta se vana is named for Adolf Engler, one of the greatest contributors to our knowledge of the family, author of the treatment of the family for Martius’ Flora Brasiliensiy. C = hylaeae - s0ly, nom. nov. Oedematopus weberbanert Engl. iy , Bot. Jahrb. au (4/ a 0):6. 1923, non Clusia weberbauert Engl., Bot. ane Syst. 58 (4, an { ion. Type: PERU. Lorero: near Moyobamba, 800—900 m, 14 Rue ona a . : Weberbaner 4526 (noLoryrr: B- destroyed; isorype: F). Distribution.—Clusia hylaeae occurs at the lowland wet and premontane forest interface in the Amazon of Peru and Ecuador to approximately | ,000 m elevation. Etymology.—The specific epithet refers to the species’ habitat, in forests at the Hylaea/Andean interface. Clusia polyandra (Vesque) ee comb. nov. Oedematopus polyandrus Vesque, fonogr. Phan. 8:164. 1893, 7 > COLOMBIA. Antrioguia: between Caldas and Amaga, 2,000 m, without date, 7 ion 3220 (HOLOTYPE: P: IsoTyPE: F), Distribution. —Clusia polyandra occurs in premontane wet forests of the colombian Choco Floristic Province, 1,600—2,200 m. Clusia sect. Pilosperma (Planch. & Pee Pipoly, soa et stat. nov. Pilosperma Planch. & Triana, Ann. Sci. Nat. Bot. Ser. 4, 13:315. 1860. Typr SPECIES (by monotypy): Clusia candata (P anch. & ney Pipoly. An updated circumscription of the secion will be presented in the up- coming treatment of the species formerly placed in Ocedematopus (Pipoly & Boom, in prep.). Clusia caudata (P] ae .& Triana) Pipoly, comb. nov. Pilosperma caudatum Planch. Triana, Ann. Sci. Nat. Bot. Ser. 4, 14:244. 1860. Type: COLOMBIA. Cuoco: Pacific coast, i 11857 (fl), J. Triana s.n. (OLOTYPE: P, n.v. 5 lsorypPe: P), Distribution and ecology.—Clusia caudata is endemic to the Chocé Floris- tic Province, occurring from the Panamanian/Colombian border to the Colombian/Ecuadorian border, from 1,100—1,700 m. It is a common epi- phytic shrub. Clusia sect. Havetia (Kunth) Pipoly, comb. nov. Haveria Kunth in H.B.K., Nov. Gen. Sp. 5:203. 1819. Type Sprcirs (by monotypy): Clusia colombiana Pipoly. Pipoty, Neotropical Clusieae 767 Clusia colombiana Pipoly, nom. nov. Havetia laurifolia Kunth in H.B.K., Nov. Gen. Sp. 5:204, 1819, non Clusia pete Planch. & Triana, oy Sci. Nat. Bot. 4, 13:344. 1860. Type: COLOMBIA [Nueva Granada]. uca: Andes de Popayan, entre ie Vega de os Lorenzo y Pansitara, 2,530 m, ere date (fl), A. Humboldt & A. Bonpland s.n. (P-BON, n.v.). Clusia colombiana is a common shrub in the subpéramo formations of Colombia and adjacent northern Ecuador, from 1 ,600—2,200 m elevation DOUBTFUL OR EXCLUDED NAMES oe tr: martii Spruce ex Planch. & Triana, Ann. 3 — Bot. Ser. 4, -246. 1860. Havetia laurifolia Mart. Nov. Gen. Sp. Pl. . 166, 297. 1829, non H.B.K. Havetiopsis laurifolia (Mart.) Engl., Fl. poe. 13:438. 1858. Type: BRA- ZIL. AMazonas: Rio Negro, without date, C. Martius 5.2. (HOLOTYPE: M, n.v.). I have not had an opportunity to examine the type material, housed in the Martius Herbarium (M). From the descriptions available, it may be a synonym of what has traditionally been recognized as Havetiopsis flexilis Spruce ex Planch. & Triana. ACKNOWLEDGMENTS I chank the curators of the cited herbaria who made the specimens avail- able to me. Barney Lipscomb, Charlotte Taylor and Jon Ricketson reviewed the manuscript. REFERENCES Berry, P., B.K. Horst, and K. Yarskievycu (eds.) . Flora of the Venezuelan Guayana. Mis- souri Botanical Garden (In Press). COMMENTARIES ON ASA GRAY’S BOTANICAL TEXTBOOKS BY GRAY AND HIS CONTEMPORARIES (1836-1887) RONALD L. STUCKEY Herbarium, Museum of Biological Diversity Department of Plant Biology The Ohio State University 1315 Kinnear Road Columbus, OH 43212-1192, U.S.A. ABSTRACT Asa Gray (1810-1888) authored 11 different editions of his textbooks of botany during the second half of the nineteenth century. His books were considered by his contemporar- ies as models, the best ones written from the views of current botanical knowledge and for covering the entire scope of the subject as known in his day. This paper describes Gray’s textbooks by using information from Gray himself as taken from (1) the books’ prefaces and his correspondence with botanical friends, and from (2) commentaries from other writers in published reviews of his textbooks. Based on a titles, contents, and ae Gray’s textbooks may be placed into three categories: those that (1) discuss the basic ments or provide the essential elements, written for beginning students in schools, “ lished in 1836, 1857, and 1887; (2) have the word “textbook” in the title, writcen for students at the collegiate and secondary levels, published in six rewritten editions, 1842, 1845, 1850, 1853, 1858, and 1879; and (3) have a morphological- Solana orienta- tion written for use in elementary schools, published in 1858 and 1872. Gray's textbooks jos were original concributions that organized botany into a more a science and fulfilled the educational needs of the time. They were in demand for use in schools and the public in general, but after Gray's death, no one continued preparing revisions, and thus both their use and usefulness ceased. RESUMEN Asa Gray (1810-1888) fue el autor de 11 ediciones diferentes de libros de texto de botanica durante la segunda mitad del siglo diecinueve. Fueron considerados por sus contempordneos como modelos de libros bien escritos — os panes. de vista del conocimiento botanico de su tiempo y de cubrir el espectr ida en la época. Este articulo describe los libros de Gay mediante iomaciones aero del mismo Gray tomadas de (1) los prélogos de los libros y de su correspondencia con amigos botanicos, y (2) de comentarios de otros autores en revisiones publicadas de sus libros. B Basindonos en sus s titulos, contenidos e neeneionalidad: los libros de mek: pueden ss ] agruparse en es (1) losaqu liscuren elementos ba SICOS u sfrece py bdsicos, escritos para Ssccuclagies principiantes en las escuelas, poblicados en 1836, 1857, 1887; (2) los que llevan la palabra “textbook” (libro de texto) en el titulo, escritos para scant de Ses nivel y universitarios, publicados en seis ne revisadas, 1842, 850, 1853, 1858, y 1879; y (3) los que tienen una orientacién morfoldgica- fisiol6gica escritos para ser nsados en escuelas elementales, publicados en 1858 y 1872. Los libros de Sipa 17(4): 769-802. 1997 —~ TiO Stipa 17(4) Gray fueron contribuciones onetnalcs ane: estructuraron la botainica en una ciencia mas util y sacisficieron las necesidad le la €poca. Tuvieron demanda para ser usados en escuelas y por el niiblico en pai pero weer de la muerte de Gray, nadie continud haciendo las revisiones, y por ello cesaron tanto su uso como utilidad, Asa Gray (1810-1888), considered the foremost nineteenth century bota- nist in the Ae States, authored 11 different editions of his textbooks of botany, regarded as among the best at that time in the United States. Pub- lished during the second half of the nineteenth century, Gray’s books were considered by his contemporaries as models, because they covered the views of current botanical knowledge and the entire scope of the subject as known in his day. He wrote them at a level of comprehension and complexity different from most textbooks, in order that they would be available and useful to all ages and knowledge-levels of individuals wanting to have in- formation about plants. Despite their widely recognized merit, no discus- sion, so far as is known, has ever been prepared on Gray’s botanical text- books. When one sees the various titles and dates of publication, either from a collection of the books in the library or in lists of textbooks, the complexity and confusion of this subject become apparent. Gray's biogra- pher, A. Hunter Dupree (1959) offers very little information about Gray’s textbooks, but his book has provided the chronological backdrop for the organization of this paper’s text. Because of the merit of Gray's textbooks, they are worthy of being identified and described. The procedure in this discussion follows the chro- nological order in which they were written and published.! Information about them is taken from two primary sources: (1) Gray himself, as the writer of the prefaces and correspondence with his botanical friends, and (2) commentaries in published reviews of his textbooks. Both of these sources contribute significantly and realistically coward an understanding of the content and quality of the books. The comments quoted from the reviews are the reviewers’ opinions of Gray’s books. Information on the content and quoted passages from the books, customarily used in the writing of book | The inspiration for writing this paper on Asa Gray's botanical textbooks came while reviewing abstracts and manuscripts of papers that Emanuel Rudolph (1927-1992) presented in 1971 and 1975 at two history of science meetings. This effort was part of the ae project of editing Rudolph’s unpublished papers for their publication as a memorial vol- ume (Stuckey and Burk, eds. In Press). A short paper of Rudolph’s research on Gray’s text- books (Paper No. 3) appears in that volume. This paper is in part written from notes and quoted portions of Gray's letters published by Jane Loring Gray (1893) that Rudolph had collected, with the remainder based on research by Stuckey with generous bibliographic and word processing assistance from William R. Bile Biology Librarian at the University of North Carolina, Chapel Hill. The first draft of this paper was written in December 1995 and revised with additions in December 1996, while at the U niversity of North Carolina, Chapel Hill. Stuckey, Asa Gray’s botanical textbooks 771 notices and reviews during the nineteenth century, are excluded from these commentaries. The reviews examined in this study were taken from the reference list compiled by Stuckey and Burk (1998, in press). Based on their titles, contents, and purposes, Gray’s textbooks may be placed into three categories: those that (1) discuss the basic elements or provide the essential lessons, written for beginning students in schools, published in 1836, 1857, and 1887; (2) are identified with the word “text- book” in the title, written for individuals in colleges and secondary schools, or for private students, published in six rewritten editions, 1842, 1845, 1850, 1853, 1858, 1879; (3) are the books which have a morphological- physiological orientation written for young people in elementary schools, published in 1858 and 1872. Gray’s textbooks are described and discussed in chronological order of their writing and publication, rather chan by the grouped topic. A list of their full citations at the time of their first publica- tion is arranged in chronological order (Appendix I). Known reprint edi- tions based on citations published in The National Union Catalog, Pre-1956 Imprints (Compilers 1972) are noted at the end of each citation of the corresponding original edition. Selected features of Gray’s books not dis- cussed in the text appear as an added summary (Table 1). A true textbook is, according to the American Textbook Publishers In- stitute (1949, p. 19), a published work “especially prepared for che use of pupil and teacher in a school or class, presenting a course of study in a single subject, or closely related subjects.” Textbooks, particularly those o wide use, according to Rudolph (1971, a, b), are by their nature reposito- ries of generally accepted knowledge in any discipline at a particular pe- a) tiod. The botanical textbook is here defined as a work that introduces stu- dents to current aspects of plane science, rather than a manual for the identification of particular plants. Textbooks are useful for the historian of science because they present in a concise way the generally held paradigms” of a period. Rudolph (1975) further noted: “They {textbooks} should pro- vide an insight into the ‘normal science’ of the time. For textbooks are windows to the posture that the science takes when being presented to the initiate.” The concept of “normal science” comes from Kuhn (1970), who defined it for the history of science.” The origins of textbooks are to be 2 “ . ” Ape Paradigms,” according to Kuhn (1970, p. viii) are “universally recognized scientific achievements ak for a time provide model problems and solutions to a community of practitioners.’ > “Normal science,” according to Kuhn (1970, p. 10) is “research firmly based upon one or more past scientific achievements, achievements that some particular scientific commu- nity aon ee for a time as supplying the foundation for its further practice. Today such achievements are recounted, though seldom in their original form, by science text- Taste 1. Summary of Selected Features of Asa Gray’s Botanical Textbooks Abbreviated Title of Book Year on Title Page Year of Copyright Copyright Date of Preface Number of Illustrations Illustrator Number of Additional Printings Changes from Previous Edition Group 1. Elements and Lessons: Elements 1836 First Lessons 1857 Elements Group 2. “True” Textbooks: Textbook ed. 1. 4 Textbook ed. 2. 1845 Texthook ed. 3. L850 Texthook ed Mie l 853 Texthook ed. 5. 1858 Textbook ed. 6. > 1842 1842 1850 Group 3. “Botany tor Young People:” J § How Plants Grou How Plants Behave 1872 1900 Asa Gray Wiley & Putnam Wiley & Putnam George P. Putnam George P. Asa Gray Ivison & Jane L. Gray April 1836 1 January 1857 March 1887 No Preface March 1845 April 1850 March 1853 September 1857 10 April 1879 No Pretace 2 February 1872 Isaac Sprague Isaac Sprague Miss Agnes Mitchell; illustrations by Isaac Sprague same as edition 3 Isaac Sprague Isaac Sprague Isaac Sprague none none none none none “new and much sed portions “entirely written’ “rewritten and arge “corrections and minor alterations” “almost entirely rewritten “entirely rewritten” ‘Figures are numbered by two methods: “Each figure contains one or more individual illustrations, each designated with the | etters a, b, c, etc. "Each individual illustration is given a figure number. Stuckey, Asa Gray's botanical textbooks PLS, found in works such as Linnaeus’ Philosophia Botanica (1751). The Linnaean model persisted throughout the eighteenth century and stressed gross struc- ture and history as a tool for classification. This model, which was used in books in England and on the European Continent, was gradually replaced by works having greater emphases on plant function and geography. Asa Gray's botanical textbooks were examples of these latter works. Of Gray's textbooks, Joseph C. Arthur (1895, p. 362) of Purdue Univer- sity in his vice-presidential address of 29 August 1895 to Section G, Botany, of the American Association for the Advancement of Science spoke as follows: Botany, as a substantial part of the curriculum, cannot be said to have received recognized standing in the American educational system until the time of Asa Gray. In the latter part of the decade of the thirties his first text-book, the “Elements of Botany, appeared, and in the decade following the ‘Text-book for Colleges’ and the ‘Manual,’ all of which works showed a true appreciation of the best features of the science and the needs of the time. They were so well conceived, and so much in demand, that new editions rapidly succeeded one another; and to the present oe they hold a high place in the estimation of botanical teachers. These works po sessed a specially potent element of virility in being the expression of know] ae at first hand, the words of the master. In so far as inspiration was drawn from foreign sources it came chiefly from French and English scholars, of whom De Candolle the eldest and Robert Brown were the representatives. WRITING THE ELEMENTS OF BOTANY (1836) By early 1835, the 24-year-old Asa Gray (Fig. 1), having graduated from medical school four years earlier, had concluded he did not want to practice medicine but become an authoritative botanist. Unemployed and living at home with his father in Sauquoit, New York, he “was requested to superin- tend the republication of some one of the most approved European trea- tises” (Gray 1836, p. x). As Gray began to work, he soon concluded that the European works were much too large, illustrated with expensive en- gravings, or contained a considerable quantity of information which did not belong in an elementary work. Gray decided “to engage in the more formidable task of preparing an original work, expressly adapted to the use of the student of North American Botany” (Gray 1836, p. x) He resolved to write an elementary textbook in botany. Gray needed money, but of equal importance was his desire to establish himself as a repucable botanist. A books, elementary and advanced. These textbooks expound the body of accepted theory, illustrate many or all of its successful ie and compare these applications with exe suplany observations and experimen science” and “paradigms” are scientific achievement that is yet still open-ended to allow for various related problems to be studied or resolved by future research scientists, to whom Kuhn (1970, p. 10) refers to as the “redefined group of practitioners.” similar in that both share in expounding the 74 Sipa 17(4) Fic. 1. Asa Gray (1810-1888), from a photograph in the Emanuel D. Rudolph collection, Archives, Herbarium, The Ohio State University, Columbus. 3 good textbook might initially earn $300, but the project also might be of greater value to botanical science, for he believed that a new and original textbook should be made available. The book would be his own, like an American Lindley or a French de Candolle, arranged on the Natural Sys- tem, rather than the antiquated, inferior Linnaean Sexual System as em- ployed in the books written by Amos Eaton, and his student, Mrs. Almira Hart Lincoln Phelps. In this creative endeavor, Gray hoped to challenge Stuckey, Asa Gray’s botanical textbooks 775 the European masters with a new standard of botanical excellence (Dupree 1959, p. 48). Despite differences, distractions, and doubts, Gray forged ahead and wrote the manuscript during the summer and fall of 1835 and on into the winter of 1836. The book was to have a balance of information on form, function, and classification which allowed students to understand the entire science of Botany. His main concepts came from Augustin P. de Candolle of Geneva, Switzerland, the master of all fields of botany. During the course of writing, Gray left home in the autumn of 1835 to reside in a boarding house in New York City, where he could be close to a scientific community and to publishers of textbooks. On 28 September he wrote his father: “As to my book, I am trying to make a bargain with two publishers; the prospects seem pretty fair, and I shall probably get $300, which is the sum I insist on. I shall have a definite answer in a few days” (J.L. Gray, 1:54. 1893). Some years later while reflecting on those days, Gray noted: “{I} arranged with Carvill & Company to take my book. I think they gave one hundred and fifty dollars, which was a great sum for me” (jb. tary; 1520... 13899); To assist in proofreading and adding much to the clear, crisp, business- like prose, Gray (1880) had his 12-years older English friend John Carey, a gentleman of considerable botanical knowledge, revise the proof pages and help bring the manuscript to completion through the press. Gray signed the preface in April 1836, and 10 or 12 days later he expected the printed book. On 3 May 1836, Gray notified his friend Nathan W. Folwell, as quoted in Dupree (1959, p. 55), saying, “This preparatory training ... has prepared me most thoroughly for future progress, and if I happen to pursue Botany undividedly for a liccle time I shall (entre nous) be soon the best botanist in this country;” a strong boast for a young man who was soon to be age 25. Completion of Gray’s first textbook came before his initial ap- pointment to a university teaching position, his nonresident professorship appointment in 1838 at The University of Michigan (Bartlett 1941). An original work, Gray's Elements was adapted to the student of North American Botany (Fig. 2). As Gray (1836, pp. x—xi) stated in the preface, its execution was not easy, chiefly because of the “difficulty of combining rigid perspicuity and philosophical accuracy with a popular form and a familiar style.” To achieve this goal, Gray further wrote in the preface that he freely made use of all the most approved works and original memoirs upon the subject that were within his reach, and has more particularly and constantly con- sulted the Théorie Elémentaire, Organographie Végétale, and Physiologie Végétale of De Candolle; the Traité d’Anatomie Végétales, and the more recent Elémens de Physiologie Vévétale et de Botanique of Mirbel; the Introduction to Botany, the Introduction to the Natural System of Botany, and the Ladies Botany of Prof. Lindley; the Article Botany Se - ee Fic. 2 Title page of Gray's Elements of Botany (1836 Sipa 17¢ ELEMENTS By ASA GRAY, M.D. MEMBER OF THE C¥SAR. LEOPOLD.-CAR. ACAD, NATUR.& CURIOSORUM, AND y¥ TUE LYCEUM OF NATURAL UWisTORY, NEW YORK. natura modos poimim dedit: his genus omne j sylvan, fruticunque viret f quod ipse vid sibi repperit usus. Vino. Geora. II, 20, NEW-YORK : G. & €. CARVILL & CO. 1836. ann ees ore wrens = eran sae ete sesso et PRR 4) Stuckey, Asa Gray’s botanical textbooks 777 in the seventh edition of the Encyclopaedia Britannica, by Mr. Arnott; and the fifth edition of the Nowveaux Elémens de Botanique of Achille Richard. The Elements’ (1836) preface outlined and described the contents, which basically contained three components: (1) Structural Botany, (2) Physiologi- cal Botany, and (3) Systematic Botany. The third component was further divided into (a) Taxonomy, the Principles of Classification, in which he fa- vored the Natural System over the Artificial System of Linnaeus, (b) Phytography, an exposition of the Rules of Botanical Nomenclature with the idea of directing students’ attention to this phase of botany whose 1m- portance had been undervalued and needlessly violated in this country, and (c) Glossology, the definition of the technical language used in describing plants, here confined to adjectival terms. An Appendix provided general directions for collecting and preserving plants for the herbarium, and con- tained a list of the plant families arranged in the Natural System. Two reviews of Gray’s Elements have been located. In the American Journal of Science (30:399. 1836), an anonymous writer, probably its editor Ben- jamin Silliman, wrote that Gray’s book was “the best work on the philoso- phy of Botany that has appeared in this country, and ... that its merits will be appreciated by the numerous students of this science among us.” The other review, signed “D,” is believed to be Gray’s friend, the physician William Darlington of West Chester, an authority on the flora of Chester County, Pennsylvania, who wrote in The American Gardener's Magazine (2:421-424. 1836): _.. the aim of the author seems to have been to exhibit a full view of the present state of the science, with all the recent improvements and discoveries, condensed into a clear and perspicuous treatise, which, being sufficiently popular in expres- sion, should at the same time retain that rigid accuracy so indispensably necessary to so extensive a branch of natural science. In this Dr. Gray has succeeded perfectly . {by the} excellent arrangement of the subject, and the lucid manner in which the whole is illustrated and explained. Those persons who are noc familiar with the large and expensive treatises published abroad, will find a great mass of new facts for study and digestion, and to those who are endeavoring to attain the elements of jon botany, we recommend this volume as a text book of the highest merit. In later years, an anonymously written review in the Bu/letin of the Torrey Botanical Club (6:317. 1879) referred to Gray’s Elements of Botany (1836) as the book that provided this writer's “first insight into the science. We well recall the delight with which we read the clear elucidation of the subject— delight in the method, for we had no particular experience in the matter.” Gray's British friend Joseph D. Hooker (Nature 37:376. 1888) noted: The “Elements” is a noteworthy book; it was at once accepted as the best that had appeared in the States, and as second to none in the English language; its only rival was {the American edition of} Lindley’s “troduction to the Natural System of Botany,” ... {which was used in} American schools. 778 Sipa 17(4) PREPARING THE BOTANICAL TEXT-BOOK (1842) AND ITS REVISION (1845) In 1841, Gray began to prepare a new textbook, even though he was in the midst of a pape John Torrey to write a Flora of North America (Torrey & Gray 183 844). Since botany was widely taught in the schools, he could at once aes some needed cash, and because of Amos Eaton's death in 1842, Gray would be able to reach students previously served by Eaton and his followers. Gray could also nourish a new group of botanists capable of using, but who only incidentally were purchasing, his Flora. During the course of writing his textbook, Gray was offered and accepted on 30 April 1842, the appointment as the Fisher Professor of Natural History in Harvard University. With an added incentive of writ- ing a good text for his forchcoming classes, and realizing the distinguished community he was about to join, he respectfully inscribed the book to the most eminent botanist of Boston, “Jacob Bigelow, M.D., E.L.S., Professor of Materia Medica in Harvard University; author of the Flora Bostoniensis, and of the American Medical Botany.” He completed writing the textbook in late July, and arrived in Cambridge in sufficient time to take on his new position in September 1842. At Harvard the demands of science and the rewards from society came into balance for Gray, and the Fisher Professor- ship marked a real beginning there of a continuous scientific study of botany, as botany now was most important in Gray’s life. Near the end of 1842, The Botanical Text-book for Colleges, Schools, and Private Students was pub- lished, having two parts, comprising “An Introduction to Structural and Physiological Botany,” and the “Principles of Systematic Botany.” On the title page was also the identification chat Gray was Fisher Professor of Natu- ral History in Harvard University. He noted that “it was in che course of printing when I was appointed to the Fisher professorship, so chat I could put that title on the title-page, and have a text-book for my class” (J).L. Gray 1:28. 1893). As was the situation earlier when writing the Elements of Botany (1836), Gray met with difficulties and delays in preparing The Botanical Text-book (Fig. 3). He wrote (J.L. Gray 1:282. 1893) co George Engelmann in St. Louis, 30 March 1842, saying: Owing to illness I have as yet written almost nothing, and besides have to superin- tend all the drawings, as they must be made by a person unacquainted with botany; one at the same time I have to correct the proofs of about thirteen sheets yet of the Flora,” so that r am almost distracted when I think how Iam to Pee it here, where I have to see personally to almost every detail. But I must do it, as I hope to lay the foundation for a popular and—what is of consequence to me—a profitable vork Stuckey, Asa Gray’s botanical textbooks Fic. 3 THE BOTANICAL TEXT-BOOK FOR COLLEGES, SCHOOLS, AND PRIVATE STUDENTS: COMPRISING PART I. AN ceases TO STRUCTURAL AND ANY. HYSIOLOGICAL BOT PART IL. THE PRINCIPLES OF SYSTEMATIC BOTANY WITH AN ACCOUNT OF THE CHIEF NATURAL FAMILIES OF THE VEGETABLE KINGDOM, AND NOTICES OF THE PRINCIPAL OFFICINAL OR OTHERWISE USEFUL PLANTS Ellustrated with wumerous Wigrabings ow woor. BY ASA GRAY, M. D. PISHER-PR HISTORY IVERSITY: 1 ature Curjosorum, of the B. of Ratisbon ; Honorary M A Academy of Arts nd Scien: f the B g ¢ | he. NEW-YORK WILEY AND PUTNAM. BOSTON; LITTLE AND BROWN 1842. Title page of Gray's The Botanical Text-book (1842). nce TE ta 779 780 Stipa 17(4) No preface was written for this book, but Gray provided some com- ments about the first edition (1842) in the preface of the second edition (1845). Both students and botanists using the book gave favorable opin- ions which satisfied the author, even through the book was prepared in great haste and carried rapidly through the press. Gray agreed that the text arrangement was satisfactory, but he desired to prepare an even better book. Atter the Text-book appeared, Gray enthusiastically wrote QJ.L. Gray 1:297. 1893) to John Torrey, 3 January 1843, commenting on a review from a British journal: The December number of [che] “Annals and Mz igazine of Natural History” {10:352- 354. 1842] (of which Professor Balfour is the botanical ed itor) Contains a very com- plimentary notice of the “Botanical Text-Book,” accompanied with a few judicious selections, which shows that the writer has looked it over carefully; and winds up by terming it the best elementary treatise (as to structural eee in the English language. So easy is it to get praise where it is not particularly deserved! The major points of the review were: It gives a comprehensive view of the present state of botanical science, and is writ- ten ina clear and lucid style, so as to render it accessible to all classes of readers. ... The work is illustrated with engravings on wood, which are highly useful to the student ... a good view ts given of the principle of classification, and the Artificial and Natural methods are well explained ... The nomenclature of botany receives a due share of attention, ... Upon the whole, we look upon this work as one of the best Text-Books which ... has yet appeared. — Additionally, on the foreign scene, Gray’s British friend William ay Hooker, in the London Journal of Botany (1:636. 1842) referred to Gray’s second book as . an immense mass of useful information, ... most perspicuously detailed, and rendered still more intelligible to the student, by the well executed figures, which are judiciously worked in with the type. We are much mistaken if this dofes} not become a popular book in Our country, A short review in The Gardener's Magazine (2, 3rd ser. 634. 1842) con- ducted by J.C. Loudon of London stated that Gray's Botanical-Texthook “ap- pears to us a very excellent work... . {with} numerous woodcuts, which are very well executed; and the paper, ene, and printing, are equal to those of any London publication.” The German botanist D.E.L. von Schlechtendal (Botanische Zeitung 1:50— S51. 1843) made several comments, as noted in translation: (1) the printing and paper were superb making a positive impression, (2) the woodcuts were very clean, clear, and distinct, (3) the representative plants, taken from the North American flora, made the book especially interesting to European botanists, and (4) although concisely conceived and only intended for the beginner, Gray's text book touched upon everything interesting and worth knowing. Stuckey, Asa Gray’s botanical textbooks 781 In the United states, two reviews are known: First, payin friend Wil- liam Darlington (American Journal of Science and Arts 43:388—389. 1842) wrote: The Text Book of Dr. Gray, affords at once the most compendious and satisfactory view of the vegetable kingdom which ras yet been offered, in an elementary treatise, to the American public. In a style remarkable for its correctness and perspicuity, the author has traced and unfolded the vegetable structure, from its simplest forms up to its most complicated and elaborate developments. He has presented us with the first principles of the science, in accordance with the ... {“admirable doctrine of vegetable metamorphosis”} ... by which the external organs of plants are gradually rodified, or ed ek from the crude cotyledons of the germinating seed, to the most delicate component parts of the flower and the fruit. [Presentation of this doctrine} has, indeed, given to the science of botany an entirely new aspect. Darlington also pointed out that with this textbook, “the teachers of botany ... may speedily elevate the study to its legitimate rank among the natural sciences... . [The student’s} faculties of observation and compari- son ... [will be} exercised according to the strictest rules of logic and phi- losophy.” As Darlington further elaborated, the student can learn the true characters of the plants, to judge their economic value, and derive pleasure from studying the science itself. These advantages “may be confidently expected from the general introduction and proper use of the Botanical Text Book, . Second, in hike North American Review (56:192—207. 1843), George B. Emerson discussed and analyzed in detail various topics about plants in Gray’s Textbook, including the movement of fluids in roots and stems, the theory of the transformation of leaves, the structure and symmetry doc- trine of leaf arrangement, food and nutrition, and essential elements. The reviewer questioned Gray’s introduction of the Natural System of plant classification in a basic textbook, when in reality the long-used Linnaean System of Classification had proven efficient and dependable for the begin- ning botanist in learning the organization of plants and their binomial names. — Reviewer Emerson, however, did recognize Gray’s efforts to the extent that botany has become more than what Linnaeus had provided to his audi- ence of that time period, stating that Gray’s _ book puts within the reach of thousands the enlarged views and vast and exact knowledge of the Jussieus, of Linnaeus, of Richard, DeCandolle, and Robert Brown, men who deserve to be known to common fame, as among the greatest observers [of plants] and most original thinkers of the last hundred years. Through his efforts of writing The Botanical Text-book (1842) Gray could serve science, and at the same time, also serve the general public. His sec- ond edition appeared in 1845, and Gray referred to this book in the preface (p. [9]}) as a 782 Sipa 17(4) - compendious treatise ... designed to furnish classes in our schools and colleges with a suitable text-book, as well as private students with a convenient introduc- tory manual, adapted to the present condition of botanical science Believing that systematic botany is best studied when grounded upon struc- tural and physiological botany, Gray wrote (1845) in the preface (p. 10) that these latter topics have been . amplified to nearly twice ... [cheir} former extent, [and], ... have been almost entirely rewritten; ... the principal topics ... as far as is practicable in a brief and strictly elementary treatise ..., instead of having been rendered more abstruse by the enlargement, will rather be found to be more simple and generally intelligible than before. The chapters upon the Principles of Classification, and of the Natural System, have also been recast and somewhat enlarged. An extensive, detailed account of this second edition appeared in the North American Review (61:254—258. 1845). The reviewer, George B. Emerson, stated that this new edition was necessary artly on the account of the rapid advancement which physiological botany . ae} 1S 2 neni making; and partly because Dr. Gray's experience as a teacher , Ais enabled him to discern ... the points which were defective, or which seamed more detailed explanation... . We are glad to see, therefore, that the author ... has made a good use of the pe cecaes and of his professional experience, and has recom- posed, and almost entirely rewritten, the first, and, in the author's view, the most essential, part of the work, chat upon structural and physiological botany, ... Among other changes were the doubling of the number of illustrative wood- cuts and changes in style to render a more flowing narrative. The book was noted in the same review as being . very neat and tasteful [in the} manner in which the mechanical part ... is finished. Its outside is as attractive as the subject of which it treats, the pindine:: eke paper, & and engravings being not merely unexceptionable, but elegant. This is such a rare merit 19 a mere manual of instruction ... ,a point of so much importance, that it deserves especial notice and commendation. Later, in the July issue of the North American Review (67:180—182. 1848), John Carey, botanist who aided Gray with writing and proofreading his Elements of Botany (1836) concluded concerning Gray’s 1845 edition, as well as Gray's previous elementary works, that they were entirely worthy of his well-established reputation. His subject is botany, but it is evident that he writes and feels as a general naturalist, and this is che pointe which we wish to have constantly kept in view. The details are minute and ample, and, for the most part, very concisely expressed; buc he never loses sight of the great gener- alizations which must ever accompany progressive science. We think Dr. Gray has fairly realized his expressed design, “to furnish classes in our schools and colleges with a suitable text-book, al (Botanische Zeitung 4:98-100. 1846), the German botanist, repeated that the examples from the North American flora serve on D.EL. von Schlechtenc Stuckey, Asa Gray's botanical textbooks 783 to recommend the work, but he was highly critical of the text on certain anatomical features, as indicated in translation: _ we find a substantial ae uaerne in that the lesson on cells and vessels, as well as vue sections that cover the main plant organs, are explained much more exactly with more consideration for al relationships and through more plentiful accompa- nying woodcuts. However, it seems to us that some sections are in need of elabora- tion, for example the one on cell contents, in which the green color should appear only in the places exposed to light and is derived from small grains. Moreover, no mention is made of either the starch nucleus of these grains or of the uniform green fluid; nor of any other colors, essential oils, or ocher substances that often fill entire cells. The trichome formations also are dealt with perfunctorily, and moreover no discussion is given about stellate trichomes, scales, or even of the waxy precipitates on the surface of the epidermis. ertainly this significantly reworked edition will also find a better reception than the first, which makes this second edition necessary after only three years and at the same time serves as a witness to the ever expanding study of botany even in these regions, which, invigorated and made more common by his handbook, re- dounds to the author's joy and honor. Printing and paper are excellent THIRD EDITION OF THE TEXTBOOK (1850); COMPETITOR WITH WOOD'S CLASS-BOOK (1845) About 1843 or 1844, Alphonso Wood, a teacher at an academy in Meriden, New Hampshire, who professed an interest in botany, came to talk with Gray on two occasions, proposing a joint-effort in writing a bo- tanical textbook. Wood wanted to have a textbook for secondary schools which would also serve as a manual based on the Natural System of classification for the plants of the region. Like Amos Eaton, he thought practical plant collecting and identification, rather chan structural-physi- ological botany was the proper approach for a beginning course, and hence Gray’s Botanical Text-book (1842) was the primary target for replacement (Lyon 1939, p. 18; Willis 1881, pp. 54-55). Wood's production was his Class-book of Botany (1845), and upon its appearance was destined to do more damage to the discipline of botany than threatening the income from Gray's text-book. Wood's book, accord- ing to Lyon (1939, p. 18), was fresh, complete, and original for its empha- sis upon field work. It included the fundamentals of form and function of plants and was notable for its descriptions of plants in common language that was simple and accurate. The book was appropriate for Wood's “edu- cational scheme of its time and aroused the interest of younger people in the plants,” (Lyon 1939, p. 18). In the preface of the second and sometimes later editions of his Class-Book, Wood (1847, p. 3) stated that his book treated the basic elements of botany according to the latest authorities, and was written in the form of simple propositions, briefly illustrated, with 784 Sipa 17(4) short paragraphs for the convenience of the learner. For students who aim toward a much higher level of botanical understanding, Wood wrote, “it affords us pleasure to be able to recommend ... the full and elaborate ‘Text Book’ of Dr. Asa Gray,—an American work of the highest meric.” Gray did not have a glowing opinion of Wood's Class-book (1845). He noticed that it gave only scant coverage to anatomical and physiological botany and fea- tured the flora of the northeastern portion of the United States, not as a mere list of families, as in Grays’ Text-book, but with a brief English de- scription of each species based on several sometimes unreliable sources. In Gray's mind, Wood's classification should not be allowed to become the standard guide for American plants. Gray immediately went to work to produce his own competing botanical Manval (1848) and followed soon after with a rewritten and enlarged third edition of his Botanical Text-hook (1850), only five years from the issuance of the second edition (Gray 1845). The beginning of the preface (p. [v]) in this third edition is the same as in the second edition (1845), with “the structural and physiological part ... again almost entirely rewritten ... much enlarged ... the symmetry and morphology of the flower {has} ... been altogether recast and greatly ex- tended.” Ina letter of 2 April 1850 to William J. Hooker (J.-L. Gray 1:368. 1893), Gray noted: The re ane of all the structural parts of my 3d edition of the “Botanical Text- Book,” which I was inadvertently drawn into, has proved a most time consuming business. It : not yet through the press By the time this third edition appeared, Gray had been to Europe a second time, primarily to study specimens of North American plants de- posited in European herbaria. The only review seen of this third edition is by von Schlechtendal (Botanische Zeitung 9:157—158. 1851), who stated in translation: The author has used the sources that he found in Europe, as far as they were acces- sible to him, but he has noc incorporated all of the new views into his own, rather he followed in many points older views, namely those of A. P. de Candolle. This textbook will also enjoy a wide circulation. EDITIONS FOUR AND FIVE OF THE TEXT-BOOK (1853, 1858) New editions of the textbook, the fourth (1853) and the fifth (1858), the atter under the title Latroduction to Structural and Systematic Botany, appeared during the next decade. The preface to the fourth edition (1853, p. vi) 1s the same as the one in the third (1850), but with the added notation: — The changes ... are comparatively small; consisting of corrections and minor alter- ations, especially in the parts which relate to Vegetable Anatomy and Physiology, and in the addition of a short Chapter on the Fecundation of Cryptogamous or Flowerless Plant Stuckey, Asa Gray's botanical textbooks 785 The preface to the fifth edition (1858, p. [iii}) is similar to the one in the fourth edition (1853), but contains the following additional information: _, the structural and physiological part of the work, and the chapters on the Pracinles of Classification and of the Natural System, have been again almost e tirely rewritten, and such changes made as the advanced state of our oes required, or the author's continued experience in teaching has suggested. This has been done without increasing the extent of this part of the volume, which, consid- ering the limited time devoted to the study in our colleges, &c., is found to be as full as is desirable for a text-book. No reviews have been located for the fourth edition; a short review no- tice for the fifth edition (1858) appeared in the April number of the North American Review (86:587. 1858). The anonymous writer noted the immea- surable superiority of the natural system ona scientific basis, but that only a very brief skeleton-outline of the Linnaean system was provided in the book. The reviewer believed a more extensive account of the Linnaean sys- tem should have been included, because of an old love for that system. The Linnaean system was helpful in the identification of plants in flower by the novice in botany. Despite this exception, “Dr. Gray's Text-Book tn its present form exists ... [as a] more perfect or satisfying manual for the student or the proficient {individual} in botanical science.” Another writer, Isabella James, in the October number of the North American Review (87:326. 1858) compared this fifth edition with Gray’s first edition of 1836, pointing out that this edition (1858) is twice the size, printed on better paper, entirely rewritten, and supplied with numerous illustrations, “from the hand of Mr. Isaac Sprague, who has received the well-merited title of the most accurate of living botanical artists.” This writer also stated that Gray's Bo- tanical Text-book gained consideration abroad and “that for some years it was used as the class-book in the University of Edinburgh—a compliment seldom paid to American works, Gray’s textbook was essentially written for college students and pre- oping and changing ideas in botanical science. — sented the rapidly deve Consequently, a great void existed for younger students who needed a sim- pler work, and by the 1850s most secondary and elementary schools in the country were incorporating botanical study into their curriculum. By 1855 Alphonso Wood was industriously reaping profits with his C/ass-Book, now in its 41st printing (Lyon 1939, p. 82). What was Gray to do about this gap at the elementary level of the textbook series? WRITING BOOKS FOR YOUNGER MINDS (1857, 1858) Gray's publisher, G.P. Putnam and Company, had financial difficulties in 1854, and Gray shifted his books to Ivison and Phinney, New York, who were active textbook publishers. Under pressure from them, Gray wrote 796 Sipa 17(4) his Ferst Lessons in Botany and Vegetable Physiology (1857), designed for high school students (Fig. 4). It was reprinted over several years, with the title changed to Gray's Lessons in Botany, and later combined with his book Field, Forest, and Garden Botany (1868) to make a single volume called Gray's School and Field Book of Botany (1868) (Appendix II). Gray believed that the basics of Botany was “one of the most generally interesting of the Natural Sciences, {and} surely ought to be taught, and to be taught correctly, as far as the instruction proceeds.” These words came from his Fvrst Lessons (1857, preface (p. [iut). The book was . intended for the use of beginners, and for classes in the common and higher schools, ... {The} Lessons are made as plain and simple as they well can be, . _ have fan mancheals far enough to make the book a genuine Grammar of Botany ee Veg- etable Physiology, and a sufficient introduction to those works 1 in which the plants Thi are described. . of a country—especially of our own vork is complete in itself, as a school-book for younger classes, and even ee the Se of our higher seminaries... . {The book} comprises a pretty full account of the structure, organs, growth, ane reproduction of plants, and of their important uses in the scheme of creation... . The book is also intended to serve as an introduction to the author's Manual if ‘ihe Botany of the Northern United States ... and to be to it what a grammar and dictionary are to a classical author... . a full Glossary, or Ditiinan of Terms used im describing Plants, is added to the volume. Pennsylvania botanist Thomas C. Porter (American Journal of Science and Arts 73:439—440. 1857) wrote a most favorable review of Gray's First Les- sons (1857): Good ae books in any branch of science are rare, ... Indeed, a good text- book of science, like a picture or a poem, is a work of art, the creation of which requires extensive knowledge, abundant resources and a generous and enthusiastic ove of nature. And, even where these exist, none but a master can rightly seize the leading facts and principles and exhibit them in language at once precise, clear and simple—can be brief without being obscure and thorough without being prolix— can so fashion his teachings as co charm the youthful mind and lead it on impercep- tibly into the very heart of the science—can furnish fresh and original illustrations of familiar objects in drawings and engravings, and give them that perfect accuracy which only an eye trained to a close scrutin y of forms can fully appreciate. Hence really excellent elementary books are rare - ed; There is a special es tion, therefore. when these requisites are fulfilled, as 1s true in the case of the “First Lessons in Botany and Vegetable Physiology” from the pen of Dr. Gray. It is a model of its kind; and meeting a want long and widely elt, it must sooner or later win its way into all our schools and seminaries of learn- ing... . Considering the intrinsic value of these books, the great care, labor and talent which have been devoted to their preparation, the good style in which they are published, and the great number and excellence of the their illustrations (from drawings by Mr. Sprague), the wonder is, that they can be sold at so moderate rates. We trust that they may meet with a circulation, proportioned to their real merit. 1 England, an anonymous reviewer writing in the Journal of Botany and Kew Garden Nie (9:154—160. 1857) stated: Stuckey, Asa Gray’s botanical textbooks 787 sornegagrmeger pment anata” \ THE OHIO STATE UNIVERSITY . HERBARIUM LIBRARY GIFT OF EMANUEL D )LPH j FIRST | LESSONS IN BOTANY VEGETABLE PHYSIOLOGY, ILLUSTRATED BY OVER 3860 WOOD ENGRAVINGS, FROM ORIGINAL DRAWINGS, BY ISAAC TO WHICH 19 ADDED A COPIOUS GLOSSARY, 1 DICTIONARY OF BOTANICAL TERMS. 4 By ASA GRAY, FISHER PROPESSOR OF NATURAL HISTORY IN HARVARD UNIVERSITY. { Se eons NEW YORK: 4 IVISON & PHINNEY asp G. P. PUTNAM & CO., 32) BROADWAY CHICAGO: S.C. GRIGGS & CO., 11) Lake Street. Fic. 4. Title page of Gray’s First Lessons in Botany (1857). { | 788 Sipa 17(4) Professor Gray has a thorough knowledge of his subject; he is a successful teacher, a lucid and accurate writer, and a most careful compiler and analyst ... The genera — character of his work ... {is} both scientific and i and as perfectly suited to its object as any work of the kind we have seen... . It is che first good indigenous American work on Elementary Botany, and it is written to meet an urgent want 00k adapted for the use of the classes in the Common and higher schools of se United States; ... The illustrations are good ... of the subjects ... , and are said to ye, almost tone exception, original, ... [wit ig the skill and botanical accuracy of r. Sprague, who stands at the head a ee in botanical artistes. We ... highly praise Mr, Sprague’s exertions in making the illustrations, wherever possible, from American plants. — ofa > M The reviewer made some critical remarks about the illustrations, the order of presentation for certain selected terms referring to the organs of plants, che failure to define a few words used, and the sketchy account of the procedures for collecting plants and making an herbarium. D.EL. von Schlechtendal (Botanische Zeitung 16:151. 1858), the German botanist, made several specific comments saying that the book: (1) Explained everything as clearly and correctly as possible, (2) Offered a true gram- mar of botany and plant physio ogy, (3) Provided a satisfactory introduction to those books in which the plants of an area are described, (4) Drew examples from the North American flora, and (5) Would serve quite well for instruction, _— ut for readers in Germany, the dose of anatomy and physiology should be richet Because Gray’s textbooks were considered by many educators as too ad- vanced for a large portion of the textbook market, Gray worked many eve- nings preparing a simpler book, How Plants Grow (1858), written for “young people and common schools,” as stated on the title page (Fig. 5). It too, was reprinted for many years. These two elementary books differed from the advanced textbook only in being less complex, but not in content or mode of presentation. Gray’s effort was to make real science easy and simple for the younger minds. Gray realized $500 to $600 per year from the sale of his textbooks during this period. Most of his intelleccual pursuits during the 1850s were involved with the writing of these textbooks. Gray’s atti- tude about producing them was stated ina communication (J.L. Gray 2:438. 1893) of 26 April 1856 to his British botanical friend George Bentham: My last book [How Plants Grow (1858)} in elementary botany is now just off my hands, and will be out ina fortnight. Phope it will be of use. Forgive me for writing horn-books, and lam now done with that sort of work. There were several convinc- ing reasons for doing it. Gray's reasons for writing textbooks for the elementary student are fur- ther elaborated in a letter of 15 May 1857 to the Rev. R.W. Church, who was the rector of Whaley, a village in Scotland J.L. Gray 2:429—430. 1893): An acquaintance en route for Scotland has offered to take some small parcels for me. Among them is one I have taken the liberty to address to you, a copy of a very StucKEY, Asa Gray’s botanical textbooks bat eR? Notany for Poung People and Common Srhools. HOW PLANTS GROW, A SIMPLE INTRODUCTION TO STRUCTURAL BOTANY. WITH { A POPULAR FLORA, OR AN ARRANGEMENT AND DESCRIPTION OF COMMON PLANTS, BOTH WILD AND CULTIVATED | ILLUSTRATED BY 500 WOOD ENGRAVINGS. i AERA, 4 oe \ 4 by ; : Sy By ASA GRAY, MD, &. | FISHER PROFESSOR OF NATURAL HISTORY IN HARVARD UNIVERSIaY. ay 6 rs re" 2 cae ie >| oe 3 § y 4 oy ‘) | ge i NEW YORK: IVISON, BLAKEMAN, TAYLOR, & CU., Fic. 5. Title page of Gray's How Plants Grow (1858). elementary book [First Lessons (1857)]} I have prepared as an introduction to my favorite science, finding there was no one in use here which I chought fit to put into the hands of young beginners. Here botany is taught, somehow or other, in most schools, and generally by incompetent teachers from wretched books, 1.e., those S 9 — used in the ordinary schools and for young peop I have endeavored, in the little book I send you, to make real science as easy 790 Sipa 17(4) and simple as possible. I doubt if I have yet aimed low enough; but the book seems to take, and promises to be useful. As stated in Gray's introductory comments (p. 2) to How Plants Grow (1858), “this book is intended to teach Young People how to begin to read” about the vegetable kingdom. Conceived as the first of a two-part work on “Botany for Young People and Common Schools,” this book has two major parts. The “Firse Part” considers how plants grow, and what their parts or organs are; how plants are propagated or multiplied in numbers; why plants grow; and how plants are classified, named, and studied. The “Second Part” consists of a popular flora for beginners for classifying and describing com- mon wild and cultivated plants of the country. Following these parts is a dictionary of botanical terms used in this book Slightly over a year later on 1 June 1858, after How Plants Grow (1858) had been published, Gray sent the Rev. ms a coy of that book, with the following letter JJ.L. Gray 2:444—445. 1893 Last week the publishers at my request, sence ... a copy of a new and more elemen- tary book [How Plants Grow (1858)} of mine than the one you are pleased to compli- ment. I intended that as a kind of horn-book, which Dr. Hooker insists it is not: and as something more simple was wanted here, to lead the way both to the “Les- sons” and especially to the “Manual,” which is rather strong for beginners, I have tried again, and you will see the result. I should have made the little ‘ ‘Popular Flora” fuller if the eens had allowed more room. Having last year reédited my “Botanical Textbook” ..., 1 have now done n my part in elementary botanical writing, and | return with zest to my drier investigations An anonymous reviewer (1858) in the American Journal of Science (76, 2nd ser., 26:139—140. 1858) wrote concerning How Plants Grow (1858): The work is simple in style, and beautiful in ics illustrations. While teaching with clearness the details of the subject, ic is constantly bearing the mind, by simple ne itions, above these details to higher ee and principles, and preparing it fo uller survey of the science in the more extended works of the author's series. ce excellence of the volume consists in its being really “science made easy’—not by culling out “interesting facts” to attract, and tying them artfully together, but by presenting che system of fundamental truths in a manner intell igible and attractive to the young mind. In the October number of North American Review (87:322—326. 1858), Isabella James stated with reference to Gray’s two books for young people, The First Lessons (1857) and the How Plants Grow (1858 The excellence of these elementary books is what every one who has had any knowledge of Dr. Gray’s previous works must have expected; for toa perfect knowl- edge of his subject he adds a clearness and exactness of style seldom met with, and the power of condensing in a few words a great amount of information. As an accu- rate analyst, he has received a just meed of praise from all foreign botanists. Stand- ing as he does at the head of the science in our own country, and scarcely inferior to Stuckey, Asa Gray’s botanical textbooks 134 any botanist of the Old World, we consider it a subject of congratulations that he has found time, among his multifarious avocations of a high order, to write two books expressly for the young. The urgent need of a botanical primer, to introduce so charming a study into the school-room in a form attractive to children, has long been felt. The habits of observation, research, memory, and judgment ... are the very qualities which are most needed in every day’s practical life; and if by proper educational training these discriminating powers are quickened at an early age, they bs continue a constant and unfailing source of instruction and delight. v Plants Grow (1858)} ... may be truly called a wonderful book. The style is fee hly scientific, and yet so clear and simple that even a little child can understand it, and become interested in the subject. It is not, like some popular — works, written so far down to the comprehension of the young as to become a weak mixture, where science is diluted to the lowest possible standard, as if to keep the young mind for ever in swaddling-bands; but it is arranged and designed so to educate and expand the intellect as that a young person of ordinary capacity 1s soon able to seize upon, and comprehend, the principal features of the vegetable world, When a work like this, combining in so rare a manner the two sterling quali- ties of excellence and cheapness, adapted expressly to young persons, and arranged for the use of schools, emanates from the highest botanical authority, we trust a discriminating public will show a just appreciation of its merits, by discarding al the trashy volumes now in use, and adopting one which aa in a small space su the learning and research of a life devoted to the subje Botanical historian Frederick Brendel (1879, p. 8 wrote, “For until Prof. A. Gray’s popular book, ‘How Plants Grow’ appeared in 1858, nota single work of any importance [for young people} was published in this country.” In France, in the Bulletin de la Societé Botanique (15:655—656. 1858), an anonymous writer noted that “Lexecution typographique en est fort remarquable,” for the 500 figures drawn by Isaac Sprague, the most accurate botanical illustrator of that time (Rudolph 1990). FOUR-PARTS PROJECTED FOR THE TENT-BOOK $ SIXTH EDITION (1879, 1885) cy) years passed before Gray returned in the late 1870s to the task of revising the fifth edition of his Text-book (1858). In 1872, Gray contem- plated doing the revision, but time proving insufficient, he asked George L. Goodale, his new colleague, to help in its preparation. Goodale came to Harvard University in the fall of 1872 to assume some of the regular classes taught by Gray. During the following year Gray retired from classroom teaching, and Goodale received the appointment as Fisher Professor of Natural History. Much new information had become available in the now- expanding and more widely developing botanical discipline since the fifth edition was published, and Gray conceived that the new version would need to be expanded into several parts. A four-volume effort was envisioned, and the first volume on Structural 792 SIDA 17(4) Botany (1879), concerning the morphology, taxonomy, and phytography of phanerogams was, compared to previous editions, entirely rewritten by Gray According to the preface (p. [iit}): The present treatise is intended to serve as a text-book for the hig instruction. To secure the — rer and completer whole range of subjects, it has been decided to divide the work into distinct volumes, each a treatise by itself, which may be indepen- dently used, while the whole will compose a comprehensive botanical course. This volume, on the Structural and Morphological Botany on Phaenogamous Plants properly comes first. It should thoroughly equip a boranise for the scientific pros- ecution of Systematic Botany, and furnish needful preparation to those who proceed » the study of Vegetable Physiology and Anatomy, and to the wide and variec department of Cryptogamic Botany. After the book was available, Gray mailed one to George Engelmann, 22 May 1879 (J.L. Gray 2:687. 1893), noting: “I send you by mail a copy of my new “Text-Book.’ You see I relegate to other hands the anatomy, physi- ology, and cryptogamia,—glad to be rid of them.” Indeed usual botanical information was included in the sixth edition, , not all of che first volume of the new In the extensive review of the Sixth Edition, Gray’s colleague George L. Goodale (American Journal of Science and Arts 118:73—76. 1879), reminded the reader that in Gray’s first book, Te Elements of Botany (1836): Phe plan of this early work was generous in its scope, and was philosophically developed. A morphological basis was adopted as the only safe one on which to build, and upon this a symmetrical superstructure was erected. It was no ordinary sagacity which led a young botanist, without experience in teaching, to select a method which has needed no essential change for forty years, and which is to-day generally accepted as best adapted to elementary and advanced instruction. “Text-book,” which was developed from the earlier through several editions .. The “Elements of Botany,” has passed . A still further development of tl a division into separate volumes, ... [of whic umes planned}... . The re plan ... necessitated ch this volume is the first of four vol- e present ae ... 18 adapted ... to the wants of the ad- cae student and the working botanist alike. An eu ee author in the Bu/letin of the Torrey Botanical Club (6:317— 320; 328-329. 1879) wrote that Gray’s Part I of the Sixth Edition (1879) will undoubtedly for years to come be the main text book on the science in our colleges and scientific schools, containing as it oes the latest results in chis depart- ment, treated with a mastery and a clearness of which we know no equal Henry Trimen, the editor of The Journal of Botany British and Foreign new ser. (8:253—254. 1879), published at the British Museum in London also reviewed Gray's sixth edition, Part 1 (1879). After noting that the book “is strictly devoted to the morphological anatomy of Phanerogams with sec- tions on the principles regulating their classification, description, and no- menclature,” Trimen wrote: Stuckey, Asa Gray’s botanical textbooks 793 It was, indeed, the masterly and philosophical treatment of an sections of the science which always gave to the Text-Book its special value; they have been particu- larly the branches to which the author’s long and laborious life has been devoted, botanists will receive this fuller treatment of — and it is with great satisfaction that them at his hands. The whole has been entirely re-written, and it 1s scarcely necessary to say, well written. Dr. Gray is able to convey strictly technical instruction in the attractive and easy style only reached by a few masters. Charles E. Bessey, one of Gray’s former students who became a distin- guished teacher of botany, wrote in the American Naturalist (14:870. 1880), that Gray’s Sixth Edition (1879) ranked “as one of the best books on struc- tural botany extant.” An anonymous reviewer in the Bulletin de la Societé Botanique de France (26:165. 1879) commented on its successful perfection, as were each of the previous editions. Later, George L. Goodale published the second volume on Physiological Botany (1885), but the two remaining planned volumes were never pub- lished. Gray’s other colleague William G. Farlow was to write the third volume on an Introduction to Cryptogamic Botany, both Structural and System- atic, and cee. the fourth volume on a Sketch of the Natural Orders of Phane 's Plants, Including Their Special Morphology, Classification, Dis- atone a Products. Farlow (1890) wrote: “It is deeply to be regretted that he [Gray} was never able to write this volume, for it would have en- abled him to present the general views on classification derived from a long and exceptionally rich experience.” He did not comment on why his own volume number three also was not written An anonymous reviewer of Goodale’s Phystological Botany (1885) 1n the Botanical Gazette (10:392—393. 1885) wrote: ere is a really new book; not new in its facts, of course, but essentially new in end the arrangement, and thoroughly original in the presentation of them } ver wide range of literature from which these facts have been gathered ... [and] ae references to the literature, and in many cases a citation of the passage 1n point . The reader will also be impressed with the comprehensiveness of the treatment. J.C. Arthur (1895, pp. 369-370) spoke of Goodale’s Part II as a work covering a broad scope, including histology, anatomy, and ecology, as well as physiology proper. Of the last item, he recognized it as ._ by no means the most conspicuous part of the book. The encyclopedic fulf{l}ness of the work better adapted it for a reference-book to accompany a course of lectures than as a textbook. It greatly helped the science in America however, especially as it stimulated experimental study by a set of laboratory exercises given as an appen- dix... . [Goodale’s book} a . first, and to the present {is} the only treatise on physiological botany by an American author. 794 Stipa 17(4) MORE BOOKS FOR ELEMENTARY STUDENTS (1872, 1887) Gray’s earlier book How Plants Grow (1858), was as stated on the title page, a simple introduction to structural botany with a popular flora illus- trated with 500 wood engravings. Furthermore, the title page identified the book as “Botany for Young People and Common Schools.” Having years earlier written to the Rev. R. W. Church of Scotland that he had finished his writing of textbooks for children, the urge; however, must have re- turned. Gray now wrote a companion book to How Plants Grow, calling it How Plants Behave (1872), which as stated in his preface ( pp. [vii—viti}) was a topic to which he “long wished to introduce pupils of an early age.” He described that topic as being . admirably adapted, while exciting a lively curiosity, to stimulate both observa- tion and thought, ... Not only {should young people ask about] ‘How Plants Grow,’ but f[also} ‘How Plants Act.’ [The plant phenomena} in certain important respects {are] easy to be observed,—everywhere open to observation, but (ike other com- mon things and common doings) are very seldom seen or attended to. [It adds} some very interesting chapters to the popular history of Planc-life... . written with a view to elementary instruction ... with all practicable plainness, the subjects here presented are likely to be as novel, and perhaps as interesting to older as to younger readers. [Gray hoped that} this little creatise, designed to open the way for the young stu- denc into this new, and, I trust, ateractive field, may be regarded as a supplement to the now well-known book, [How Plants Grow. Gray discussed three major topics in How Plants Behave (1872): (1) The motions of plants including climbing, (2) plants that are carnivorous, and (3) che fertilization of plants by insects. These themes are discussed in “the simplest and {yet} most comprehensive statement” made by the author, as noted by the Rev. C. E. Bolles in the American Naturalist (6:475—477. 1872). This reviewer stated further that Dr. Gray at least he is careful, all che way .. appears to recognize a personality in plants through, to show that the actions which he explains are che result of the plant’s will; and just as far as botanical science allows, he assigns the reasons for them. =) An anonymous reviewer in the By/letin of the Torrey Botanical Club (3:35. 2), noted: he book ts attractive in form, and we hope {it} may lure the young to our favorite study. To more mature minds, imbued with a love of Nature, it cannot fail to prove of great interest. Following Gray's death in 1888, Joseph D. Hooker (1888) in his account of Gray’s life wrote that these two smaller books “for charm of matter and style, have no equal in botanical literature ... In 1887, just before departing on his last European journey, Gray finished Sruckry, Asa Gray’s botanical textbooks ces. a small book, The Elements of Botany ... (1887) containing in an abbreviated format the substance of the earlier Structural Botany Textbook (1879), as well as some chapters on cryptogams. He chose the same title for this book as he had for his earliest book in 1836, but since those copies were long out-of- print and rarely seen, he saw no inconvenience by reusing the former title The book was identified on the title page as being “for beginners and for schools.” It also replaced his Lessons in Botany and Vegetable Physiology (1857), published 30 years previously. As stated in the E/ements’ pretace (pp. {111} iv), the book was for beginners learning structural botany and the prin- ciples of plant life, mostly as they pertained to flowering plants, “with which botanical instruction should always begin” to quote Gray. The book was also to be a companion and interpreter for his Manuals and Floras this treatise should teach that che study of botany is not the learning of names and terms, but the be - knowledge and ideas... . the Glossary ... has Gee enlarged. udef{s} not merely the common terms of botanical description but also many w a are unusual or obsolete George L. Goodale, in his review in the American Journal of Science and Arts (134:495. 1887), described Gray’s E/ements (1887), as a new work differing in many features from any of its predecessors. It embodies the grammar of Orga- nography, the first principles of Vegetable Physiology, and of Botanical classification, and gives with the help of an adequate glossary, directions for the description of plants. The bringing so much within so narrow a compass has been accomplished by giving the kernel of every topic with out any husks ... It is ... adapted to the wants of classes and of those who are obliged to study gi themselves. Charles E. Bessey, recognized as a master of the botany teaching profes- sion, wrote a review in the American Naturalist (22:46-47. 1888), compar- ing in numerous details Gray’s first E/ements (1836) and this E/ements (1887) For example, in the first book, 18 pages were written on the Linnaean Sexual System, and 44 pages were devoted to the Natural System, which was nec- essary to justify its inclusion. In this book a short paragraph is all chat remains of the discussion of the Linnaean System, and less than two pages suffice for the Natural System... . Not only do these con- trasts show us what advances have been made in botany in half a century, but a comparison of these two books shows, still more, the remarkable ae and peren- t is not given to many men to live ou nial youth of the master-mind who wrote them. to see such great changes in the aspect of science as has been the see: fortune of Dr. Gray, and still fewer have had the strength or ability to adapt themselves to the new views and theories. Walter Deane (1888, p. 65), Gray’s friend and obituary author, noted that these “two books are a most fitting Alpha and Omega to his industri- ous life. [Of the latter}, he always spoke with much enthusiasm in regard to this revised work and seemed much pleased with the result 596 Sipa 17(4) jon In the Botanical Gazette (13:41. 1888), an anonymous reviewer statec concerning Gray's E/ements (1887): No text-book has been more widely used than the one of which this is a revision; and nothing could be more appropriate than for the author himself to give a final revision, which has been long demanded by our rapid advance in knowledge. There can be no doubt that this is the best book for its purpose that we now have. This writer pointed out errors in the use of anatomical terminology and the uded with a final statement: — choice of poorly illustrated “old figures,” bue conc Although more concession to the results of modern botanical work might have ~~ deen expected, the auchor may have been wisely conservative, and his justly great name insures a book that can be trusted. British botanist James Britten, writing in the Journal of Botany British and Forergn (26:92—93. 1888), provided a review of Gray's Elements (1887): — This new edition of an old favourite ... {is} che last which will come from the hand of the venerable and kindly writer... . Dr. Gray's work, throughout his long and active botanical career, has been eaneked by clearness and consistent usefulness; and these characteristics are as noticeable in a work like the present as in the greater undertakings by which his position among botanists has been securec FURTHER CONTEMPORARY COMMENTS OF EVALUATION Charles Bessey (1888, p. 280) considered Asa Gray “a great man.” Asa New York village school boy, Gray was “so far removed from all incentives to the study of science, {nevertheless he became} the future leader of one of the great branches of science in America.” With Gray’s death on 30 January 1888, the pen of nineteenth century America’s greatest botanical textbook writer was laid to rest. Joseph D. Hooker (1888) wrote after Gray’s death: . [the Elements ea (1836)} ... was the first of a series of editions of a work dhach ras been for nearly half a century the text-book of schools and colleges through- out the United States, and the latter issues of which have been generally recom- mended by the botanical professors of the United Kingdom as the best of its class. Gray's colleague in cryptogamic botany at Harvard University, William G. Farlow (1890, pp. 772-773), summed Gray's contributions to the writ- ing of botanical textbooks as follows: No better text-book on the subject had ever been written in the English language than Gray’s “Text-Book” in the original form; and, although botanical instruction 1s now very different from what it used co be, it is still crue chat, as an introduction to the study of Phaenograms [i.e. Phanerogams}, the group to which beginners naturally turn their attention, the later “Structural Botany,” is likely co hold its own for some time to come. Dr. Gray had the rare faculty of being able to nee himself to all classes of readers. With the scientific he was learned, to the student he was instructive and suggestive, and he charmed the general reader by the many beauty of his style, Stuckey, Asa Gray’s botanical textbooks 197 while to the children he was simplicity itself. The little books, “How Plants Grow,” and “How Plants Behave,” found their way where botany as botany could not have gained an entrance, and they set in motion a current which moved in the general direction of a higher science with a force which can hardly be estimated. His: scientific friends, especially those abroad, sometimes blamed him for spending time in popu- lar writing; but he may have understood himself and his surroundings better than they. With him botany was a pleasure as well as a business. Few wrote as easily as he, and, so long as he spent most of his time in higher work, he certainly had a right to amuse himself with writings of a popular character if he chose. As it was, he interested a multitude of readers in the subjects which he had at heart ... . Later, Farlow (1895, pp. 173, 174-175) further wrote: The numerous editions of Gray's Text-Book made his name known in all the higher schools in the country, ... He left a great nation in which, very largely through his exertions, the value of botany had become generally recognized and in which a crowd of young workers had arisen anxious to carry out the good work ... . SUMMATION Asa Gray’s recognition as the foremost nineteenth-century botanist of North America was earned not only because of his creative research and investigations on the flora of that continent, but also as an author of seven editions of his Manual of the a of the Northern United States (1848— 1908). These books described the species and provided keys for their identification. Gray also may be recognized with this distinguished utle because of his authorship of 11 different editions of textbooks of botany. In these books Gray developed botany into a more precise science, as he drew from the great Swedish botanist, Carolus Linnaeus, the French botanists A. P. de Candolle, A. L. de Jussieu, C. F. Mirbel, and Achille Richard, and the Englishmen Robert Brown and John Lindley. He ignored or chose not to extract from the German writers, who were well advanced in experimental laboratory approaches in the teaching and researching of botany. These ap- proaches had not yet been inaugurated in the schools teaching botany in the United States except for the early efforts of Charles E. Bessey. Gray’s textbooks were original and fulfilled che needs of the time. They were in demand as both the purchasing population and the scientific infor- mation about the plants expanded. New editions with broader dimensions for different audiences became a necessity, and consequently previous edi- tions were revised, rewritten, or expanded. After 1858, his books under- went many printings. As an example of the usefulness and demand for Gray’s textbooks, Stuart G. Noble in his A Hastory of American Education (1954) wrote that Gray’s How Plants Grow (1858) was continued in use until 1900. From 1896 to 1900 it was used in more schools than any other text... . che chief emphasis of the text was upon anatomical structure and classification. Numerous technical terms were used and 798 Stipa 17(4) the logical treatment was maintained. Under this leadership instruction in botany ——-. became formal and highly technical. After Gray's death, no one continued preparing revisions of his textbooks, and thus both their use and usefulness ceased. At the end of the nineteenth century, the German modeled botanical teaching laboratory was developed in the Uniced States (Rudolph 1996). New textbooks were designed for these new teaching situations. ACKNOWLEDGMENTS Special thanks go to: (1) William R. Burk, Biology Librarian, Univer- sity of North Carolina at Chapel Hill, who located most of the book re- views and read, edited, and word processed the manuscript; (2) Judith A. Warnement, Librarian, Harvard University Botany Libraries, Cambridge, who provided some of the book reviews and photocopies of prefatory pages of several textbooks; (3) Jimmy Dickerson, Chemistry Librarian, Univer- sity of North Carolina at Chapel Hill, who translated the reviews of D.EL. von Schlechcendal into English. Most of Gray’s textbooks discussed in this paper are in the Emanuel D. Rudolph collection in the Herbarium Library at The Ohio State University. Some of the reviews and textbooks were ob- tained for examination from the Marston Science Library at the University of Florida, Gainesville. on — APPENDIX | A CHRONOLOGICALLY ARRANGED LIST OF ASA GRAY’S TEXTBOOKS OF BOTANY 1836. Elements of Botany. G. & C, Carvill & Co., New-York. xiv, 428 pp 1842. The Botanical Text-book for € olleges, Schools, and Private Students: Comprising Pare 1. An Botany. Part IH. The Principles of Systematic Botany; Introduction to ena and Physiologica With an Account of the Chief Natural Families of the Vegetable Kingdom, and Notices of the Principal Officinal or Otherwise Useful Plants. Hlustrated with numerous engravings on wood, Wiley and en New-York; Little and Brown, Boston. xii, {13]—-413 pp. 1845. The Botanical Text-book, for Colleges, Schools, and Private Students: C omprising Part I. An Introduction to Structural and Physiological Botany. Pare II. The Principles of Systematic Botany; I Wich an Account of the Chief Natural Families of the Vegetable Kingdom, and Notices of the Principal Usetul Planes. 2nd ed. oe ated with more than a thousand engr gs on wood. Wiley and Putnam, New-York. {1-9}, 10-509 pp. 1850. The Botanical Text-book, an oe to Scientific Botany, Both Struccural and System- atic. For Colleges, Schools, and Private Students. 3rd ed., rewritten and enk arged., iiusteaked with twelve hundred engravings on wood. George P. Putnam, New York. xii, {13]—-520 yp. 1853. The Botanical Text-book, An ee to Scientific Botany, Both Structural and System- atic. For Colleges, Schools, and Private Students. 4th ed. eae with twelve hundred engrav- ings on wood. George P. Pucnam & Co. one York. xit, [13}-5 8 pp. 1857. Firse Lessons in Botany ine Ve pei ene ee by over 360 woodcuts, from aoe drawings, by Isaac Sprague. To which is added ; a COprlous a or dictionary of borani- cal terms. Ivison & & Phinney and G. P. Putnam & Co., New York: 8. ©, Gr riggs & Co., Chicago. xii 236 p pp. [Reprinted as Gray's Lessons in Botany beer alas in 1868; “a rints listed in The National on Catan (1 ee ieee. 1860, 1862, 1863, 1864, 1865, 1866, 1867 . 1868, 1869, 1870, 1873, 1874, 1875; Reprint also listed in WorldCat (OCLC First Search): 1858, 1886}. Sruckgy, Asa Gray’s botanical textbooks 799 1858. Introduction co Struccural and Systematic Botany, and Vegetable Physiology, Being a Fifth and Revised Edition of The Botanical Text-book. Illustraced with over thirteen ena woodcuts. th ed. Ivison and Phinney, New York; S. C. Griggs & Co., Chicago. xii, [13]-555 pp. {Title 1858, copyright 18 : ] age peas in ie Neel si Catalog (1972): eee 1862 1865, 1866, 1869, 74,1876, , 1878; Reprint also listed in the ee University Library ae aoe se . aa in WorldCat (OCLC First Search): en 1858. ow Plants Grow, A Simple Introduction to Structural Botany. With a Popular Flora, or an Arrangement and Description of Common Plants, Both Wild and Cultivated. Botany for Young aan and Common Schools. Illustrated by 500 wood engravings. Ivison, Blakeman, Taylor, & Co., New York. 233 pp. [Also published in 1858 by American eae G see New York, with a. ».]. [Reprints listed in The Ne ie : nion Catalog (1972):1859, 1860, 1861, 1862, 1863, 1864, 1865, 1866, 1867, 1868, 1869, 1870, 1871, 1872, reo 1874, ; 6, 1879; Reprints also listed in the ae University Libary Online Catalog:1880, Isso; Re- aes also listed in WorldCat (OCLC First Search): 1881, 1906 1872 v Plants Behave: How They Move, Climb, Employ ae To Work for Them, &c. Botany tor oe People. Part II. Ivison, Blakeman, Taylor, and Company, New York and Chicago. viii, [9]-46 pp. {Also published in 1872 by American Book C pare New York, with 46 pp.]}. [Re- prints listed in The National Union Cate alog (1972):1873, 1874, 1875, 1900}. 1879. Gray's Botanical Text-book. 6th ed. Vol{ume} I. een Botany, or Organography on the Basis of Morphology; to Which Is Added the Principles of Taxonomy and Phytography, and a Glossary of Botanical Terms. Ivison, Blakeman, Taylor & Company, New York and Chicago. xil, 142 pp. [Reprints listed in The National Union Catalog (1972):1880, 1881, 1885, 1890, 1907 1887. The Elements of Botany for eee and for Schools. Gray’s Lessons in Botany. Revised e American Book Com », New York, Cincinnati, Chicago. vit, 9-226 pp. eee listed in The National Union C ‘he Ron a 1890, 1901] 875, 187 877 , 187 78 APPENDIX I GRAY’S SCHOOL AND FIELD BOOK OF BOTANY (1868), TWO BOOKS IN ONE Gray's School and Field Book of Botany (1868) consisted of two of Gray's books bound together in the Lessons in Botany (1857) and the Field. Forest, and Garden Botany (18068). First issued in L868, che two together formed a popular and eon hensive botanical work that could be used by begin- ning and advanced classes (Fig. 6). In reality, chis book was Gray's answer to Alphonso Wood's earlier challenge for a textbook that presented the terminology of botany along with a manual for field identification. Wood's Class-book (1845), issued over 20 years sages to 1868, and still being re- printed, was scill available for class use. ow. however, it was time fora book having a fresh eo t one => a that combined botanical lessons with floristic and eee information. According to publisher's preface: [Gray's book was} adapted to beginners and advanced classes, to Agriculture Colleges and Schools, as w ell as to all other grades in which the science is taught; ... also adapted ... asa hand-book to assist in analyzing plants and flowers in field study of botany, either by [students in] classes or individuals It is a Grammar and Dictionary of Botany, and supplies a great desideratum to the Botanist anc Bot sofeal Teacher, there being no similar class-book published in this country. REFERENCE 1868. Gray's School and Field Book of Botany. Consisting of “Lessons in Botany {18571," and “Field, Forest, and Garden Botany {1868],” bound in one volume. Ivison, Blakeman, Taylor & Co., New York and Chicago. Lessons, xii, 236 pp.; Field, 7-386 pp. {Reprints listed in The National Union Catalog 197 2):1869, 1870, 1871, 1872, 1873, 1874, 187 1876, 1877, 1878, 1879, 1880, 1881, 1887, 1890, 1895, 190?}. iRencints from 1887 and the following years, consist of Gray’s Elements of Botany ({1887)), rather than ¢ — als S00 Stipa 17(4) GRAY’S SCHOOL AND FIELD BOOK BOTANY. CONSISTING OF “LESSONS. IN BOTANY,” AND “FIELD, FOREST, AND GARDEN BOTANY,” BOUND IN ONE VOLUME. ASA GRAY, FISHER PROFESSOR OF NATURAL HISTORE IN HARVARD UNIVERSITY, IVISON, BLAKEMAN, TAYLOR & CO., NEW YORK AND CHICAGO, al =~ » 6. Title page of Gray's School and Field Book of Botany (1868). Stuckey, Asa Gray's botanical textbooks 801 previous Lessons in Botany (1857). The title page, however, for the entire book referring to the two combined books in printings after 1887 continued to carry the subcicles, Lessons in Botany and Field, Forest, and Garden Botany). REFERENCES AMERICAN TEXTBOOK PuBLIsSHERS INstTITUTE, THE. 1949. Textbooks in Education. A Report from the American Textbook Publishers Institute to Its Membership, Its Friends, and Others ... The American Textbook Publishers Institute, New York City. x1, 139 ete , J.C. 1895. Development of vegetable physiolog ry. Science, n.s. 2:360—373. [Also published in in Proc. Amer. Assoc. Advancem. Sci. 44:{163]-184. 1896]. BARTLETT, 1941. Asa Gray’s nonresident professorship. The ia es of the Uni- versity in ae Arbor. ane oe Alumnus Quart. Rev. 47:215- Bessey, C.E. 1888. Asa Gray. Amer. Naturalist 22:280—281. BreENDEL, FE. 1879-1880. Hiscorical sketch of the ee coe in North America — » 1840. Amer. Naturalist 13:754—771. 1879; 1880. Reprinted, 1978. De- nee net of Botany in Selected Regions of ane peas before 1900. Edited ae an introduction [4 ee } by Ronald L. Stuckey. Arno Press, A New York Times pany, New York. 1 volume (variously paged). A collection of papers, published ened 1879-1909, aoe - Ronald L. Stuckey and reprinted in book form as part of a series, oe and their World ComPILers. 1¢ he National one Catalog Pre-1956 Imprints. Volume 211. Mansell eer Limited, London; The American Library Association, Chicago, Illinois. on Deane, W. 1888. ie Gray. Bull. Torrey Bot. Club 15:59-7 ae A.H. 1959. Asa Gray 1810-1888. The Belknap Pies: ot Harvard University Press, Cambridge, Massachusetts. x, 505 pp. Fartow, W.G. ee Memoir of Asa Gray. Annual Rep. U. S. Natl. Mus. 1888:763-783. «1895. Memoir of Asa Gray, 1810-1888. hee before the National Acad- emy, April ro soe Bio. Mem. Nat. Acad. Sci. U.S 3:161-175. GoobaLe, G.L. 5. Physiological botany. I. Outlines i af histology of phaenogamous plants. II. ene pegs ig 's botanical textbook. Sixth edition. Vol. II. Ivison, Blakeman, Taylor & Co., New York. xxi, 499 pp. + Practical Exercises, 36 pp. Gray, A. 1848. Manual of es of the northern United States, from New England to Wisconsin and south to Ohio and Pennsylvania inclusive, (The mosses and liverworts by Wn Sullivant,) Arranged according to the natural system, ... James Munroe and Cue Peach & Cambridge, [Massachusetts]; John Chapman London. Ixxii, 710 pp. _____.«s1 868. Field, forest and garden botany: A simple introduction to the com- mon plants of the United States, east of the MaskepeL Both wild and cultivated. Ivison, Phinney, Blakeman & Co., New York; S. C. Griggs & Co., Chicago. 386 p Revised 1870. _. «1880. John Carey. Amer. J. Sci. Arts, 3rd ser., 19:421-422. [Also Sad lished in Scientific Papers of Asa Gray Selected by Charles Sprague Sargent, vol. 2, pp. 417-418. Houghton, Mifflin and Company, Boston and New York. 2 vols. (l:viti, 397 p.s 2: iv, 503 pp.).4 Gray, J.L. oe 1893. Letters of a Gray. — Mifflin and Company, Boston and New York. 2 wae (1: 368 pp.; 2: pp. 369-83 H[OOKE RI, ole PH] D. 1888. ae Asa pat Nature 37:375— Kuun, T.S. 1970. The structure of scientific revolutions. Sal ed., crt The Univer- sity of eine Press, Chicago. Peer btn: Unity Sei. 2(2):xi1, 210 7 802 Stipa 17(4) LINNAEUs, C. 1751. Philosophia Botanica; In Qua Explicantur Fundamenta Botanica ... . Godofr. Kiesewetter, Stockholm. 362 p Lyon, C.J. 1939. A 50 edition best seller: Alphonso Wood, class of 1834, wrote botanical class books that reached a circulation of 800,000 copies. Dartmouth Alumni Mag. 31(March): 18, 81-83 Nose, $.G. [1954]. A history of American education. Revised ed. Holt, Rinehart and Winston, New York. xx, 5352 pp Rupotpu, E.D. 197 La. The origin and development of the botanical textbook and its Ameri- can aspects, p. 6. In: Abstracts of Contributed Papers ena before The Ohio Acad- my of Science 1971 History of Science Meeting, Marc 1971, Battelle Memorial Institute, 505 King Avenue, Columbus, Ohio. Mimeogt ana {Abstracc. } L97 1b. “The origin and development of the bocanical textbook and its American aspects.” Handwritten notes listing titles of textbooks with notations about them. | p., 7 pp. Read 6 March 1971 at Columbus, Ohio. {Manuscript. ] [1975]. “The botanical textbook in nineteenth-century America as a reflection of botanical and cultural crends.” Handwritten, 6 pp. + 1 Table. Titles and vee from co and related information, including items on Asa Gray's Text- books. . Read 11 April 1975 ae Manhattan, Kansas. {Manuscript text and notes. } 1990. Isaac Sprague, “Delineator and Naturalise.” J. Hist. Biol. 23:91— 126. —____—___—. 1996. History of the botanical teaching ie in the U ma States. {Edited by Ronald L. Stuckey & William R. Burk.} Amer. J. Bot. 83:661—171. Stuckey, R.L. and W.R. Burk, (eds.). In press. Emanuel D. Rudolph’ S ane in the his- tory of botany; ... Introductions by Ronald L. Stuckey. Sida, Bot. Misc. Botanical Research Inscticute of Texas, Fort Worth. Torrey, J. and A. Gray, 1838-1844. A flora of North America: containing abridged de- scriptions of all the known indigenous and naturalized plants growing north of Mexico; arranged according to the natural system. Wiley & Putnam, New-York, London; Bossange x Co., Paris. 2 vols. (l:xiv, 711 pp.; 2:504 pp.). Reprinted, 1969. With an introduction by Joseph Ewan. Classica Botanica Americana, Vol. 4. Hafner Publishing Co., New York and London. Wittis, O.R. 1881. Alphonso Wood. Bull. Torrey Bot. Club 8:53—56. Woop, A. 1845. A class-book of botany, designed for colle ges, academies, and other semi- naries where the science is taught. In two parts: Part I. The elements of botanical sci- ence. Part H. The natural orders, illustrated by a flora of the northern United States, particularly New : ngland and New York. Crocker & Brewster, Boston; Simeon Ide, Claremont, N.H. 2 Parts: C 124 pp.; U:ii, 3-474 pp.). Woop, A. 1847. ne ace, pp. 3-5. Jv A class-book of botany ... Illustrated by a flora of the northern, middle, and western states; ... Second edition, eed and enlarged. Simeon Ide, Claremont, N.H. 64 4 pp EMANUEL D. RUDOLPH AS A BOOK REVIEWER FOR CHOICE WILLIAM R. BURK Jobn N. Couch Biology Library University of North Carolina CB#3280 Coker Hall Chapel Hill, NC 27599-3280, U.S.A. ABSTRACT Emanuel D. Rudolph (1927-1992), polar lichenologist and historian of botany, helc academic positions at Wellesley College (1955-1961) and The Ohio State University (1961— 1992). This paper outlines the history of Chorce, a book review serial, and Emanuel Rudolph’s role as a reviewer beginning the year that Chore was established (1964). Important factors influencing his decision to write book reviews were: a commitment to serve colleges, an ou avid interest in book collecting, and sharing a similar philosophy with science historian George Sarton, who stated that those q ualified shoul ; review books “because it was their — ‘ssional duty to do so” (Choice 26:1093. 1989). For 28 publications Rudolph wrote 195 book reviews. Of these 79 ae in Choice in the disciplines of botany (58), polar science ee history of science (6), and other biological sciences (5). His book reviews dem- onstrate analytical and descriptive writing skills as well as a breadth of knowledge in these subject areas. Through his book reviews Rudolph provided collection development and teaching enrichment assistance for college librarians and faculty. ABSTRACT Emanuel D. Rudolph (1927-1992), liquendlogo polar e historiador de botanica, ocup6 cargos académicos en era nae College (1955-1961) y la Universidad Estatal de (1961-1992). Este articulo bosqueja la historia de Choice, una serie de revisiones de libros y el papel de Emanuel Rudolph como revisor r empezando en el aflo en que se 2 establecio Choice (1964). Fueron factores imp jue influenciaron su decision de escribir revisiones de libros: el compromiso de servir a las en su gran interés en la colecci6n de libros, y compartir una filosoffa similar con el cours de la ciencia George Sarton, quien ee que las personas cualificadas deberfan revisar libros “ profesional hacerlo” (Choice 26:1093. 1989) para 28 publi hio porque era su deber . Rudolph escribié 195 revisiones de libros licaciones. De ellas 79 aparecieron en Choice en las disciplinas de botanica (58), ciencia polar (9), historia de la ciencia (6), y otras ciencias ae cae (5). Sus revisiones de libros demuestran unas habilidades analiticas y descriptivas asf como un amplio nocimiento en los temas de estas dreas. Mediante sus revisiones de libros Rudolph promovi6 el desarrollo . la coleccién y el apoyo enriquecedor a la docencia para sus companeros bibliotecarios y la facultad. Biographical accounts of Emanuel D. Rudolph (1927-1992), polar lichenologist and historian of botany, were written by Ronald L. Stuckey (1994, 1995). Additional knowledge of Rudolph’s scholarly endeavors 1s revealed through his book reviewing for Choice, a serial devoted to the pub- Stipa 17(4): 803—808. 1997 804 Sipa 17(A) lication of reviews. The book review as known today is a relatively new form of communica- tion, having emerged and developed after the 16th century. Its beginnings precede the advent of printing when oral criticisms of an individual’s work were made. For example, on the Athenian stage during Aristophanes’ com- edy, The Frogs, a dramatized critique was made of the works of Aeschylus and Euripides. During the late 17th century when newspapers appeared, scholarly notation about books took the form of book notices to acquaint scholars with their colleagues’ publications (Stueart 1978). A book review, an article published in a newspaper or periodical, announces a new book and describes and evaluates it (Donovan 1996). The North American Review, established in 1815, was the first periodical in the United States to provide book reviews. Beginning mid-1963 librarians tn liberal arts colleges throughout the United States received letters explaining reasons for publishing a new re- view journal, CAosce, aimed at undergraduate libraries. They were asked to assist by recommending faculty at their campuses for the purpose of writ- ing reviews. Similar announcements appeared in educational journals. CHOICE, A PERIODICAL PUBLISHING BOOK REVIEWS Although in the planning stages during the late 1950s, Choice did not begin publication unti : — March 1964 under an inicial grant from the Coun- cil on Library Resources through the Ford Foundation (Clark 1964). First housed in Olin Library, Wesleyan University, Middletown, CT, Chovce edi- torial offices subsequently moved to 42 Broad Street, and later to 100 Riverview Center, both in Middletown. Richard K. Gardner was the found- ing editor of Choice, and the “architect” of its reviewing program (Sabosik 1994). Choice, a monthly review periodical, publishes book reviews for the As- sociation of College and Research Libraries, a division of the American Library Association. Although most of the reviews in Choice are for books, electronic products, and microform materials are also covered. These re- views primarily support undergraduate library collection development. Choice reviews are also now available on CD-ROM and online. In 1988 a publishing plan for Choice was approved (Sabosik 1989) and a selection policy was published in 1993 (Anonymous 1993). The editors of Chore give reviewers suggestions to assist them in look- ing at a book. One of the most important services of the reviewer, or sub- ject consultant, is to place a book in the literature of its subject area and to identify its undergraduate audience: freshman, senior, or advanced student. In noting the book’s contents consultants consider (Anonymous [1965 }): (1) What is the standard work with which the book invites comparison?; Burk, Emanuel D. Rudolph 805 (2) What is the relationship of the book to it? Portion of subject covered? Approach?; (3) What is the authority of the author?; (4) On what principle does the author select and order his material?; (5) What is the author's point of view?; (6) How well does he write?; (7) What is the quality of references and illustrative features (bibliography, index, map, illustrations)?, and (8) Is the book a new edition? If so, is the revision significant to war- rant a new edition? The importance and impact of volunteer reviewers were remembered and recognized by the Choice administration. On at least an annual basis the Choice editor communicated news and extended thanks to its reviewers in a memorandum, issued through the early 1970s as “Report to Consult- ants,” and later as a cover letter accompanied by an annual “Reviewer Up- date Form.” Sentiments of gratitude for the contribution of reviewers were expressed on numerous occasions by Choice editors. Especially poignant were the comments of Editor Richard K. Gardner, who stated ina letter dated 9 April 1973 that “your loyalty is much appreciated by the staff of Chose and also by the library community at large. . .. Without your unselfish help Choice could not continue to produce its 6,500—7,000 reviews a year.” Re- viewers were a respected and integral component of the publishing ven- ture. Their comments and opinions were often solicited by Chovce editors if new policies were being considered. When new consultants were needed for reviewing, existing reviewers were asked for their recommendations of colleagues. Besides keeping the books that were reviewed, consultants also enjoyed the intrinsic benefits of reading books in their area of research, and keeping up-to-date in their area of specialty. In 1991 reviewers were of- fered special discounts on subscriptions to Choice. EMANUEL D. RUDOLPH AS A BOOK REVIEWER Rudolph read about Choice and its need for reviewers in the April 1964 issue of Scholarly Books in America. He wrote a letter, dated 1 May 1964, to the editors of Choice, informing them of his keen interest in writing re- views. He mentioned that he had a strong interest in undergraduate librar- ies, having had some experience with them at Wellesley College, where he taught for six years, and at The Ohio State University, where he then was working with a committee to purchase books in the history of science. Although not mentioned in his letter, Rudolph also was an avid book col- lector. He and his wife Ann amassed an impressive, personal library of 53,000 volumes of books in botany and natural history, including an im- portant children’s collection. Twenty-five years later Rudolph wrote about another factor which influenced his interest in writing reviews. He stated that “many years ago, I read a paper by the late George Sarton (1950), founder of the field of the history of science in this country, that urged 806 Sipa 17(4) those qualified to review books because it was their professional duty to do so’ (Clark 1989). Rudolph took Sarton’s comments seriously, as he eventu- ally wrote 195 book reviews for 28 publications. While an assistant profes- sor of Botany at Wellesley College, Massachusetts, Emanuel Rudolph pub- lished his first book review in the January 1959 issue of Ecology (Rudolph 1959). Rudolph wrote his first review for Choice in October 1965 (Rudolph 1965). For Choice he wrote 79 reviews in the disciplines of botany (58), polar science (9), history of science (6), and other biological sciences (5). The language used by Rudolph in his reviews was pointed, but descrip- tive. In citing the weaknesses of a book, he would refer to its “stiff transla- tion, “inaccurate statements,” “dry” style, or “poor bibliography.” In stat- ing the strengths of a book, he would comment that it had “authoritative coverage, “superior, useful illustrations,” “stimulating” text, ora “master- ful summary” of the field. Sometimes Rudolph extolled a book for under- graduates, such as the monograph Fungi: Delight of Curiosity by Harold J. Brodie (1978). About this book, he said, “All libraries . . . should have this volume that could easily spark a future mycologist” (Rudolph 1978). Rudolph’s last review for Choice was published posthumously (Rudolph 19D 2). Besides writing reviews for Choice, Rudolph participated in three other projects: (1) providing bibliographical citations for the revision of Books for College Libraries (Association of College and Research Libraries 1975), (2) submitting entries for Opening Day Collection, a special supplement to Choice (Gardner et al. 1974), and (3) contri suting his comments to the silver anniversary issue of Choice (Clark 1989). Concerning the latter he stated that: — A distinct advantage of writing book reviews is that the books are examined with a critical eye for the audience for whom the review is written. In the case of Choice, my thoughts always are, would this be a book that I would want various kinds of students to find in their libraries? This perspective takes me outside of my own limited professional interests and expands my horizon. The second advantage of Choice reviewing is that it forces me sometimes to read critically books in my discipline that are not exactly in my particular limited field of research. This too has broadened my horizon. In short, being a Choice reviewer has made me a more rounded teacher and researcher, and for that I am very grateful. i Although Emanuel Rudolph’s predominant book reviewing role was with Choice, he also held other related responsibilities. He was the book review editor of The Ohio Journal of Scrence (1965-1974). As editor (1982-1 985) of The Plant Science Bulletin, he oversaw the book review section. Rudolph’s book reviews demonstrate analytical and descriptive writing skills and a breadth of knowledge of books in subject areas of botany, polar science, and history of biology. Through his book reviews Rudolph pro- Burk, Emanuel D. Rudolph 807 vided collection development and teaching enrichment assistance for col- lege librarians and faculty. ACKNOWLEDGMENTS I thank Judith A. Douville, science editor, Choice, for information on Choice and her comments on the paper. Jeff Beam and Karen Collins, h- brary assistants, Couch Biology Library, University of North Carolina at Chapel Hill, provided critical readings and valuable comments. Special gratitude is extended to Ronald L. Stuckey, who provided access to Emanuel D. Rudolph’s archives and reviewed the paper. ARCHIVAL RESOURCES CITED AND CONSULTED Emanuel D. Rudolph’s archives dealing with his association with Chovce are housed in the Herbarium, Museum of Biological pwe, The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212-1192 and curated by Dr. Ronald L. Stuckey, Professor Emeritus and Somter Emeri- tus, Department of Plant Biology, The Ohio State University, Columbus, Ohio. REFERENCES ANonymous. {ca. 1965}. Suggestions for Choice consultants. Choice: Books for college libraries, Olin Library, Wesleyan University, Middletown, Connecticut. 1 p. (mimeo- graphed). _ «1.993. Choice’s selection policy 1993 revision. Choice 30:1577—1583 ASSOCIATION OF Coun GE AND ReEseARCH Liprarirs. 1975. Books for college libraries: A core collection of 40,000 titles. 2d ed. A project of the eee of College and Research Libraries. American Library Association, Chicago. 6 vols. [Volume V., Psychology. Sci- ence. Technology. Bibliography, contains the reference citations recommended by Rudolph] Ciark, V. 1964. Letter to Emanuel D. ae: Department of Botany, The Ohio State University, Columbus, Ohio, 20 May. 2 pp. _. (ed.)}. 1989. Feature. Cibice at 25: Charter reviewers remember. Choice 26:1085-1 097. (Giore, p. 1093). Donovan, D. 1996. Book review, pp. 469-470. In: The World Book Encyclopedia, Vol. 1996 ed. World Book, Inc., Chicago. 22 vols. GARDNER, R.K. and L.F. Lock woop (eds.); K. MCLINTOCK, M.V. Patrucco, K.1]. WERNER, and L.J. Woops (asst. eds.). 197 74. Opening day collection. 3d ed. Choice, Middletown Connecticut. iv, 59 pp. {This is a reprint with corrections and some updating of four rere all tee Opening Day Collection, that first appeared in Choice (Decem- 3, January, February, and March 1974). The science and technology section that ane entries recommended by Rudolph appears in the February 1974 issue. } Rupotpn, E.D. 1959. [Book review: a of as pathology. By E.C. cae and J. George ene 1957. Ronald Press Co.}. Ecology 40:17 1— «1965. [Book review: Guide to the ee of a By Benjamin Daydon Jackson. 1964. Facsimile ed. Hafner}. Choice 2:85. 808 Sipa 17(4) 1978. [Book review: Fungi; delight of curiosity. By Harold Johnston Brodie 1978. pune of Toronto Press}. Choice 15:1395. came eee eeS 2. [Book review: see life at low temperature. By John Davenport. 1992. Chapman & Hall}. Choice 30:3 S{aposik]}, P. E. 1989. Editorial. Choice ae plan. Choice 26:1626. 994. Celebrating 30 years of reviewing: An inside view. Choice 31:1063— 1067. SARTON, George. learned books. Isis 41:14 Stuckey, R.L. 1994. Emanuel (Dada Rudolph (1927-1992): Polar lichenologist and his- torian of botany. Bryologist 97:437—446. {Appendix. Publications and reports of Emanuel D. Rudolph (Arranged chronologically), pp. 443-446, with William R. Burk]. ee, 1999; Emanuel David Rudolph be, Book collector and library friend. Nich, Bot. 34:4—23. STucart, R.D. 1978. Reviews and reviewing. Introduction, pp. 314—324. In: A. Kent, H. Lancour and J.E. Daily (exec. eds.) and W. Nasri (asst. ed.). Encyclopedia of library and 25. Marcel Dekker, Inc., New York, Basel. (35 volumes, pub- 1950. aes preface to volume 41 of [s/s. Notes on the reviewing of 158. information science, Vol. lished 1968-1983; supplements have been issued). FLORA VASCULAR DE LA LAGUNA DE BABICORA, CHIHUAHUA, MEXICO A. EDUARDO ESTRADA C. Facultad de Ctencias Fovestales Universidad Autonoma de Nuevo Leén A.P. 41, Linares, N. L., MEXICO RICHARD SPELLENBERG Department of Biology New Mexico State University Las Cruces, NM 88003-8001, U.S.A. TOUTCHA LEBGUE Facultad de Zootecnia Universidad Auténoma de Chihuahua Cd. Universitaria, A.P. F-28 Chihuahua, Chihuahua, MEXICO RESUMEN La Laguna de Babicora es una cuenca endorreica. Se localiza en la porcion noroeste del estado de Chihuahua enclavada en I ees SS Sierras y Liane ee uras en la Provincia F oo tae de la Sierra Madre Occidental. El clima e La altitud es de 2150 m.s.n.m. en los valles y peer m.s.n.m. en eine picos, con suelos aluviales y Poe respectivemente. Las rocas predominantes son igneas. Se presentan 2 tipos de vegetacién: Bosque de Pino-Encino y Pastizal Hal6fito. La flora esta constituida por aproximadamente 69 familias, 247 géneros : 489 e see pees a los taxa infraespecificos. Las familias mas diveras son Asteraceae, Poaceae y Fabaceae. los géneros con mayor numero de especies son Mahlenbergia, . see Die. ee y oe ABSTRACT La Laguna de Babicora es una cuenca endorreica. Se localiza en la ae noroeste del estado de Chihuahua enclavada en la Subprovincia Fisiogrdfica Sierras y Llanuras Tarahumaras en la Provincia Fisiografica de la Sierra Madre Occidental. El ee es semifrio subhtimedo. La altitud es de 2150 m.s.n.m. en los valles y 2500-2700 m.s.n.m. eeu) con suelos aluv “ales y licosoles respectivamente. Las rocas predominantes son fries Se presentan 2 tipos de vegetacién: Bosque de Pino-Encino y Pastizal Hal6fito. La flora esta pee por aproximadamente 69 familias, 247 ee y 489 oF ademas de los taxa infraespeciticos. Las familias mds diversas son Asteraceae, Poaceae y Fa baceae. los géneros con mayor numero de especies son ee oie US, Enphorbia, Dalea y Desmodium. Sipa 17(4): 809-827. 1997 SLO Stipa 17(4) INTRODUCCION La Laguna de Babfcora pertenece a las dreas denominadas humedales (INEGI 1990). Es una cuenca endorreica con escurrimientos de agua semipermanentes. Se localiza al noroeste de la ciudad de Chihuahua, en los municipios de G6mez Farfas y Madera en la Provincia Fisiogrdfica de la Sierra Madre Occidental en la Subprovincia Fisiografica Sierras y Llanuras Tarahumaras (INEGI 1987). A la fecha no se presentan estudios floristicos de la zona ni de areas aledafias por lo que el presente trabajo tiene el propésito de contribuir al conocimiento de la flora de la Sierra Madre Occidental. La cuenca de la Laguna de Babjcora se localiza entre las coordenadas 29°05'- 29°40°N y 107°30'-108°15 W. Abarca una superficie aproximada de 1882 km cuadrados. Su elevacién va desde los 2150 m en los valles hasta los 2700 m en los picos montafiosos. La zona de estudio tiene forma semicircu- lar y queda circundada por las vias de comunicacién que unen los poblados El Alamillo, Pablo Amaya, El Porvenir, Gomez Farias, Pefia Blanca, San José de Babicora y La Libertad (Fig. 1). La f6rmula climdtica del area presentada por INEGI (1986) es C (E)(w1)(b\(e’), semifrio subhtimedo, con Iluvias en verano de tipo torrencial, verano fresco largo, y oscilaciones de temperatura media mensual muy altas. La precipitaci6n promedio anual es de 630 mm. La temperatura media anual es de 11.5° C. Las rocas dominantes en la zona de estudio son de tipo igneo extrusivo del Periodo Terciario, Los suelos dominantes son race y lacustres, estos ultimos ricos en materia organica, de color negro obscuro a café pardo con profundidades de hasta 2 m (Montufar 1987, INEGI 1986). La Laguna presenta un ojo de agua permanente que mana del subsuelo. Presenta escurrimientos de agua semipermanentes de arroyos que fluyen hacia e — interior de la cuenca. VEGETACION La vegetacién de la La Laguna de Babfcora queda clasificada dentro de dos tipos principales: Bosque de Pino—Encino y Pastizal Hal6fito (INEGI 1986) Bosque de Pino-Encino.—Es la comunidad vegetal dominance en la zona, abarcando una extensién de 1075 kil6metros cuadrados, presentandose en todos los picos montafosos que rodean a la laguna. Entre las especies dominantes de pinos estén Pinus cembroides, P. engelmanii, y P. chibuabnana: los encinos mas frecuentes son Quercus emoryi, O. arizonica, Q. depressipes, QO. hypolencoides y QO. rugosa. Pastizal Halofito.—Constituye la vegetaci6n predominance en las partes bajas o Ilanuras, abarcando una superficie aproximada de 237 km cuadrados, alcanzando una altura promedio de 30 cm, donde las especies dominantes son Dastichlis spicata var. stricta, Mublenbergia utilis, Sporobolus eryptandrus, Estraba C., ET AL., Flora de la Laguna de Babjicora 811 \ 08°00! Cuenca de la Laguna de Babicora Afio de Hhdalgo A Aguaje L Pint a Pinta SIs DE ihe GEOGRAF FICA oe NAT oe EOD EESMIA (U ACH.) 29°22'- TOUTCHALFEFRG Carr. & a ee INEGI, 1983 ay ] . Localizacion del area de estudio, escala 1: SOO,000 Fic. | Eragrostis mexicana, Cyperus odoratus, C. sphaerolepis, Eleocharis tenuis y Physa- lis sulphurea. FLORA Los registros de plantas vasculares de la Laguna d e Babicora comprenden 489 especies y taxa infraespecificos, englobadas en 247 an y 69 familias (Tabla 1). Las familias mas diversas se presentan en la Tabla 2. Los géneros con mayor numero de especies se presentan en la Tabla 3. on una especie es endémica en la zona de estudio (Hydropectis estradii B.L. Turner). Todas los registros de especies se circunscriben exclusivamente a las colectas realizadas por los autores (Mayo 1994—Agosto 1995). El arreglo seguido para el listado de plantas en los grupos taxonoOmicos mayores para angiospermas es el propuesto por Cronquist (1988). Se sigue una secuencia alfabética en la disposicién de familias géneros y especies. ABREVIACION EN LA LISTA FLORISTICA Colector (EE= Eduardo Estrada) y herbario donde estan depositados las colecciones son BRIT, NMC, NY, TEX (Holmgren & Holmgren 1993) 812 Sipa 17(4) TasLa |. Flora vascular de la Laguna de Babscora. Division Familias Generos Especies Polypodiophyca \ 7 LO Pinophyta 2 3 7 Magnoliophyta Clase Magnoliopsida 34 189 3533 Clase Liliopsida 9 18 119 TOTAL 69 247 189 TABLA 2. Familias con mayor numero de géneros. Familia Generos Especies Asteraceae 48 80 Poaceae 28 79 Fabaceae i) 18 Cyperaceae { 16 Scrophulariaceae 7 13 Solanaceae i 13 Euphorbiaceae { 13 TABLA 3. Géneros con mayor ntimero de especies. Genero Especies Genero Especies Muhtlenbergta 15 Dalea 9 Cyperus 10 Desmodium 8 Euphorbia 10 Aristida 8 NOTA: Los numeros de colecta en su totalidad y los que no estan precedidos por abreviaturas de herbario, son ejemplares que se encuentran depositados en el herbario de la Facultad de Ciencias Forestales, Universidad Auténoma de Nuevo Leon. EstrRADA C., ET AL., Flora de la Laguna de Babicora LISTA DE LAS PLANTAS VASCULARES DE LA LAGUNA DE BABICORA POLYPODIOPHYTA ADIANTHACEAE Se sinuata 1 Hook (Lag. ex Swartz) am Benha Boemmeria pe ee Undew., EE 3041 NM Cheilanthe alabamensis (Buckl.) Kunze, EE O5 C; EE 3018 Che aie villosa Maxon., EE 3011 NMC; EE 3418 NMC. Cheilanthes wootont Maxon, EE 3040 NMC. Cheilanthes wrightit Hook., EE 3676 NMC. Pellaea atropurpurea (L.) Link, EE 3¢ 74 NMC EQUISETACEAE Equisetum laevigatum A. Braun, EE 2945 NMC, NY OPHIOGLOSSACEAE Ophioglosum engelmannii Prantl., EE 3504A POLY PODIACEAE Pleopeltis polylepis (Kunze) T. Moore var. erythrolepis (Weatherby) T. Went, EE 3704 NMC, PINOPHYTA (Gimnospermas) CUPRESSACEAE Steud, EE 2489, 3394 NY. lynihveyus debh, efor L ~ de PINACEAE Pinus arizonica Engelm Pinus ayatabuite Ehr. var. ae Shaw, 3221a. Pinus embod: Zucc., EE 345 2a. Pinus chihuahuana Engelm. EE 2491. , EE 3657. Pinus engelmanit Carr. Psendotsuga menziesit (Mirb.) Franco, EE 36 MAGNOLIOPHYTA (Angiospermas) MAGNOLIOPSIDA (Dicotiled6neas) AMARANTHACEAE Amaranthus palmert 8. Wats., EE 3272 NMC; EE 3515 BRIT, NMC, NY ; EE 3482 BRIT, NMC, NY; EE 3342 NMC; EE ag NMC; EE 3195 NMC; EE 3751 NM oe gracilis (Hook.) Moq,. EE 3113 NY; EE 3127 NMC. Gomphrena sonorae Torr., EE 3341 NMC, NY. Guilleminia densa (Willd. ) Mog., EE 3486 N MC Gossypianthus Tinie (Poir.) Mog. var. lanuginosus , 333) , NY. Tidestroemta lanuginosa (Nutt.) Standl. var. lanuginosa, EE 3391 NMC, NY. tH S ANACARDIACEAE Rhus trilobata Nutt. ex Torr. & A. Gray var. pilosisstma Engler in DC., EE 3020, EE 3361 NMC APIACEAE Daucus montana Humb. & Bonpl., EE 3742 [C. ie lemmonti Coulc. & Rose, EE 3016 Eryngium —— Hlum Engelm., 3396 NMC ; EE 3290 BRIT, pe EE 3134. Prinosciadinum madrense 8. Wats., EE 3058 NMC ASCLEPIADACEAE Asclepias contrayerba Sesse & Moc., EE 3140 N Ades ‘ie: illata (A. Gray) Vail, EE RIT, NMC, NY. ae chibuahuensis A. Gray EE 3206a NMC. ASTERACEAE sae eee L. EE 7 TEX ore nee a (A. Gray) Greene, EE 370 1C, NY, TEX 3338 NMC, NY, A ies ne eabina (A. Gray) King & Rob., NMC, NY. 814 Ageratina oe (Gray) King & Rob., El 3705 7 Ambrosia Camescens - Gray, EE 3292 BRIT, Ny LES Ambrosta Pusch DC., EEB-S627a TEX: EE 33 EX; EE 3482a TEX: EE 3484; ee ete EE 3267a; EE 3779 TEX. Artemisia dracunculus var. glauca (Pall.) Munz, EE 3672 NMC, NY, TEX. Artemivia ludoviciana Nutt., EE 3095 TEX: 7TI2NMC, NY, TEX. Ages ae var. parviflorus (Nees) ee ae 32069 NMC, NY, TEX; EE > EE 2920 NMC, NY, TEX. re auved ate Sherff ~ LE. 3068, 32284 TEX; EE eran EE 3518a; EE 353- Bidens lemmonii A ce 3E 3592 N ~ TEX. — epln tha Q} Sherff, EE 3669 NY, TEX. Bidew morte Cav. var. odorata Gray, EE TEX; EE 3258 TEX; EE 3259: EE a 1 NMC, NY, TEX; EE 3375 TEX; EE 3399 TEX. — a Gray, EE 3560 BRIT, NMC, NY, TEX Brille ones var. choi olepis (Woot. Shinners, EE 3620 TEX; EE EX; EE 3620 NMC, NY, TEX. Bal ne Gray, EE 3692 BRIT, NMC, NY, TEX. Brill snplex A. Gray, EE 3563 NMC, TEX Bide; a ley & Ss hoe en ere, (S. Wats.) Grashoff ex. B, Turne er var. privgler EE 3470 NMC, Y," oe americana Nutt., EE 3678 BRIT, NM IC , TEX iH) eo ent ae Ce, se a BRIT, “NMC. NY, TEX; EE Conyza idpuda (L.) - ronq., a 3313 LEE EE S523 LEX Cosmos « parciforn (Jacq.) Pers., , EE 3256 TEX; EE 3260 | TEX 607a TEX: EE 3616 TEX. Cosmos palmeri B.L. Rob., EE 3444 Cosmos pringle: BLL. Rob. & Fern., LEX, EE 3253 ) TEX; EE 1 TEX. , EE 3427 NMC, TEX; EE EE 3573 Stpa 17(4) Dahlia coccinea Cav., EE 3015 TEX. LE rigeron COVONAVIMS Green EE 3326 BRIT, TEX; 3406 TEX; EE 3457 Evigeron fraternus E. Greene, EE 2942 TEX. Erigeron neomexicanus Gray, EE 3038 TEX. Es rigeron oreophilus Greenm., EE 3625 NMC, Erigeron strigulosus Greene, EE 3789 TEX; BE 3790 THX; BE 3700 NY, TEX. Evigeron versicolor (Greenm.) Sac e i E 3203 NMC, TEX; EE TE 5 34 TEX; EE 3498 TEX. Evrigeron ae Gray) Greene, EE 343 16) TEX; EE ae ie BRIT, TEX, NMC; EE 3373 oT Galo paid Torr., EE 3122 BRIT, yee lll Cav. var. see EE 3312 EX; EE 347 oe hii Cav. var. ae A. , EE 3640 TEX, Ghali aff _arizonicum A, Gray, EE 3002 , EE 3190 NMC, NY, TEX; EE 3 a NMC, j Guaphalinm viscosum Kunth EE 3129 NMC, NY, TEX; EE 3327 TEX; EE 3404 TEX. Guapbalium sp. “EE. 3708 TEX. Grindelia cn Steyerm., EE 3316 TEX. a (A. Gray) B.L. Rob., EE 3078 NY, aoe EE 3756 ve TEX. Gutierrezia eae Nera ) M. Lane, EE 3257 NMC, NY,” Gutierrezia rari A. EE 3595 TEX: EE 5S NMC, TEX. pee aerate Bierner, EE 3287 NY, TEX; EE 3501 NY, TEX. Heterosperima pinnatum Cav., EE 3066, 324 TEX; EE 3476 NMC, NY, TEX. 3609 TEX; Guardiola ar git Hreracium carnenm Greene, EE SE 3572 TEX; S571 T Hreracium pringler A. Gray EE 3164 TEX. Hydropectis estradti B.L. Turner, EE 3499 Hyme ope pit mexteanus Gray, EE 3709 TEX. /; Gray, EE 3693 BRIT, EE 3517 NY: EE 361 Lactuca serriola LL. TEX ESTRADA C., ET AL., Laennecia schiediana (Less.) Nesom (= Conyza schiedeanus (Less.) Cronq.), EE 3645 NM EX; EE 3138 TEX. Laennecia sophiaefolta Boras ae (= 5 Conyza ais Kunth)., 3179 NY, TEX: EE 3247 NMC, - ae EE 2) 313 38 a Lasianthaea po piipialelt Gray) Benth., EE 3056 T Leibnitzia yaa Don) Nesom, EE 2939 T E 3659 NMC, TEX eset gracilis (Nutt. Shinn rs, EE 3405A TEX; EE 3353 NMC, NY, TEX. 343 TEX: ao SEVIQOSUM Stuessy, E3 ie EE 3464 TEX. Pectis ae Cav., EE 3090 T Psilactis astertoides A, Gray, ae 42 i NY; 3524 TEX. — mexicana (Wats.) Sharp, EE 3681 ie pinnata (Lam.) Kuntze var. cuatemalensis (Rydb.) McVaugh, EE 3281 BRIT, NMC, NY, TEX; EE 3449a TEX; EE 3477 NMC, NY, TEX; EE 3534 NMC, NY, TEX; EE 3539; EE 3540 TEX; EE 3643 TEX. Senecio flaccidus Less. var. flaccidus, EE 3365 NMC, NY, TEX. Senecio octobracteatus Barkley & Turner, EE 2927 Simista amplexicaults (Sims) Pers., EE 3389 NMC, NY, TEX; EE 3459 TEX; EE S511 NMC, NY, TEX; EE 3516 NMC, , TED Stevic “planer i) var. plumerae, EE 7 TEX; EE 3575 TE Stevia serrata C av. var. serrata, EE 3055 ce EE 3240 NY, TEX; EE 3215 TEX.; EE 2793 TEX Stevia viscida Rundi: EE 3469 NMC, NY, EE 3557 NMC; EE 3791 TEX. EE 3135 T TE Tavetes lncida Cav., 5 TEX. EE 3284 BRIT, Tagetes i er Cav., NM , TED a9, a Be Ge ray, EE 3538 NMC, NY, 39 TEX ee io: L., EE 3355a NY. seks ee, ace .) Cass., EE 3445 > EE 3504 TEX; EE 3525 TEX. Flora de la Laguna de Babjscora Tragopogon porrifolius L 815 ., EE 3283 TEX; EE me a Verbesina longifolia Gray, EE 3446 NMC, TEX; EE 3673 NMC, NY, 7 pee aiansmaae B.K.) Blake, EE 3079 a Bcp a — a a Gray var. cordifolia, EE RIT, NMC, NY, TEX. re es s A. Gray, EE 3085 TEX. Vigwiera nario (Nutt.) Blake, EE 3282 E 3315 TEX. Nanthinm streumarium L., EE 3528 NMC, Nantl ee lum &) spermotdes (A. Gray) Benth., EE 3295 NMC, NY, TEX Zinnia tenuis 28 Wats.) Strother, EE 3480 ’ TEX; EE 3686 TEX. BORAGINACEAE Amsinckia micrantha Sask., EE 3362 NMC, Heliotrapinn curassavicum L. var. curassavicnm, EE 3231 NMC, NY. EE 3/28 BRIT, NMC, NY Lithospermum quleaiovain Gray, EE 3249 Lithospermum cobrense Greene, EE 3249a NMC BRASSICACEAE Brassica campestris L., EE 3271 BRIT, NMC, NY. Brassica rapa L., EE 3483 NMC. Descurainia impatiens (Cham. & Schlecht.) schulz. EE. 3792 TEX Jescurainia pinnata (Walt.) Brit. var. prnnata, EE 2492. Draba petrophylla Greene, EE 3671 NMC, NY. Lepidium sordidum L., EE 3289 NMC. Lepidium virginicum L., EE 3685 NMC. Penellia longifolia Benth., EE 2999, 3168 NMC, NY; EE 3670 NMC; EE 3766 NMC. nase micrantha (Gray) Nieuwl., EE 3325 NW EE Sisy oe tum linearifolium Nutt., 3458 MC EE 3360. Sisymbrinn sp., 816 CACTACEAE Echinocereus polyacanthus Engelm., EE. 4241. Mammillaria sp. EE. 4244. — CAMPANULACEAE Lobelia anatina ¥.. Wimm, EE 3460 cae IT, TEX; EE 3762 NMC; 3621 BRIT, NMC, NY. Lobelia fenestralis Cav., EE 3139 NMC; EE 3296 NMC; EE 3403 TEX. CARYOPHYLLACEAE Arenaria bourgaer Hemsl., EE 3503 BRIT, MC, NY. Arenaria Ree Gas (Michx.) Rohrb., EE 3104 IC, NY. ee nutans is Raf. , EE 2948 BRIT, NMC, N 2 BRIT, NMC, NY, eon C orvigiola | pe Tr. & Planch., EE NMC, NY. Drymaria leptophyHa (Cham. & Schlecht.) fenzl., EE 3145 BRIT. ae a EE 3212 NMC: EE 3711 NMC, Drymaria molluginea ( Lag.) Didr. NMC, NY. Silene laciniata Cav. var. daciniata, EE NY 51 BRIT, NMC, TEX. var. pringle: (Wats.) 4 BRIT, NMC, NY. 3157 DO4T pi ae Hook. tche., EE 357 CHENOPODIACEAE Atriplex texana 8S. Wats., MC, TEX. EE 3730 BRIT, Chenopodium album L., EE 3123 BRIT, NMC, NY; EE 3270 NY) EE 3522 NMC, pac ae Vala Mogq., EE 3726 NMC, NY. Chenin graveolens Willd., EE 3628 NY; ae i 3/ Chenopodium incisum Poir., BE 3124 NMC, NY. Shale tragus L., EE 3392 NY. CISTACEAE g ) Dun., EE Helianthemum vlomeratum (Lag. C,NY. 3378 NM Helianthemum ae S. Wats., E 3380 NMC Sipa 17(4) CONVOLVULACEAE Convolvulus equitans Benth., EE 3328 NMC, NY. Evolvulus sericens Sw. EE 3329 BRIT, NMC, NY imo apace (H.B.K.) G. Don, EE 3317 ea a mm ee Torr. , EE 3161 NMC, NY; EE 3200 NN Y, Tbamnen lindbetmiert Gray, EE 3569 BRIT, NMC, NY. |pomoea ineainale Jacq. ie 35360 NMC, NY. Ipomoea madrensis 8. Wats., EE 3072; EE 3150 NMC [pomoea pubescens Lam., EE 3351 NY. Ipomoea tenutloba Torr., EE 3032 NMC. CRASSULACEAE Sedum alamosanum M.E. Jones, EE 3042 CUCURBITACEAE Sicyos lig L., EE 3725 BRIT, >; EE 3236 NMC. Cucur en i ale NMC, (obs. pers.). CUSCUTACEAE Cuscuta applanata Engelm., EE 3547, 3126 ERICACEAE Arbutus xalapensis H.B.K., 2496. Arctostaphyllos pungens H.B.K., EE 3152 NY. Monotropa latisquama (Rydb.) Hult., EE iE 3059; EE EUPHORBIACEAE Acalypha neomexicana Muell., EE 3172 1) GE 3209 NIMC, NY:-EE 3519 NY. Chamesey se e serphyllifolia (Pers.) Small, I'T, NMC; EE 3530 NMC TEX: EE 3028. Eaphitinva barrancana Mayfield & Steinm., 3440 NY, TEX. Exphorbia inane Boiss., EE 3387 NMC. Euphorbia dentata Michx., EE 3242 BRIT, NMC, NY. NY, 49 ESTRADA C., ET AL., Euphorbia heterophylla L., EE 3776 NMC. Euphorbia hexagona Nuce., EE 3596. ae hia eis o & Gke.) Boiss., EE 4N Pes nutans a EE 3183 NMC. oo sphaerorhiza Benth., EE 3320 , NMC, NY; EE 3596a NMC. Er . vil Scheele, EE 2925 NMC, MC. a < ne Kunth, EE 3321 TEX, Tragia ramosa Torr., EE 3337 NY. FABACEAE FE 2229. NY. EE NMC, Astragus ig Wats., 3428 NMC, NY; 3526 BRIT, NY. A stragatus Aumistratus Gray var. humistratus, 5 , NMC, NY. Astragalus longissimus (M.E. Jones) Barneby, 3433 NY. As iragalus scalaris Wats., EE 2929 NMC, NY. es vaccarum Gray, EE 2928 NMC, E 3224 BRIT, NMC, NY. Colagania a ae ELB Bs, BE 3057 TE 1 NY; EE 3376, 3408 NY. Cologania aren Schlecht., EE 3049, 3644 NY; EE 3767 NY; EE 3069, 3442. Crotalaria Nes Riley, EE 3050; EE 3606 NY; EE 3757 NY; ae e: | ie NY, TEX Crotalaria rotundifolia (Walt.) Gmelin var. vulgaris Grear, EE 3160 NY; EE 3412. Coursetia cies (Gray) Lavin, EE 3082 NY; E 3177 NY; EE 3211 NY; EE 3454 ek , NY. Datea albiflora Gray, EE 3128 NMC, NY; EE 3582 NMC, NY; EE 3622 BRIT, NMC, NY, TEX; EE 3733 NMC, NY. Datlea exigua Barneby, EE 3154 NY; EE 3201 NMC, NY; EE 3584 BRIT, NMC, 3760 NY. Datlea filiformis Gray, EE 3186 NY; EE 3206 T, NMC, NY; EE 3479 NMC, NY, BRIT; EE — NMC, NY; EE 3736 BRIT, N Datea ee (Vail) L. o Williams, EE 3044; | BRIT, NMC, NY; EE 3574 NY. Dalz fe (Aiton) Bullock, EE 3535 Flora de la Laguna de Babjscora 817 NMC, NY; EE 3778 NY; EE 3453 NY. — ihe SS sie EE 3613 a prs ee a ie 3] NY: EE 3216 NY; EE 3694 NY; EE 3740 NY; EE 3777 NY. aes fee Cov ) Willd., EE 3614 BRIT, E 3735 BRIT, NMC, NY. _ msi ae Greene var. wrceolata, EE 3615 NY. Desmodium arizonicum §. Wats., EE 3710 NMC, NY. Desmodium batocanlon Gray, EE 3025 TEX. Desmodium gees Gray, EE 3012; EE 43 TEX; EE 3196 NMC, NY, TEX; EE 3390 NY, sae EE 3597 NY; EE 3715 TEX Desmodium hartwegtanum Hemsl. var. hartwegianum, EE 3045, EE 3153 NY; Desmodium madrense Hems|., EE 3455 NMC, Desmodium neomexicanum Gray, EE 3424 RIT, NMC, NY, TEX; EE 3115 NMC, NY, TEX; EE 3254 NY; EE 3388 NY, TEX; EE 3688 NMC, NY; EE 3753 NMC, NY; EE 3773 BRIT, NMC, NY, TEX fio PE 3390 BRIT, it Desmodinm retinens Schlech NMC, NY, phere sp., EE 3¢ 423 NY,TEX; EE 3054 EE 3021 TEX. ae ‘a montana Rose, EE 4243. Lathyrus graminifolius se White, EE 3562 BRIT, NMC, Lotus chihuahuanus oo a EE 3071. Lotus oroboides (H.B.K.) Ottley ex Kearney & Se es, EE 2935 NY; EE 3425 NY; EE 3 Lupinus bark C.P. Smith, EE 3063 TEX; EE 3262 NY,TEX; EE 3422 NMC,TEX; oye’ re NY,TEX; EE 3145 BRIT, NMC, NY, TEX. a he ae nn i C.P. Smith, EE 3184 EX; EB 3209 BRIT, NMC, NY, = 3441 BRIT, NMC, NY, TEX. Macroptilium gibossifolium (Ort.) Delgado, EE : NY: EE 334 Medicago sativa L., EE 3353. Medicago lupulina L. (observ. pers.). S18 Minkelersia galactotdes Mart. & Gal., EE 3608 BRIT, NMC, NY, TEX. Phaseolus parvulus Greene, 210; EE 3212a BRIT, NMC, NY, EE 2856 TEX Phaseolus pedicelatus Benth. var. gr —- Woot. & Standl., EE 3383 NY, SEE ; EE 3064. EE 3051; EE TEX; 3447 TEX; Phaseolus ritensis Jones, EE 2852 NY. Phaseolus vulgaris L.., EE 3264 NY, yachosta macrocarpa Benth., EE 3359 BRE I, NMC, Sophora ee Tu ner, EE 3 oe Trifolium amabile WBAK. var. onabile 3602 NY; EE 3223. Trifolium dad H.B.K. var. Br Cee Hemsl., EE 3331 NY; EE NY, BRIT; EE 3495 NMC, EE 3738 BRIT, NMC, NY. Trifolium wormskioldii Lahm. var. ortegae (Greene) Barneby, EE 3314 BRIT, NY. Viera pulchella W. B.K. ssp. mexicana (Hemsl.) &. Gunn, EE 3500 pees NY, TEX; EE 3550 BRIT, NMC, sea os K. ve pulchell £ E 2936 a YDS ie ere es oa 31 Zornia reticulata Sm., = 3453 3 BRIT, NMC, FAGACEAE Quercus artzonica Sarg., EE 3024; EE 3439 BRIT, NMC, NY. Onercus Dae Trel., EE 3702 BRIT, NMC, NY; EE 3716 BRIT, ne NY; EE 3720 NMC, NY. Quercus emory? Torr, EE 3471 BRIT, NMC, Fr Quercus gambelti Nutt., EE 2934 NMC, NY: EE 3687 BRIT, NMC, NY. » EE 3472 BRIT, NMC, 3701 BRIT, Onercus gra : iebm., OO NN AC; EE Quercus hypolencordes A. Camus, EE 3023; EE 3474 NMC see nevaughit Sual lenberg, EE 3237 C; EE 3438 BRIT, NMC, NY: EE NW aes NMC, NY, BRIT; EE 3627 NMC, NY. Sipa 17(4) Ouercis rugosa Nee, EE 2490; EE 3626 NMC. Quercus sideroxyla Humb. & Bonpl., EE 3088 BRIT, NMC GENTIANACEAE ae amarella (L.) Boerner, EE 3677 NMC, NY, TEX. GERANIACEAE . Her, EE 2494; EE i moet cicntarinm (L.) L Geranium atropurpurenm Heller, EE 3244 NMC, NY; EE 3381 NMC: EE 3684 NMC, EE 3656 BRIT, NMC, NY. Geranimm nivenin S, Wats., EE 3771 NMC: EE 3619 NY Geranium richardsonit James, EE 3787 NMC, Geranium vubulaton gaged Kunth, 3679 NMC, NY; i NMC EE 3783 NMC. Geranium wislizeni ee EE 3006, EE 3099 > EE 3220 NMC, NY GUTIFERAE Hypericum pauctfloram H.B.K., NMC, NY HY DRANGEACEAE Philadelphus oo A. Gray, EE BRIT, NM fe E 3662 HYDROPHYLLACEAE — fichi WOW I (R uIzZ& Pavon) Choisy, EE IS NY, T LAMIACEAE A gastache aan (Lindl.) Cory var. cortacea - . Sanders, EE 3091 BRIT, NMC 3 a BRIT, NY. ae oblongifolium eee eller var. mexican Irving, EE 3 ae NMC, Y; EE 3092 NMC, — Monarda austromontana Epling, EE 3255 ’; EE 3084 city. Salvia prunelloides H.B.K., EE 3166 NMC, NY EstRADA C., ET AL., Salvia reflexa Hornem, EE 37 NMC, NY, TEX. Salvia tiliaefolia Vahl., NMC, NY, TEX; EE 3426 NMC, EE 3213 BRIT, NMC NY. Salvia sp., Stachys coccinea Jacq., EE 3576 BRIT, NMC, ne LINACEAE Linum aristatum Engelm., EE 3158 NMC. Linum sp., EE 3600 NMC MALVACEAE EE 3230; EE ) BRIT, NMC, Anoda 1 cristata (L ) a ENG 351 NY; EE 3 387 NMC Matlva ae Waller. EE 32 78 NY; EE 3 BRIT, NM Sida neomexicana Gray, EE 3411 NMC. Sidalcea neomexicana A. Gray, EE 3291 BRIT, M > EE 3610 Glaeiahes angustifolia ( (Cav.) Don var. angustifolia, EE 3137 BRIT, NMC, NY. Sphaeralcea fendleri Gray, EE 3529 NMC; MIMOSACEAE Acacia angustissima (P. Miller) Kuntze var. texensts (Torr. & Gray) Isely, EE 3363 NY. Calliandra humilis Benth., EE 3463 BRIT, NMC, NY; EE 3623. — saree: Gray, EE 3007, EE re dysocarpa es cs: 3368 BRIT, NMC, N NYCTAGINACEAE Mirabilis longiflora L., EE 3033 NMC. eee multiflora oo Gray, EE 3117 \ NM rAi5) Minis Roce Heimerl, EE 3169 BR MC, NY; EE 3698 NMC. OLEACEAE 7 NMC, NY. Fraxinus velutina Torr., EE 3437 ONAGRACEAE Epilobium ciliatum Rat.. EE 3274 BRIT, NMC, NY. Flora de la Laguna de Babicora 32 BRIT, EE 3731 BRIT, 819 Cane coccinea Pursh., EE 3047 TEX; EE ; EE 3448 TEX; EE 3588 NY, TEX; EE 3005 THX: 3507 TES: Gaura hexandra Gomez Otrega ssp. gracilis oc. & Standl.) Raven & Gregory, EE 3136 NMC, NY,TEX. Lopezta gracilis Wats., EE 3637 NMC, NY. Ocnothera brachycarpa Gray, EE 3497 TEX. Oenothera coe es ssp. laciniata, EE 94 NMC 3553 NMC; EE 3703 NMC; EE 3 on EE 3449 NY, TEX. OX ALIDACEAE Oxalis alpina ( egy EE 32 Oxalis corniculata I 1 NY; EE 365 NMC, NY; EE ee ty ace — Bk Be a2l7 IMG. i aaa Gray, EE 3268 NMC. PLANTAGINACEAE Plantago australis Lam., EE 2922 NMC. Plantago hirtella H.B.K., EE 3630 NMC, NY. Plantago linearis H.B.K. var. mexicana (Link) Pilger, EE 3246 BRIT, NMC, NY, TEX. Plantago patagonica Jacq., EE 3456 NMC. POLEMONIACEAE a a pinnata (Cav.) V. Grant, EE 3793 ; EE 3377 NMC; EE 3719 NMC, Ipomopsis thurberi (Gray) V. Grant, EE 3432 BRIT, NMC, NY. EE one E 3346 NMC. [pomopsis sp., Phlox nana Nutt., POLYGALACEAE — rl ee A. Gray, EE 3027; EE 3180 3605 NMC, NY. my a Nutty hE 2519: ER 3132NYX, Polygala es Benth., EE 3587 NMC; EE 3593 NMC, NY. Polygala tu ee Britt., EE 3364 NMC, NY. POLYGONACEAE 3992 Eriogonum atrorubens Engelm., EE C 820 Polygonum aviculare L., EE 3277 BRIT, NMC, NY; EE 3531 BRIT, NMC, NY. Polygonum ia al L., EE 3307a BRIT, NMC Pabysonan ~ an anicum L., EE 3548 NY. Polygonum sp., EE 3140a. E 3646 ey we Rumex mexicans Meiens. E 7 NMC, NY; EE 3280 NMC; EE ee NMC. BE 365 Rumex acetosella L. Rumex sp., SONY. PORTULACACEAE Portulaca oleracea L., RE 3273 NMC. RANUNCULACEAE Agquilegra skinnert Hook., EE 3674 BRIT, NMC, NY, TEX. Ml andesit ola Ewan, EE 2943 NMC. iC: Ranunculus sp., EE 3617 NI alah pinnatum Wats., EE 3682; EE 3382 NMC RHAMNACEAE Ceanothus fendleri Gray var. venosus Trel., Ek 3000; EE 3060; EE 3093a NMC, NY iE 3 x 372 ROSACEAE Alchemilla sii L., EE 3304 BRIT, NMC, NY oa montanns Raf. var. as (Wats.) EL. Martin, EE 3022. pct ea Via Maxis EE 3691 NY, TEX. Pail ararny L. 3227 BRIT, NMC, 3521 BRIT. NY. as noe iis EE 3461 NMC. Potentilla s sp. 495: EE 2518. Prunus serotina aes EE.4242. Rosa sp., EE 2932 NY, NMC. RUBIACEAE eens ternifolia (Cav.) Schlecht., EE 305 Crusea ae (H.B.K.) Anderson, EE 34066 NY Crusea ci (Willd. ex Roem & Shult.) Anderson, EE 3478 BRIT, NMC, NY. Galinm mexicanum H.B.K. var. asperulum Stipa L7(A4) (Gray) Dempst., 3668 BRIT, NM Galiun microphyHum ee E E 300 onsis L., EE 3108 NY; - 3208 3318 NMC, NY; EE 3400 NY; EE ee NY. RE — NMC, NY; EE IC, he a arve SALICACEAE Populus tremuloides Michx., EE 2933 NMC, NY; EE 3658 NMC, NY. Salix lastolepis Benth. (obs. pers.). SAXIFRAGACEAE Heuchera sanguinea Engelm., EE 3030 SCROPHULARIACEAE soe patriotica Fern., EE 2941 NMC, ’; EE 3683 TEX 7 aw. meee tenuiflora ae on 3717 BRIT, MC, NY. 4 TEX. eo sp., BE 3080 sara Mecardine ieee tees H.B.K.) Pennell, EE 16 BRIT, , NY. Mimulus gutatus ¥ oy ex DC. EE 3772a1 , EE 2924 NY, Penstemon barbatus Ke av.) Roth., 3076 aes N EE 3188 NMC, fee EE | NMC. Pee campanitlatus (Cav.) Willd. ssp. i hibuahuensis Straw, EE 307 : 7 +3165 NMC; EE 3250 oe NMC, NY 5 Penstemon lanceolatus Benth., EE 3251 NMC, Penstemon mintata Lindl. ssp. apateticus G oy) Straw, EE 3419 BRIT, NMC, a stenophyllus Gray, EE 3010; EE 5 2 3603 NMC a intermedta (A. Gray) Pennell, EE 3112 NMC. Seymeria a Seem., EE 3062; EE 74 BRIT, NMC, NY. . Wats., EE 2940 NMC, Veronica mexicana § TEX. SOLANACEAE Datura quercifolia A.B.K., EE 3125 NMC, NY. EsTRADA C., ET AL., ae parviflora Juss., EE 3228 BRIT, , NY. ae soudella Standl., EE 3075 NMC, NY Phy sais hederaefolia Gray var. hederaefolia, EE 3355; EE 3204 NMC, NY, TEX Phy satis Leetela Gray var. puberitla Gray, EE 3402 NMC. Physalis pubescens L., EE 3 Physalis pubescens L. var. oe EE 3171 sop scene Rydb. var. svbulata, EE T, NMC, NY. Phys sats phi la Waterfall, EE 352 RIT, NMC, NY; EE oe Ea 3230 NY. ea SP -~ sae NMC, NY. n A. Br., EE 3266 BRIT NMC, NY. Solanum fendleri A. Gray, EE 3061; EE 3649 05 NMC; EE 3218 NMC, EE 3386. Solanum nigrescens Mart. & Gal., EE 3001 EE 3191 BRIT, ; y. Solanum rostratum Dunal, EE 3334 NMC, NY. Solanum sp., EE 3086 NMC. ULMACEAE Ulmus americana L.., EE 2493 (cultivado). URTICACEAE Urtica chamaedryoides Pursh, EE 3780 BRIT, NY. VALERIANACEAE Valeriana edulis Nutt., EE NMC; EE 3665 NMC. sail iana sorbifolia Kunth X; EE 3648 BRIT, NMC 3655 BRIT, , EE 2950 NY, NY, TEX. VERBENACEAE Verbena dee Nutt., a 3 OO BRIT, NMC; EE 3330 NY; EE 3494 NY. Verbena gracilis Dest., EE 3 285 cae NMC, Verbena neomexicana (Gray) Small var. a Perry, EE 3146 BRIT, NMC, EE 3589 NY; EE 3202 NMC, NY. Flora de la Laguna de Babjcora 821 Verbena caroliniana L., EE 3734 NMC. Verbena sp., EE 3788 NMC. VIOLACEAE Viola umbraticola H.B.K., EE 3004, 3586 NY. VISCACEAE Arceuthobium vaginatum (Willd.) Presl., EE 3081 NMC, NY. a. juniperinum Gray, EE 3690 T, NMC, NY. ae ee (Nuct.) Nute., EE 3468a BRIT, NMC, NY. VITACEAE Vitis arizonica Engelm. EE. 4241 LILIOPSIDA (Monocotiledoneas) AGAVACEAE ele singuliflora (S. Wats.) Rose, EE 7I8 NMC. Mee: Engelm. var. parryi, EE 3718A. AMARYLLIDACEAE a hirsuta (L.) Cav., EE 3048 TEX, hp mexicana Schult., EE 3452 NMC, EX; EE 3580 BRIT, NMC, NY. COMMELINACEAE Commelina ees Del., EE 3473 TEX; 3473a TE E 3758 TEX; EE 3163 EE 3194 pee EE 3443 NMC. Commelina erecta L., EE 3323 NMC; EE 3568 BRIT, NMC, NY; EE 3604 NMC, NY; EE 3607 NM — Asis ee eau (ines Rafin, EE 3458 EE 3481 NMC, NY, TEX; EE o NMC, NY, TEX; EE 3772 NMC; EE 3207 NMC; EE 3156a NMC. Tradescantia pinnetorum Greene, EE 3009 CYPERACEAE Carex lencodonta Holm., EE 3199 TEX. Cyperus esculentus L., EE 3502 NMC. 822 Cyperus fendlerianus Boeck., EE 3101 NMC, NY; EE 3176 NMC. eon Havicomus Michx., EE 3465 NMC, NY; EE 3591 NMC; EE 3008 NMC. Cyperus fugax 1. Aa EE 2926 NMC, NY; EE 3631 NN Cyperus on a , EE 3192a NMC: EE 3144 NMC, G yperus see L., EE 3175 NMC, NY; EE 3N Caperas ser H.B.K., EE 3110 NMC, 22 NMC, NY; EE 3198 fig Cyperus sphaerolepis Boeck., EE 3155 NMC, NY; EE 3156; EE 3354 NMC, NY: EE 3689 NMC, NY. Cyperus squarrosus L., EE 3632 NMC. cae sp., EE 3036; EE 3345 NY; EE ; EE 3722 NMC, NY. El macrostachya Britt., EE 3279 MC, NY; EE 3310; EE 3304a NMC, Eleocharis att. montevidensis Kunth, EE 3640 * (Willd.) Schlecht., E 3102 Eleocharis tenuis MC, NY. Scirpus acutus Muhl., EE 2946 NMC, NY. IRIDACEAE Sisyrinchium cernuum (Bickn.) eine & Peebles, EE 3769 BRIT, NMC, NY, TEX; EE 3107 NY; EE 3170 me EE 3634 NMC. Sisyrinchium pringlet B.L. Rob. & Greenm., EE 355% Sisyrinchinm

, NY; EE 3371 BRIT. oe as ae i NMC, Mublenbergi dubia (Forn. ye ex Hemsl., 1 NMC, ee mae emersley’ Vasey, EE 3034. si aiey flavida Vasey, EE 3493 BRIT, Yy Muhlenbergia leptoura (Piper) Swallen, EE NY. Muhlenbergia macroura (H.B.K.) Hitchc., EE 3298. ce ‘gia minutisstma (Steud) Swallen, EE 2923 NMC; 3749; EE 3083 ANG, NY; EE 3098 NMC, EE 3635 NMC, NY. eee ‘Clad jee M.E. Jones, EE 3713 >, NY. ae eia montana (Nutt.) Hitche., EE 3748 BRIT, 3 Muhlenbergia elise Scribn., EE 2938 NMC, NY; EE 3714 BRIT, NMC, NY. Muhlenbergia pul. i Scribn., EE 3638. Muhlenbergia utilis (Torr.) Hitche., EE 3489 BRI {C, NY > Muhlenbergia rigens (Benth.) Hitche., EE 3618 NMC, NY; EE 3121 BRIT, NMC, NY. Maeno robusta (Fourn.) Hitche., EE 4370 | NMC, NY. Nica enber ia wolfit ne ) ee EE 3629, EE 3675; EE 3639 NMC, N Mahlenber es wrightit Vasey, EE BRIT, NMC Panicum ee H.B.K., EE 3087 NMC, NY; EE 3141 NMC N Pantcum fasciculatum Swartz, ‘EE 3097. Panicum hirticaule Presl., EE 3519 NY, BRIT; EE 3087; EE 3541; EE 3546; EE 3783 C. Panicum obtusum A.B.K., EE 3520 NY. Panicum a Miche. EE 3087 G; Paspalum us L., EE 3770 NMC; EE ee vaseyanum Scribn., EE 3203. — egies (H.B.K.) Hitche., EE > E 301 3 > 4 5S NY. Piptochactin) NY; EE 3017. Poa fendleriana (Steud.) be EE 3707. Shizachyrium cirratum (Hack.) Woot., EE 3467 BRIT, NMC, NY; EE 3768 NMC; 3174 NY; EE 3131 NMC, NY. Se ee dal sanguinenm (Retz) Alston, EE 3552 BRIT, NMC, N ole i (Beal.) Parodi, EE 3148 Schyz ee sp. EE 3554 NMC, NY. $24 Sipa 17(4) Setaria grisebachit Fourn., EE 3014, 3225. Sporobolus pulvinatus Swallen, EE 3234 Bafa (Poir) Revsceten: EE 3219; NMC, NY. BRIT, NMC, NY; EE 3430 NY. — Sporobolus sp., EE 3514 BRIT, NMC, NY. Sor churn nudipes Nash., EE 3585 NY;EE — Triniocloa laxa Hitchc., EE 3666 NMC S NMC, NY. Lease ae (H.B.K.) Kunth, EE Spor boli atroides (Torr.) Torr., EE 3233. 3765 NMC, EE 3654 NY; EE 3029: Sporobolus cryptandrus (Torr.) A. Gray, EE 3559 re 34 Y AGRADECIMIENTOS Deseamos expresar nuestro agradecimiento a Rupert C. Barneby, B. L. Turner, G. Nesom, y M. Mayfield, por el apoyo en la identificacién y corroboraci6n de los materiales botanicos. REFERENCIAS Austin, D.E y R.A. Peproza. 1983. Los géneros de convolvuléceas mexicanos. Bol. Soc. Bot. Mex. 44:3—7 BarkLEY, FA. 194 ere iaceae. In: Lundell, C.L., ed. Flora of Texas. Southern Meth- odist University Press, Dallas. 3:1—150. Barnesy, R.C. 1964. Atlas of North American Astragalus. Mem. New York Bot. Gard. 3:1-1188. Barnesy, R.C. 1977. Daleae imagines. Mem. New York Bot. Gard. 27:1—892. Barnesy, R.C. 1989. Intermountain Flora vol. 3, part B, Fabales. New York Bot. Gard. Bicker, K.M. 1979. A monograph of the genus Lasianthaea (Asteraceae). Mem. New York BEETLE. . 1947. Poales (Cyperaceae, Scirpeae). North American Flora. New York Bot. Gard. Ig: 179-504 BLACKWELL, W.H. 1968. Revision of Bowvardia (Rubiaceae). Ann. Missouri Bot. Gard. 5:1—30. Brake, S.F. 1913. 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WILLIAMS. 1983, Field guide to orchids of North America. Uni- verse Books, em York. WINDLER, D.R. 1974. A systematic treatment . the native unifoliolate crotalarias Mes ueunesder f North America. Rhodora 76:151—204. YUNCKER, T.G. 1965. Cuscuta. North American oe a HI. Part 4. New York Bor. Gard. ZANONI, T.A. 1978. The american junipers of the section Sabina (J/aniperus-Cupressaceae). A century later. Phytologia 38:433-453. ZOHARY, M. and D. HeEtier. 1984. The genus Tréfol/imm. the Israel Academy of Sciences and Humanities. Jerusalem. Israel. CONVENTIONS FOR REPORTING PLANT CHROMOSOME NUMBERS JOHN L. STROTHER University Herbarium 1001 Valley Life Sciences Building University of California Berkeley, CA 94720-2465, U.S.A. GUY L. NESOM Texas Regional Institute for Environmental Studies Sam Houston State University Huntsville, TX 77341-2640, U.S.A. ABSTRACT Original reports of chromosome numbers should reflect actual observations. To facili- tate comparisons among taxa, reports of chromosome numbers in floras and other taxo- nomic works should be uniform. RESUMEN Los recuentos originales de nimeros cromosémicos deberan reflejar las observaciones verdaderas. Con el fin de facilitar las comparaciones entre taxa, las citas de los numeros cromosémicos en floras y otros trabajos taxonémicos deberfan ser uniformes. Because of variations in ways in which chromosome numbers are re- ported and because untenable statements (such as “z = 6 II”) continue to appear in botanical literature, we outline here some guidelines for report- ing chromosome numbers in plants, in the hope of encouraging consis- tency. An important consideration in determining what to report, and how to report it, is context. We recognize two broad contexts in which chromo- some numbers are usually cited: original observations and summaries. ORIGINAL OBSERVATIONS Reports of original counts of chromosome numbers should precisely in- dicate what was seen. We report chromosome counts from normal gameto- phytic cells as 2 numbers and counts from normal sporophytic cells as 2n numbers. We give examples for a group of plants with a base number of x = 9: 1. n = 9 for counts from mitotic figures in gametophytic cells (e.g., pollen tubes, cells from thalli of fern gamecophytes) of a plant of a diploid taxon. Counts of 7 = 9 may also be reported from observations of dyads after cell- wall formation (see also, example 3). While such use of “2 = 9” in referring Stipa 17(4); 829-831. 1997 830 Stipa 17(4) to a specific observation of chromosomes in cells of a gametophyte would be an accurate notation, a few qua — ifying words might well be included in the report because chromosome counts are seldom made from gametophytic cells. 2. 2n = 18 for counts from mitotic figures (usually metaphase) in sporo- phycic cells (e.g., root-tip cells, leaf-primordia cells) of a diploid plant with Cia = 13. 3. 2n = 9 II for counts from meiotic figures (usually diakinesis or first metaphase) in sporophytic cells (e.g., pollen parent cells, spore parent cells) in a diploid plant. To refer to such counts from sporophytic cells as “ga- metic’ or to cite such an observation as “v = 9 IL” would be incorrect. The single “”” should be used only in reference to counts from gametophytic cells. Observations of 9 chromosomes at each pole in incom pletely divided dyads of meiotic cells of a diploid plant should be reported as: “27 = 18 (9 at each pole in dyads)” — see also, example 1. In diploid hybrids, less than perfect pairing may be encountered and meiotic figures such as: 27 = 8 I] + 21,27 =711 +4], 2n = 18 /etc., may be seen in plants with 2” = 2x = 18. 4, 2n = 16 for counts from mitotic figures in sporophytic cells of a dysploid plant; 27 = 8 II for counts from meiotic figures in — celis (e.0,, pollen or spore parent cells) of the same dysploid plant. {For discussions of dysploidy and aneuploidy and differences between them, see Dyer et al. (1970), Garber (1972), and papers cited by them. In brief, dysploids are characterized by changes in chromosome number usually accompanied by insignificant, if any, changes in numbers of genes per cell. In botanical literature, dysploid series sometimes have been mistakenly called “aneup- loid series.” In aneuploid plants, one or more chromosome(s) of the basic set is represented fewer or more times than expected in each cell and the number of genes per cell is significantly different from that normally found in euploid plants of the same population or taxon. Aneuploid plants are referred to as monosomics, trisomics, etc., usually exhibit greatly reduced fertility, and usually exhibit other traits, including morphological ones, reflecting their genic imbalance. Exceptionally, trisomic plants may be fer- tile, may form reproductively isolated populations, and may even evolve into distinct taxa. | 5.2” = 19, 2n = 17, 2n = 35, etc., for counts from mitotic figures and 27 = 9IN+11,27=8I0+11 27=1711+1 I, etc., for counts from meiotic figures in sporophytic cells of aneuploid plants, which have certain chro- mosomes represented fewer or more than the expected number of times in each somatic cell (see discussion at example 4). 6. 2” = 27 for counts from mitotic figures and 27 = 911 + 91,2” = 8IL4 I, 27 = 271, etc., for counts from meiotic figures in sporophytic cells of STROTHER AND Nesom, Reporting chromosome numbers 831 triploid plants, including hybrids. For such plants, statements such as 27 = 27 may usefully be expanded to 27 = 3x = 27 to call attention to the presence of 3 sets of chromosomes in sporophytic cells. To Nee oe for such plants would be misleading. Normally, 37 cells, 47 cells, 5x cells, etc., are found only in endosperm tissues, which we consider to be neither gametophytic nor sporophytic. 7. 2n = 36 for counts from mitotic figures and 2” = 18H, 2” = 1611 + LIV, 2n = 911+ 18 I, etc., for counts from meiotic figures in sporophytic cells of tetraploid plants, including hybrids, with 2” = 4x = 8. 2n = 45 for counts from mitotic figures and 2” = [various combinations of univalents, bivalents, and/or multivalents] for counts from meiotic figures in sporophytic cells of pentaploid plants, including hybrids, with 27 = 5x = 45 And so on for higher ploidy levels. Reports of “B” or supernumerary chromosomes, or fragments, or rings or chains of chromosomes, etc., can be made by amplifying one of the ex- amples above (27 = 9 II + 2f: 2n = chain of 3 + SII + 81; 2” = ring of 4 16 II; etc.). SUMMARIES In floras and some other taxonomic works, in which chromosome num- ber is included in descriptions or in some other bases for comparison, the chromosome numbers given are commonly each derived from more than one observation and are essentially indicators of the ploidy level(s) known for each taxon. We recommend statements such as “2” = [total number of chromosomes per cell}” for each taxon regardless of the form of the original report(s). In this way a single format allows direct comparisons of chromo- some numbers among euploids (diploids and “normal” polyploids), aneup- loids, and dysploids. ACKNOWLEDGMENTS We thank FE. Flores-Pedroche for the reswmen and D.E. Johnson, D.W. Kyhos, R.L. Moe, J.S. Mooring, and A.R. Smith for comments on early drafts of this paper. REFERENCES Dyer, A.F., K. JONG, and J.A. ae aTTER. 1970. Aneuploidy: A redefinition. Notes Roy. Bot. Gard. Edinburgh 30:177- Garper, E.D. 1972. ore An introduction. McGraw-Hill Book Co., New York. NOTES NOTES ON HEMEROCALLIS MIDDENDORFFII (LILIACEAE) IN KOREA Hemerocallis middendorffit complex (Liliaceae) grows in northeastern China, northern Korea, the central and northern Japanese Archipelago, and parts of the Kuril Islands (Noguchi 1986). As the taxonomy of the complex has been extremely controversial, Noguchi (1986) conducted a detailed study of morphological analyses on 14 characters collected from 35 localities throughout the geographic range of the complex in Japan. More recently, she also investigated the historical differentiation processes of the complex in Japan with chloroplast DNA restriction site variation (Noguchi et al. 1995). For H. middendorffii in Korea, only a few locations in northern re- gions has been documented in previous studies (e.g., Fig. 2 in Matsuoka & Hotta 1966; Fig. 35 in Noguchi 1986; Noguchi et al. 1995). In an at- tempt to improve our knowledge of the genus, we have collected samples from 75 locations in South Korea during the past 10 years to determine the overall distribution patterns for Korean species and to analyze morpho- logical variation within and among species (Chung & Kang 1994, Kang & Chung 1994, Kang & Chung 1997). Based on these studies, we recognize five Hemerocallis species native in Korea: H. hakmunensis Nakai (=H. micrantha Nakai), H. thunbergii Baker (=H. coreana Nakai), H. middendorffii Tr. et Mey., H. hongdoensis M. Chung & S. Kang, H. taeanensis 8S. Kang & M. Chung. Except H. middendorffii, the variation patterns of several morphological char- acters and distribution of the four Korean species have been documented 1n these studies. From May to June 1995, while conducting field trips in the central and southeastern Korean Peninsula, we encountered three popula- tions of H. middendorffii (see arrows in Fig. 1). Samples of rootstocks were collected from the locations and were grown under uniform conditions in Biology Plant Growth Facilities at Gyeongsang National University. Voucher specimens of all collections are deposited at GNUC. From May to June 1996, the 12 morphological characters (Chung & Kang 1994) were measured for 15 individuals from the three populations. In addition, the herbaria of Seoul National University (SNU) and Tokyo University (TD were visited in order to determine the overall geographic distribution pat- terns for H. middendorffii in Korea. Here we report the distribution and description of H. middendorffii in Korea to provide an information for better understanding the complex in east Asta. Although Matsuoka and Hotta (1966) and Noguchi (1986) presented distribution maps in which the species grows only in northeastern Korea, Stipa 17(4): 833. 1997 Sipa 17(4) Fe. F re J : MN r wig S R ie a e ' vee a i ; ( Lan | ‘ a“ f ‘ ] { 5 — e ee Ses (a c ai yy | oo A ws -\ va ° x ~ < \ 131 E \ \ 0 : \ 38°N ° \ A qx val Ne we . of ADC % oh Y n 4 RES pO J 5 Oe howe ts i oe 782, : Dereh, { } a ~ x a laa yw! 0 50Km a LU Aa —~, 7 ’ “~ \ os . Documented distribution of Hemerocallis middendorffii in Korea. Arrows (A, B, and es recently discovered populations. Population A (alt. 800 m) grows along a stream- side with Hosta capitata (Koidz.) Nakai, L lium sp, Polygonatum sp., and grasses. Population B (alt. 400 m) grows on the margins of pine forest along a streamside Population C (ale 240 m) grows on the hillside of mesoic pine-oak unc erstory Stipa 17(4): 834. 1997 Nores 835 as presented in Figure 1, H. middendorffii is widely distributed from the central to northern Korean peninsula. According to Noguchi (1986), popu- lations of H. middendorffii in Japan maintain complex local variation in several morphological characters partly due to diverse ecological habitats such as alpine meadows, low grasslands, and margins of forest understo- ries. Sizes of the three Korean populations were relatively small (< 100 individuals per population) and these isolated populations were found along stream sides of hillsides. The description of H. middendorffii in Korea is as follows: Hemerocallis middendorffii Traut. & May., Flor. Och. Phaen. 94. 1856. Leaves 30—60 cm long, 0.8—-1.5 cm wide, greenish yellow. Scapes erect, 20-60 cm long. Inflorescences are basically short dichotomous or — type with 2-4 flowers. Inflorescence bracts ovate, greenish, 2.5—5.5 long, 1.1—3.0 cm wide. Flower weakly fragrant. Perianth Seca lew. tubular-funnelform, 6.8—9.6 cm long; tube short, 1.0-1.8 cm long; throat orange-yellow, inner ee lobes 4.6-6.3 cm long, 1.1—2.3 cm wide, outer perianth lobes 4.7—6.7 cm long, 0.9-1.6 cm wide. Stamens 3.5—4.5 cm long, inserted; anthers ca. 5 mm long, dark brown with purple tint. Style filiform, ca. 8.5 cm long, exserted beyond the stamens. Capsule | .8— 2.2 cm long, apex emarginate; seeds shining black, ovoid, angled below, 7 mm long, 5 mm wide. Flowering from late May to June in Korea; fruiting and aces seed in July. S| xamined: KOREA. Prov. Spero ss itt a shi, 17 Jul 1909, T. Napa sm. (TI; Mt. Kwanmyobong, 20 Jul 1936, Toh 4507 (SNU). Prov. ay sat ngs -do: Sambang, 6 Jun 1933, B.S. Toh s.n. GNU) Mt. Baekam, 15 Jul 1934, B.S & HJ. Shim 3549 (SNU); Musan, Aug 1913, De Morr 311 (T). Prov. ee ead -do: Mt. Myohwang, 2 Jul 1932, B.S. Toh, HJ. Shim en (SNU). Prov. Kyeong-do: Kwangrung, 12 Jun 1932, B.S, 900 (S . Prov. Ch’ungch’eongbuk-do: Mt. Sokri, B.S. Toh, & HJ. Shim 9596 (SNU). Prov. Kangwon do: Mc. Myeon, S.H. Of 5.7. (SNU); Mureung-ri, Jungsun-gun, 11 Jun 1995, 8. Kang, M. Chung, & G.S. Oh 2040 (GNUC),. Prov. eae k-do: Bulyoung Valley, Wuljin- gun, 29 May | fe S. Kang, M. Chung, & G.S. Ob 2038 (GNUC); Bongwha-gun, 10 Jun 995, 8. Kang, M. Chung, & G.S. Oh s056 (GNUC). ACKNOWLEDGMENTS We thank the herbaria directors and curators of SNU and TI. This resreach was supported in part by a Korea Science and Engineering Foundation grant (96-0500-006-2) to MGC. —Soon Suk Kang and *Myong Gi Chung, Department of Biology, Gyeongsang National University, Chinju 660-701, The Republic of Korea; Ki Bae Park, Sipa 17(4): 835. 1997 836 Stpa 17(4) Department of Horticulture, Ansung National University, Ansung 456-749, The Republic of Korea. *Corresponding author. REFERENCES CHUNG, M.G. and $.S. Kano. 1994. ee analysis of the Hemerocallis L. (Liliaceae) in Korea. J. Plant Res. 107:165-— KANG, S.S. and M.G. CHUNG. cn — hakuunensis (Liliaceae) in Korea. Sida 16:23-31. 1997. Hemerocallis taeanensis (Liliaceae): a new species from Korea. Syst. t. (in pre a. Mats UOKA, M. and M. Horta. ee Ol assification of Hemerocallis in Japan and its vicinity. Acta Phytotax. Geobot. 22:25—43. (in Japanese) Nocucui, J. 1986. Geographical mi ecological differentiation in the Hemerocallis dumortiert complex with special ed to its karyology. J. Sci. Hiroshima Univ., Hiroshima, Japan, Ser. B. Di. 2, Bot. 20: 93 —__—* M. Tasaka and 1 M. tae: 1995. The historical differentiation process in Panels middendorffpi (Liliaceae) of a based on restriction site variations of chloroplast DNA. J. Plant Res. 108:41—4 si Stipa 17(4): 836. 1997 CALYSTEGIA (CONVOLVULACEAE) IN TEXAS All past Texas floristic works have identified either a single Ca/ystegia spe- cies, C. sepium (L.) R. Br., or at most, two species in the state C. sepium and C. silvatica (Kit.) Griseb. (e.g., Correll & Johnston 1970; Hatch et al. 1990; Jones et al. 1997). When we recently examined specimens at BRIT for the forthcoming Shinners & Mahlers (lustrated Flora of North Central Texas (Diggs et al.), we discovered that there actually were three species, the third being C. macounti (Greene) Brummitt. Although there has been marked confu- sion and misinterpretation of Ca/ystegia species in the past (e.g., Tryon 1939), the proper delimitation and names for North American taxa were provided by Brummitt (1965, 1980). Few students of Great Plains plants other than Austin (1986, 1992a) have recognized these three taxa. Other authors have continued an incorrect usage that masked actual biodiversity (e.g., Taylor & Taylor 1989, 1994; Hatch et al. 1990; Jones et al. 1997). In this note we wish to record the data we have on these species for fucure students of the region. Details on one or more of these species are also available for the Intermountain region (Holmgren 1 984), New Mexico (Austin 1990) and Arizona (Austin 1991). While these taxa differ by what seem to be trivial characters on herbarium specimens, they are more dis- tinctive when alive, have different ecological eis and historically occupied somewhat different geographic regions. The following key 1s modified from Austin (1986). . Leaf sinus quadrate (= nearly square); blade tissues not beginning for 2—5 (-10) mm from petiole attachment (only vascular tissue near attachment) vatica subsp. fraterniflora ae _ Leaf sinus V- or U-shaped; blade tissues beginning at the point of petiole attachment bo Basal lobes of leaf blades with |—2 small, cooth-like angles; plants nor- mal — y glabrous or with a few trichomes on petioles ...... C. sepium subsp. angulata Basal lobes of leaf blades rounded; plants normally pubescent on all veg- etative parts NO a . Plants twining throughout; leaf blades narrow eae 3:1 length to widch ratio) sepium ees ucla . Plants semi-erect or sparsely twining; leaf blades (mostly acer to width ratio) . macouniti Calystegia macounii (Greene) Brummitt, Ann. Missouri Bot. Gard. 52:215. 1965. Basionym: Convolvalus macounii Greene, Pittonia 3:331. 1898. Type: CANADA. SASKATCHEWAN: Assiniboia, Milk River, Aug 1895, John Macoun 11883 (aotoryee: ND; isotype: BM). = lated floral bracts make C. macouni distinctive. Parker (1972, Fig. 111B p. 233) incorrectly illustrated this species as C. sepium. So far as known, that is the only published drawing of The semi-erect habit and strongly in Sipa 17(4): 837. 1997 838 Sipa 17(4) C. macount. This is a Plains species that is distributed from Canada to western Minnesota and south to Texas, New Mexico and Arizona. Probably the plants were spread westward along logging railroads across New Mexico and Arizona. Because so few people have identified the species, little of its ecological preferences are known. However, C. macounii is typically associ- on ated with moist sanc in woodlands. Also, this is the first species to flower in the spring, usually y soils near waterways in open grasslands or openings in May and June. Although flowering periods overlap, this species is often in fruit by the time the others begin to flower. TEXAS. Cooke Co.: Silty bank, above small stream, limestone area, 8.8 mi W of Gainesville, 143 May 1951, L.H. Shinners 13256 (BRIT-SMU). a le ae Pt (LJ ROBE ee limnophila (Greene) Brummict, Ann. ri Bot. Gard. 52:216. 1965. Basionym: Convolvulus limnophilus Greene, a 3:29. 1898. Type: CALIFORNIA. Suisun Marsh, Aug 1883, E.L. Greene sah 09 (syntypE: ND fide Brummit, cf. Austin 1992b). Convolvulus repens L., Sp. PL. 58. 1753. Type: specimen 218.55 in herb. Linnaeus (LINN, not seen; microfiche oe Calystegia sepium var. repens (L.) A. Gray, Syn. Fl. N. Amer. 2(1):215. 1878. (cf. Tryon 1939; Correll & Johnston 1970: 1247). Convolvulus nashii House, Muhlenbergia 5:66. 1909. Type: FLORIDA. Lake Co.: Eustis, Nash 609 (HoLoryeE: Colombia Univ., not seen; isorypr: NY) The leaf blades abruptly narrowed (mostly 1—2 cm wide) above the ex- ag gerated sagittate bases make this subspecies distinctive. Rarer specimens (e.g., Boon 130 & Webster 487) have leaves wider than this but usually still have the flared sagittate bases making them identifiable. This is a wetland species that ranges from Florida to California across the southern USA and even into northern Mexico. TEXAS. aveanices Co.: N of Fulton, 10 Jun 1960, FE Jones 4086 (TEX), Galveston Co.: Park on E.M. 517 (Bayshore Deve) S - Bacliff, 10 May 1976, F Waller 3810 (TEX), Harris Co.: 5 mi NW of LaPorte, 2 4 Jul 1943, E. Boon 130 (TEX) ). Jefferson Co.: Beau- mont, 9 Apr 1924, B. Thay? 3163 (TEX). Orange Co.: Port of Orange, 27 Jun 1967, D. Correl] 34279 (LL). Terrell Co.: 7 mi upstream from jct. of Independence Creek & Pecos River. 5—6 Jul 1949, G. Webster 487 (TEX). eo Reon (L.) R. Br. subsp. angulata Brummitt, Kew Bull. pe ee) 980. Type: IDAHO. Canyon - (0.: Falks Store, 28 Jun 1910, Machride eta MO; isotypes: GH, NY. Both C. sepium and C. silvatica are ies or twining plants. Alchough the floral bracts are inflated in C. si/vatica, its habit and leaf characters are different from those of C. macounii. Leaf characters, as noted in the key, distinguish C. sepiwm from C. silvatica. Calystegia sepium subsp. angulata belongs to a disturbingly variable complex chat has probably been properly segregated into taxa by Brummite (1965, 1980). Unhappily, Brummitt SIDA 17(4): 838. 1997 Nores 839 has never published a full account of how to distinguish the varieties and it is known that some intergrade (Brummitt 1980). Subspecies angilata is native from New Brunswick to British Columbia in Canada and south to Washington, Oregon, Idaho, Utah, New Mexico, Texas and South Carolina in the U.S.A. (cf., Brummitt 1980 for specimen citation). This ts a wood- land and forest taxon typically confined to openings and glades; it usually flowers between June and August. ing oO TE aes Co.: ca. 8.5 mi W of Sabine Pass, from a sand dune area near t beac h, - May 1973, J. Taylor & C. Taylor 13242 (BRIT-SMU). Calystegia ee (Kit.) Griseb. a fraterniflora (Mack. & Bush.) Brummitt, Ann. Missouri Bot. Gar 92: 216. 1965. Basionym: Convol- vulus sepium (L.) R. Br. var. fraterniflorus ee & Bush, Man. Fl. Jackson Co. Mis- souri 153. 1902. Typg: U.S.A. MISSOURI: Martin City, 9 Jul 1899, Mackenzie G Bush s.n. (not traced; ropotyPe, 28 Jun 1905, Bash 3037 (GH, MO). The quadrate sinus in the leaf blade base is the easiest way to recognize Calystegia silvatica subsp. fraterniflora. This North American subspecies, related to the Old World C. si/vatica subsp. si/vatica, is native from New Hampshire to Kansas and Florida to Oklahoma and Texas. Within that region it typically occurs along stream margins on sandy soils in deciduous forests. This species has the longest flowering season of the three, often going from May to September. TEXAS. Anderson Co.: E of Palestine, 2 Oct 1965, D. Correll & H. Correll 31748 (LL). Chambers Co.: rare along brackish marsh in the Anahuac National Wildlife Refuge, 5 of Anahuac and on shore of Galveston Bay, 5 Jun 1987, L.E. Brown 111 55 (BRIT). Ochiltree Co.: bank of Wolf Creek, 12 mi SE of Perrycon on US 83 & 5 mi E of hwy., 13 Jul 1957, C. Wallis 4849 a Potter Co.: Palo Duro Caftion-Harding Road. 15 Jun 1929, B. Tharp 6289 (TEX —Daniel F. Austin, Department of Biological Sciences, Plorida Atlantic Univer- sity, 777 Glades Road, Boca Raton, FL 33431, U.S.A., daustin@fau.edu, George M. Diggs, Jr, Department of Biology, Austin College, Sherman, Texas 75090, U.S.A. & Botanical Research Institute of Texas, gdiggs @austinc.edu, and Barney L. Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Ft, Worth, TX 76102, U.S.A., barney @ brit. org. REFERENCES Austin, D.F. 1986. Convolvulaceae. In: Great see Flora Association. Flora of the Great Plains. Univ. Press Ais Kansas, ae: Pp. 1990. d -klist of New a ican Convolvulaceae. Sida 14:273—286. —«*'991. Annotated checklist or Arizona Convolvulaceae. Sida 14:443—457. ee 1992a. Rare Convolvulaceae in the southwestern United States. Ann. Mis- souri Bot. Gard. 79:816. Sipa 17(4): 839. 1997 Sipa 17(4) —_______. 1992b. Studies of the Florida Convolvulaceae—V. Ca/ystegia. Florida Sci. 55:58-G61. Brumitr, R.K. 1965. New combinations in North American Ca/ystegia. Ann. Missouri ot. Gard. 32:214—2 —_______—.. 1980. Further new names in the genus Ca/ystegia (Convolvulaceae). Kew Bull. 35:327-334. Corre, D.S. and M.C. Jounsron. 1970. Manual of the vascular plants of Texas. Renner, s Research Foundation. ee C. M. Jr., B.L. Lipscoms, and R.J. O’KENNON. forthcoming). Shinners & Mahler's tllustrated Ara of North Central Texas. To be published in 1998 by the Botanical Re- search Institute of Texas, Fort Worth. Haren, S.L., K.N. Ganpui, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agriculture Experiment Station, Texas A&M Univ., College Station. Mis- cellaneous Publication 1655, HoLtmMGREN, N.H. 1984. Convolvulaceae. In: > Cronquist, A., A.H. Holmgren, N.H. Holmgren, J.L. Reveal, and P.K. Holmgren, eds. aaiinentintars flora. New York Bo- tanical Garden and Columbia University Press, New York. 4:74—77. Jones, 5.D., J.K. Wiper, and P.M. Monrcomery. 1997. Vascular plants of Texas: A com- prehensive checklist including synonymy, bibliography, and index. University of Texas Press, Austin. PARKER, | 1972. An illustrated guide to Arizona weeds. University Arizona Press, Tucson, Taytor, R.J. and C.E.S. Taytor. 1989. An annorated list of the ferns, fern allies, gymno- sperms flowering plants of Oklahoma. Southeastern Oklahoma State U niversity, Durant —______.. 1994. An annotated list of the ferns, fern allies, Jeena and eee plants of Oklahoma, 3rd ed. Southeastern Oklahoma State University, ee R.M. Jr. 1939. The varieties of Convolunlus spithamaens and of C. ae iste “415-423. Sipa 17(4): 840. 1997 HETEROTHECA LATIFOLIA (ASTERACEAE), NEW TO THE FLORA OF IOWA Heterotheca latifolia Buckl. (Asteraceae: Astereae) is a taprooted yellow-rayed annual native to open sandy sites from the southern Great Plains and Ozark region to northern Mexico, which is sporadically naturalized farther north and east (Wagenknecht 1960; Harms 1965; Rolfsmeier et al. 1991). Plants of this species (referable to var. macgregoris B. Wagenkn.) were recently col- lected for the first time in Iowa. Voucher specimens: IOWA: Lee Co.: Montrose Twp. T6GN RSW sect. 3 W1/2, along railroad tracks parallelling US Hwy 61, juse N of the Linger Longer Rest Area, 1 mi N of Montrose, scattered in dry sand prairie with scattered junipers, on slope overlooking the Mississippi River, 3 Oct 1997, T-G. Lammers 10499 (F, ISC, NY, TEX). The site is a relatively undisturbed dry sand prairie of a few acres extent, situated on a moderate slope from US Highway 61 to railroad tracks, on a terrace of the Mississippi River. This prairie is dominated by Panicum virgatum L. and Eragrostis trichodes (Nutt.) A. Wood (10495), with a few scattered trees of Juniperus virginiana L. (10496). [ first visiced this site nearly 20 years ago, on 17 May 1978. That early in the season, the only species in flower were Androsace occidentalis Pursh (1493), Corydalis micrantha (Engelm.) A. Gray (1494), Draba reptans (Lam.) Fern. (1492), and Viola rafinesquii Greene (1495). Another early-season stop a year later, on 2 June 1979, added only Delphininm virescens Nutt. (27135): The opportunity to make a visit later in the season did not present itself until recently. It was on this occasion that the Heterotheca was discovered. Other psammophiles collected at the same time included Ambrosia psilostachya DC. (10498), Croton glandulosus L. vat. septentrionalis Muell.- Aro (10502), Helianthus petiolaris Nutt. (10497), Polanisia dodecandra (1.) DC. (10503), and Triplasis purpurea (Walt.) Chapm. (10494). Gleason and Cronquist’s (1991) manual, which includes Iowa within its scope, does not mention Hererotheca latifolia as either an accepted species or a synonym. Apparently, these plants were included in their concept of H. subaxillaris (Lam.) Britt. & Rusby, as was done by most authors prior to Wagenknecht’s (1960) revision. In any event, no species of Heterotheca sec- tion Heterotheca was reported from Iowa in Wagenknecht’s (1960) revision of the group. Neither H. s#baxillaris nor H. latifolia was included in the recent checklist of Jowa vascular plants (Eilers & Roosa 1994). No species of the genus was reported from the southeastern corner of the state by Davidson (1959), nor from Lee County by Peck et al. (1981). Despite its occurrence as part of a relatively undisturbed native plant Sipa 17(4): 841. 1997 842 Sipa 17(4) community, the population of Heterotheca latifolia var. macgregoris in south- eastern Iowa is presumed to be naturalized and not native, based on its disjunction from the main range of the species and its proximity to road and rail transport. The nearest approach of native populations of this taxon is in eastern Kansas; more proximate populations in [linois and Missouri are considered introduced (Wagenknecht 1960; Harms 1965). The nearest Iinois population of that taxon reported by Mohlenbrock (1986) and Mohlenbrock and Ladd (1978) is in Henry County, 115 km to the north- ast. The nearest Missouri population mapped by Steyermark (1963) is in St. Louis Co., approximately 230 km south-southeast; however, those plants are referable to var. arkansana B. Wagenkn. (Wagenknecht 1960; Steyermark 1963).—Thomas G. Lammers, Department of Botany, Field Museum of Natural History, Chicago, IL 60605-2496, U.S.A. REFERENCES Davipson, R.A. 1959. The vascular flora of southeastern Iowa. Stud. Nat. Hist. lowa Univ. 29(2):1-102. Eiters, LJ. and D.M. Roosa. 1994, The vascular plants of Iowa. University of lowa Press. Iowa City, lowa. GLEaAson, | | re and A. CRONQUIST. Me |. Manual of vascular plants ee United States and adjacent Canada, ed. 2. New York Botanical Garden. he Harms, V.L. 1965. Biosystematic ae in the Heterotheca ie complex. Trans. Kansas Acad. Sci. 68:244—257, MOHLENBROCK, R.H. 1986. Guide to the vascular flora of Illinois. Southern Illinois Uni- versity Press. Carbondale. ‘oe K, R.H. and D.M. Lapp. 1978. Distribution of [linois vascular plants. South- em epee University Press. Carbondale. Peck, J.H., T.G. Lammers, B.W. HaGian, D.M. Roosa, and L.J. Eiters. pi - checklist of the vascular flora of Lee County, lowa. Proc. Iowa Acad. Sci. 88:159— Ro.rsMEIER, $.B., R.B. Kaur, and D.M. SurHertaNp. 1991. New and ete Horistic records for Nebraska—4. Trans. Nebraska Acad. Sci, 18:141—150. J.A. 1963. Flora of Missouri. Iowa State U niversity Press. Ames, lowa. STEYERMARK, Revision of Heterotheca section Heterotheca. Rhodora 62:61 —76, W AGENKNECHT, B.L. 1960. 97-107 Stipa 17(4): 842. 1997 CERASTIUM PUMILUM AND STELLARIA PALLIDA (CARYOPHYLLACEAE) NEW TO TEXAS Early-flowering annual members of the Caryophyllaceae are often overlooked in the North American flora. Cerastinm pumilum Curtis and Stellaria pallida (Dumort.) Crép. are no exception, especially since they resemble widely distributed weedy species; Cerastium fontanum Baumg. emend Jalas and y. During a collecting trip in May of 1997, Reznicek made six collections of various species of Caryophyllaceae. Of these six, two were taxa not previ- ously reported from Texas (Correll & Johnston 1970; Turner 1996; Jones et ale 1907): Cerastium pumilum Curtis (CURTIS’ or DWARF MOUSE-EAR CHICKWEED) has been reported from disturbed areas in many of the eastern and south- eastern United States (Rabeler & Cusick 1995), including Missouri (Steyermark 1963), Arkansas, and Tennessee (Rabeler & Thieret 1988). It is now being reported from Texas from the following collection: —_— Stellaria media (L.) Villars respective TEXAS. Kaufman Co.: roadside ditch, E side of FR 205, 0.7 mi NW of jet. with TX Hwy 80, W side of Terrell, 32° 44'48"N, 96°18'19"W, 2 May 1997, A.A. Reznicek 10336, S.D. Jones, & S.A. Reznicek (BRCH, MICH). Plants were found in moist, bare soil of lower slopes of a disturbed grassy/ sedgy ditch along with Cerastinm glomeratum Thuill., Sagina decumbens (Elliott) Torrey & A. Gray, and Myoswrus minimus L. Cerastium pumilum resembles a small plant of C. fontanum and will key to C. fontanum in Turner's key to Texas Cerastimm (Turner 1996). The two —_— species can be separated by the following couplet: L. Flowers 8-10 mm wide; petals deeply notched (to 1/2 the petal length); sepals pilose, seldom glandular-hairy; stamens 10, capsules mostly over 8 mm long; perennial, often with non-flowering basal shoots .......0:06 es C. fontanum — _ Flowers 5—6 mm wide; petals shallowly notched (to 1/4 the petal length); sepals glandular-hairy; stamens 5 (rarely 10); capsules less than 8 mm long; annual ... C. pumilum Stellaria pallida (Dumort.) Crép. [= Stellaria media subsp. pallida (Dumort.) Asch. & Graebn.} (LESSER CHICKWEED) 18 also a plant of disturbed areas. Like Cerastinm pumilum, it has been reported from a number of nearby southeastern United States including Arkansas (Thomas et al. 1991) and Tennessee (Chester et al. 1997). It is now being reported from Texas from the following collection: TEXAS. Hopkins Co.: roadside, NW corner of jct. of Hopkins Co. Rd. 2401 & TX Hwy 11, ca. 3 mi SE of Pickton, 32°59'5S8"N, 95°21'10"W, 3 May 1997, A.A. Reznicek 10361, S.D. Jones, & S.A. Reznicek (BRCH, MICH). Stipa 17(4): 843. 1997 844 Stipa 17(4) Plants were found in dry, sandy, open soil of a disturbed roadside growing under Petrorhagia dubia (Raf.) G. Lopéz & Romo, another somewhat recent (1960s) introduction to the Texas roadside flora (Rabeler 1985). Stellaria pallida will key to S. media in many floristic manuals, including Correll and Johnston (1970). The two species can be separated by the fol- — lowing couplet: |. Plants green, rarely yellowish green; flowers usually open, with petals; se- pals 4 mm or longer, uniformly green; seeds dark brown, (0.8—)0.9-1.3 mm in diameter, the surface covered with rounded papillae ......0..........00.... 8. media . Plants yellowish green; flowers usually cleistogamous, apetalous; sepals sel- dom over 3 mm long, often with a red basal band: seeds yellowish brown, 0.5—0.8 mm in diameter, the surface covered with acute papillae .....0...0...... S. pallida The most consistent features distinguishing §. pallida from S. media are the color, size, and surface of the seed (Morton 1972; Rabeler 1988), Un- fortunately, many specimens (including Reznicek 10361) lack the red band at the base of the calyx which, when present, allows immediate recognition of S. pallida. Rabeler and Cusick (1995) discussed the authorship of S. pallida, noting that, contrary to many sources, Piré did not publish the combination Sre//aria pallida. The next usage of the combination they were aware of was by Junger in 1878. After that paper appeared, Rabeler learned, as Kent (1997) noted, that Francois Crépin made the combination twelve years earlier (Crépin 1866) than Junger. Both of these plants are probably recent introductions and, as Rabeler has noted in other reports of these species (Rabeler 1988; Rabeler & Cusick 1995), are likely to be found elsewhere in similar disturbed areas.—R/ch- avd K. Rabeler, rabeler@umich.edu, and Anton A. Reznicek, rexnicee@umich.edu, University of oe Herbarium, North University Building, Ann Arbor, MI 48109-1057, U.S./ REFERENCES CuesTer, E.W., B.E. Worrorp, and R. Krai. 1997. Atlas of Tennessee vascular plants. volume 2. Angiosperms: Dicots. Austin Peay St. Univ. Center Field Biol. Misc. Pub J Corrett, D.S. and M.C, Jounston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner Creepin, F. 1866. Manuel de la flore de Belgique. Gustave Mayolez, Bruxelles. Jones, $.D., J.K. Wiper, and P.M. MontGomery. 1997. Vascular plants of Texas: A com- prehensive checklist including synonymy, bibliography, and index. Univ. of Texas Press. ustin Kent, D.H. 1997. The correct authority for lesser chickweed, Sre/larta pallida (Caryophyllaceae). Watsonia 21:364 Sipa 17(4): 844. 1997 845 Notes Morton, J.K. 1972. On the occurrence of Stellaria pallida in North America. Bull. Torrey Bot. Club 99:95-97. RaBELER, R.K. 1985. Petrorhagia (Caryophyllaceae) of North America. Sida 11:6—44 1988. Eurasian introductions to the Michigan flora. IV. Two additional species of ee ae in Michigan. Michigan Bot. 27:85-88 nd A.W. Cusick. 1995 [“1994"}. Comments on some introduced ~ Gaps of Ohio and nearby states. Michigan Bot. 33:95— THIERET. oe Comments on the ie of the south- pastern cana States. Sida 13:14 Cas STEYERMARK, J.A. 1963. Flora of ae The Iowa State University Press, Ame PE. Hyatt, and C. AMASON. 1991. Additions to 19-156. Tuomas, R.D., E.B. SmirH, E. SUNDELL, the flora of Arkansas. Sida 14:482—491. TuRNER, B.L. 1996 [“1995"}. Cerastiwm texanum (Caryophyllaceae) does not occur in Texas Phytologia 79:356—363. Stpa 17(4): 845. 1997 IN MEMORIAM LYNN LOWREY (1920s?—28 June 1997) Courtesy: Native Plant Society of Texas. Native Pl. Soc. Texas News 15(5), September/ October 1997 REMEMBERING LYNN LOWREY by Dr. David Creech Lynn’s passing June 28, 1997, was a hard blow to the gardening world. Another mighty oak has fallen. A living legend is gone. For many of us Lynn was one of those folks you run across in life you want to end up like. I knew Lynn from the late 60s as a TAMU Horticul- ture major working in the fruit science end of the program and, as a graduate ae at TAMU, I panes trails with him many times in the 70s. I was then looking for various native and exotic fruits adapted to East Texas conditions (pawpaws, blueberries, hawchornes, jujubes, etc.) and Lynn was the man to see. From TAMU to SFA and then the Arboretum and suddenly Lynn’s footprints and mine crossed much more often. Trips to Mexico, to the backwoods, to nurseries, to gardens or just out and about with other plant friends was part of the plan. Simply put, from the beginning, Lynn struck me as a genius with character. For many of us, Lynn really was a legend in his own time. His impact on southern landscapes reaches almost as far as his effect on people. I could never decide if Lynn was born 100 years too late or too early. Both fit. Always a quiet reservoir of plant knowledge, always eager to learn and share, Lynn was blessed with humility and great humanity. His expertise was unquestioned; whether it was a botanist, horticulturist or adventurous land- scape designer visiting the state, Lynn was always high on a list of people to see. Lynn was more a true plantsman than a true nurseryman. Hard for him to stay in one spot, hard fo one to make a firm appointment with, Lynn liked to wander here and there collecting seed or plants. He enjoyed connecting with someone who knew about this or that plant or that person who was working with Throughout his career—and chee s no point in recounting all the names and locations of Lynn’s nursery haunts—Lynn brought adventure and excitement to the nursery and landscape industry in the south. The Mexico oaks are only che tip of F the iceberg. Many gardens across the south are blessed with ie memorials to the oh-so-many plants that Lynn showered on the world. Who can’t appreciate Lynn’s big leaf yaupon? The Arboretum’s Lynn Lowrey American Holly selections—cuttings grown from superior East Texas trees— are now robust 10’ females with large berries and clean foliage ore’s chat special Oakleaf hydrangea sporting large tresses and good fall color over by the seen, border; it deserves a name—collected by Lynn near Angola Prinson in Lout- siana. Lynn felt that the Wisteria frutescens, our native Westeria—white and blue forms-—— e Japanese and jem were sae ‘of greater use and certainly easier to manage than those ruc chinese cousins. Hmmm...what about the soapberry with exceptionally narrow leaves? What about the Hersprale from Mamuluke pass? The Mahonias? For those of us who knew him, the list never ends. And that is what Lynn would want us to think about him when we thought ie him—the plants Lynn’s impact amazes many of us academic and non-academic types who write and speak for a living in the amazing gardening world of Texas. Lynn wrote little and spoke Stipa 17(4): 847. 1997 848 Sipa 17(4 c= ) less. Yet, his followers are legion. The power of his message was more than just the plants. It was an atcitude to live by. Before it was politically correct, Lynn spoke and wrote quietly on biodiversity, taking advantage of microhabitats, and understanding the importance of site preparation, plant community development, natural form in design, using superior and adapted natives and exotics. On day Lynn and I were working along a fence row south f Navasota off a red clay road, collecting seed and cutting from a bright orange-berried, deciduous holly that mixed into a swarm of native trees and shrubs and herbaceous and God knows what. Lynn said that he didn’t see how you could much improve on where we were at—kind of God's design. While Lynn was no doubt an adventurous gardener, he didn’t stop there. He also be- came a role model to live by. L always felt chat business to Lynn—even money—was just a distraction, something that had to be deale with as little as possible. Down deep, he'd rather give away plants than sell them. Lynn's goal in life was to work at what he loved to do and share it with the world. Lynn did that better than anyone. Perhaps more important than the plants or the gardens, maybe Lynn’s greatest talent was in putting people to- gether. Whether connecting medical researchers and cancer foundations with rare cancer- fighting plants like the Happy tree of China, or locating Taxus globosa in the mountains of Mexico for the Arnold Arboretum, or hunting down a particular genotype for a botanist, or chasing down a natural hybrid between two related genera—Lynn was always there to help out—to make something happen. Amazing to me, he never thought much about all this. He seemed cto think he wasn't really doing all that much, that everyone else was doing a lot of good or that others work was “really important.” That’s what Lynn saw in people—the good. I think all the people, plants and gardening did that to him. Matt Welch, one of my students, gave me this quote the day after the Lowrey memorial and the message kind of stuck. “Maybe a garden is not like an artistic creation, but more like a relationship. It is not so much designed as it is nurtured and managed. Imperfections, ity to it, and that Laas inconsistencies, and incongruities say that the relationship has some vita the gardener is engaged in a dialog and doesn't need to have the last word.” —Tom Little, 1996. Lynn Lowrey never even intended to have the last word; his gardens, his plants and his people will do it for him. See also: Bisbee, B. 1995. The Lynn Lowrey Legacy. Native Pl. Soc. Texas News 13(5):5—6. Sipa 17(4): 848. 1997 IN MEMORIAM Lynn Lowrey r). (cente Benny Simpson 1997 Stipa 17(4): 849. IN MEMORIAM (29 February 1928—27 December 1996) Courtesy: Native Plant Society of Texas. Native Pl. Soc. Texas News 15(2), March/April 1997 PIONEER OF THE Native PLANT MOVEMENT by Jill Nokes Last night a stranger called me. He was a tree trimmer by trade, and had been thumb- ing through A Feld Guide to Texas Tree by Benny Simpson, saw my name listed with others in the foreword, and called me on impulse because he wanted to know more about this man Benny. “How does he know that this particular tree is only found on gypsum out- crops? I want to find ouc more about him. Where does he live? Do you think he'd let me follow him around some day when he’s out lookin’ at trees?” When he finally paused, | once again had the sad task of telling someone that Benny had died on Friday, December 27 of complications following heart surgery. The tree trimmer man had missed knowing a most generous and special perso a) Benny was born on the leap year of 1928. Last year we joked that he was only 17 years old instead of 68, since his birthday only rolled around every four years. He was born in Northfield, in Motley County, to parents Una and Baker Simpson, who were ranchers in the rolling High Plains part of the state that always held a special place in Benny’s heart. He was one of 15 students in his graduating high school class and attended Texas Tech college at 16, majoring in agronomy and farm machinery. Later he joined the marines, barely missing action in the Korean War. He began working at Texas A&M University Research and Extension Center in Dallas in 1954 when it was the Texas Research Founda- tion, a private agricultural organization, Texas A&M took it over in Early in his career Benny was in charge of planting crops that were being studied by other researchers at the Center. Although his job required him to supervise years of futile cotton root rot studies on the Blackland Prairie, the passion and interest for native plants and the Texas landscape he had since his days as a boy on the ranch began to take shape. On his own in the 1970's he began to collect and plant various native woody plants and find left over plots at che Center to establish field trials for some of these specimens to see how they would adapt to a situation outside their normal range. In the early days, he had to C efend his work and ever-growing arboretum from being plowed under each season, be- cause at that time Texas A&M did not see the value in studying native plants. By the time of his death Benny had finally received some of the recognition for his vision- ary and steadfast work. His research led to the selection of nine superior selections of native plants released to che nursery industry (see accompanying article by Tim Kiphart). As he neared retirement, there was concern about how best to preserve his arboretum, which was by then very substantial. In a tribute to his work, plants from his collection were donated to the Heard Natural Science Museum in McKinney, Texas. A plant collection of 345 plants in 84 genera representing more than 20 years of his research was also established in his name ac the Dallas Horticultural Center at Fair Park. And finally, at the Research and Extension Center, where he’d worked for 42 years, plans began last fall to establish che Benny Simpson Outdoor Learning Center. The Center is situated in the lovely wooded area which had been part of his field trials. Apart from these gardens, Benny was also the recipient of many awards from garden clubs, universities and professional organizations. Sipa 17(4): 850. 1997 851 Perhaps Benny's greatest legacy was the influence he had over a whole generation of horticulturists, botanists, plant enthusiasts and finally, the TAMU System itself which pce supporting more research into the us and development of one of our state's most oked resources. A big factor motivating Benny's research was his understanding (as pera only one whe had lived on a ranch knows) that scarcity of water will be one of the b st challenges in Texas’ future. Today, especially after the drought of 96, the existence ae water rationing and the acceptance of xeriscape landscape programs, the public 1s be- ginning to understand what was always obvious to hin nny was one of the founders of the Native Plant Society of Texas in 1982 and was very active in the organization until his death. He donated royalties from his book, A Freld Guide to Texas Trees, to the Native Plant Society of Texas. For several years he underwrote the cost of the cover photography on the Proceedings of the NPSOT annual meeting. It was his idea to initiate the annual awards and to rotate the location of the annual meeting each year to a different vegetational region in order to increase awareness of the spectacular variety of our native flora. Benny worked hard to organize the wonderful program we had last October in El Paso. Many thought ic was our best annual meeting yet and all of us were disappointed when he could not attend. I was a bewildered graduate student in 1973 when I first met Benny. I had a shoebox full of 3x5 index cards with notes scrawled about propagation and use of a random number of native woody plants. I had a vague idea that gathering as much of the scattered informa- tion from books and oral sources about these plants would be useful somehow, but I had no scientific background and no clue how to go about it. From the beginning Benny saw the value in this project beyond what I had imagined. He supported me and instilled confidence. He offered all the knowledge that he had painstakingly accumulated throughout the years. He encouraged me as I slogged through a GOO page masters thesis, which eventually became a book. He read both manuscripts and let me know that he was very proud of me. He was the kind of friend who can see what you can accomplish and has faith in you even when you have trouble believing in yourself. We all made the common mistake of assuming we would have Benny with us forever. Benny was working on a book on native shrubs, as a companion to his tree book. As of now we don't know how close it was to being finished or if we will have a chance to benefit from all che work he put into ic. Sally Wasowski has about a dozen more books to write and was depending on having the help he had always offered on her other projects. Mary Anne Pickens, as president of NPSOT, was counting on his guidance beginning her new term. We assumed he would shape the program for NPSOT’s annual meeting in Uvalde next October, highlighting one of his favorite parts of the state, the Rio Grande Plains. Addi- tionally, Dr. Ted Davis and Dr. Wayne MacKay intended to video tape Benny touring his dics, his- — arboretum hoping to capture information about his plants, their provenance, ha tory and cultivation he held in his he Benny’s legacy to us is still naitoldinis and being revealed. Just as the plants in his gardens grow, so will his influence on new students and researchers. It’s clear that even the general population won't go back to being indifferent about what landscape plants they will use. For me, my consolation comes everytime I look at one of his photographs (or see someone planting a shrub he selected and grew) and I hold that picture in my heart of him standing next to the plant che first time he saw it in the field. He is wearing a beat up straw hat, a white shirt with suspenders, khaki pants scuffed into Justin boots and a big smile on his face. Thank you, Benny. You gave us so much. Thank you and farewell Note: Benny is survived by his mother. two sisters. a brother, four nieces and one nephew. The Sipa 17(4): 851. 1997 852 Sipa 17(4) family asked that memorials be made in Benny Simpson's name to the Native Plant Soc iety of Llexas. 9 A memorial service to celebrate Benny's life was held on February 8 at Texas AGM University Research and Extension Center in Dallas. BENNY SIMPSON’S LEGACY TO NPSOT by Mary Anne Pickens Benny Simpson was a founding fat — yer and former president of NPSOT. He ne define the goals of NPSOT and wet the organization in motion. He served as the Master of Cer- emonies of our annual meetings. He put together our symposium proceedings. He had a clarity of vision that he didn’t mind sharing. He kept us on cot a And what ts the course? Please refer to his President’s Corner of September/October 1994, Benny would noc like for me to take up Coo much space quoting him, but he spoke of fiscal responsibility as being the Board of Direccor’s number one task. Second on his list for us was an “attractive, informative newsletter.” Number three was “our annual State Meer- ing at which the flora of one vegetational area” is studied each year. Fourth on his list of duties was “to conduct the quarterly business meetings with meaningful decorum and dispatch.” Upon becoming president of NPSOT he had made three proposals: (1) Seek a memoran- dum of understanding with Texas Parks and Wildlife with the view of selecting certain parks, conducting plant surveys of those parks and publishing a vascular plant inventory; (2) Each Chapter should work up a vascular plant inventory of garden worthy native plants within a 50-mile radius of that chapter; (3) Using the above list, each chapter should plant asmall flower garden or arboretum By the end of his term, he said, “Proposal | is out ... too time consuming ... we are not really qualified to do. Proposal 2 should be easy and a lot of fun. Proposal 3 hopefully youll chink about and one or two chapters may do it. So where are we on Benny’s proposals? We altered Proposal No. |, and with Tim Kiphart’s diplomatic skills simply signed a two year memorandum of agreement (MOA) with Texas Parks and Wildlife encouraging continued or increased NPSOT chapter members’ partici- pation in TPW D's pone stewarding native plant resources. Proposal 2 1s still easy, but how many of us have done it? If your chapter has done it, has the chapter filed the inventory with the NPSOT state office so that it is a matter of public record? Proposal 3 should be one of our memorials to Benny. With che stimulus provided by the Wray Foundation in our mini-grant program to encourage native plane demonstration gardens, we can move forward. Recognizing Benny's legacy to NPSOT, David Todd, board member of the Wray Foundation, asked that this year’s grant money be recognized as a memorial to Benny Simpson. I suspect Benny would say, “Great! Seart planting those gardens—with natives found within a 50 mile radius!” I don't always see things as clearly as Benny did. Black and white frequently blend into shades of gray for me. But for clarity of vision, I always go back to NPSOT’s mission statement: to promote the conservation, research and utilization of the native plants and plant habitats of Texas through ec — a cation, outreach and example. Benny recognized two ways to clearly accomplish the mission—produce a first class newsletter and provide an outstanding, stimulating, ever challenging annual symposium. On a state level, we do those two things. But through our local chapters there are many SIDA : 7(4): 852. 1997 IN MEMORIAM 853 ways of accomplishing the mission and meeting the objectives. Brazos Valley meets it when it collects acorns to propagate trees for Mother Neff State Park. Houston, Mercer, and Dallas Chapters meet it when they man a booth at the TAN show. Trinity Forks meets it when it restores a wildflower meadow in a burned area within a city park. Bandera Chapter meets it when it gets the Highway Department to designate an area as a Natiy Area not to be mowed. Kerrville Chapter meets it when it installs and maintains a native garden at che public library. Columbus Chapter meets it when it does the same in Beason’s Park on the Colorado River. Austin Chapter meets it when it has a booth at Eco Farr. Within each chapter lies the seed of potential growth for meeting NPSOT’s goals. I chink our mission is quite simple—plants. We study them, we encourage the research — on them, we respect them, we promote their conservation. We encourage others to do the same. If we carry out this mission, we do not need to waste time and money on anything e. Keep Benny in mind. Clarity of vision is something we all need. I will work on conducting “quarterly business with decorum and dispatch!” BENNY SIMPSON’S PLANT LEGACY by Tim Kiphart Combine botany and horticulture and what do you get? The life of a research horticul- turist. Our guest: Make Mother Nature becter and look for the botanical oddities she produces. It’s what we thrive on! Plant selection takes a keen eye. Benny Simpson worked on that for 45 years. Working for the same place your whole life - how bizarre. The things you do for love! Benny was responsible for introducing about a dozen plants into the nursery trade. He looked for qualities that made a great plant even better: form, color, Hower size, resistance (insect, disease, heat, cold)—you name it. All the selections Benny made were of western (Trans-Pecos) species. This is where he felt most at home. The most well known of Benny's introductions are the Lewcophylums - the “Cloud” se- ries. In almost every part of the state someone has planted Texas an sage. It certainly is the evergreen backbone in the western half of Texas. Two selections of L. frutescens, two of i. tandidum and a hybrid species all show Benny's thumbprint. Leacophy lien frutescens, “Green en leaf form of this typically gray leaf shrub. It is much more tolerant of irrigation—a oe plus for the nursery industry. Millions of “Green Cloud” have sold. Be- cause of the atypical white fower “White Cloud” is unique. You can imagine how choice the 5-6! shrubs. oe porcelain white flowers look against the gray foliage. Both selections are large A stunning shrub in the wild, L candidum, is difficult to grow in a landscape. Dwart silverleaf sage is about 3' tall and has iridescent silver foliage. In addition, its grape col- ated flowers make an awesome landscape display. The two selections of L. candidum are ver Cloud” and “Thunder Cloud.” They are respectfully named for their exceptional foliar quality and explosive bloom The last of the series is “Rain Cloud”, a hybrid between L. minus and L. frutescens. It caught on more in Arizona than Texas. What ashame! It has the height of L. /ratescens and the darker | to rain it produces masses of fay purple flowers of L. minus. Responding very w low was another species Benny made selections on. This shrubby tree is na- listed in nursery Desert wil tive to arroyos but has proven quite adaptable in the landscape. It is even catalogs from the 1860's, making it a native heritage plant. Chilopsis linearis, “Dark Knight’, is purple rather than the typical pink and “White Knight” is a fine, pure white, showing a yellowish throat. Sipa 17(4): 853. 1997 854 Stipa 17(4) nother multi- trunkec I shrub, Amorpha fraiteosa var. angustifolia,’ ‘Dark Lance,” is a much underutilized plant. The bloom is reminiscent of a fox tail, but displays dark indigo- purple with or He anthers. It is a wonderful habitat plant, not to mention a big hum- mingbird attraction. Benny was never much on perennials. He was a manly man! But, why then did he love frog fruit? Two perennials come to mind though—one official and one non-official, both Salvias. Officially speaking, Sa/ria regla,* Mount Emory” is unsurpassed. Not only does “Mount Emory” have more cold tolerance, the leaves are a darker, more lacquered green. The other perennial Benny spread around was S. grege//, “Lancaster Hill” (not official). A ee bloom with a white throat sets this gregg// apart. It’s like a beacon shining in the landsca Other ae Benny was working on include Mexican buckeye, Evergreen sumac, Mexi- can redbud, Wax myrtle and a hybrid Kidneywood. These plants are still under evaluation. The plant introductions were only the tip of the iceberg. Benny was responsible for turning people on to many more aan than he introduced. He loved his plants and it was this love that propelled him through lite. Well, my friend, may you forever walk the desert of your dreams, BOONDOCKING WITH BENNY by John Mac Carpenter Ben else in the history of the state. He explored every corner of it with his friends, who counted themselves lucky to be along. Many of you I know, but many I don't. Our common friends included Mary Buchanan, an uncommon woman, Dean Ricer and Mark Rosacker of New Mexico's Living Desert State Park, John White, formerly of El Paso, and Wayne MacKay, currently of El Paso, and most important of all, Dr. Barton H. Warnock (Doc) of Alpine. Benny and Doc and I spent many precious days together studying and collecting plants, ry J. Simpson was almost certainly familiar with more of Texas Wild than anyone walking desert washes, climbing hillsides, checking along watercourses. The gypsum sand dunes west of the Guadalupes in Culberson county were one of our most spectacular visits. On one memorable visic Benny and I met Wayne MacKay, Dean Ricer and Mark Rosacker at the Lynch Ranch offices in Dell City co pick up the key and get directions. It drizzled all day, but the temperature stayed in the upper 70's so it was comfortable. Mountains surround the broad plain containing the dunes, El Capitan just to the east the most spectacular, but the Cornudas being the most inte resting looking. The snowy white sand was a perfect backdrop for the rosemary mint, supposedly Poliomintha incana but reaching five feet call. In the moist air, the scent of the leaves was delicious. Benny kept saying “I could live here.” For peace and beauty, these dunes are near the top of the lise. Benny always called when it had rained. He knew spring comes in the Trans-Pecos whenever rain comes. He'd arrive about three weeks after a rain and we'd go looking. picked him and Sue Tracy up at che airport once in Iran and we went to Lancaster Hil overlooking the Pecos River in Crockett county. This is where Hill country meets Big Bend country. This is a prime site for cliff Fendler bush, one of Benny’s favorites. From this spectacular spot you can look across old Fort Lancaster into che burned mesa country chat [ love with its crooked streak of green in the foreground that is the river. Benny and Doc and I checked out Mohr oaks and Mexican redbuds on the launch in Terrell county while smelling the sweet scent of the algerita in full bloom. Some of th Sipa 17(4): 854. 1997 IN MEMORIAM 855 Mohr oaks along Big Canyon are true trees instead of small shrubs. He found desert ceanothus in Pinto Canyon in Presidio county, Quercus polymorphy on Dolan’s Falls Ranc! on the Devil’s River, roses in the Guadalupe and Davis Mountains, and cenizo that no one else could propagate, he moved from the dry hills of the Trans-Pecos to the nurseries and to our yards. A favorite place in the western part of the state for us is Mesquite Ranch on tl west slope of the Chinati Mountains in Presidio county. Doc and Benny and I spent many days there, walking hillsides, canyons, arroyos and climbing up Chinati peak. San Fran- cisco is the prettiest place, with its Thurber penstemon and frequent water. All the washes V. angustata, and canyon re south- and canyons on the ranch are lined with esperanza, Tecoma stans, senna, Senna wislizenii, which is spectacular in bloom. There is also zona cockroach plant, asa crooksii, smaller but just as pretty and poisonous to cockroaches. The Pelillas arroya are intermediate between Juglans major and Juglans microcarpa. walnut trees 1 visiting with Al Real, ¢ We were eating lunch one day at the ranch headquarters and foreman, while Benny told us about his new hearing aid. We'd noticed he was hearing better than usual and he told us how it differed from earlier models. When he finished I asked him what kind it was and he, pulling up his cuff, said, “A quarter till two Benny was always the emcee at our annual meetings until 1996 when he didn’t feel well enough. For a field hand, he cleaned up real good and was as much at home at the podium s ina swamp, a woodland or a desert wash. He looked a lot better too and could charm a garden club or educate scientists Benny left us a lot to work with and a lot of memories. Most of us are growing at least one of the plants he moved from nature to nurture and will always have something of him around, I regret the days I had expected to spend with him in the breaks along the edge of the Caprock when he retired to home in Motley county. Una, you gave us a winner and we appreciate the gift. In loving memory, John Mac Carpenter Courtesy: Dr. Tim Davis, Texas A&M University Research & Extension Center Texas A&M University System, Dallas, Texas. BENNY J. SIMPSON Research Scientist Texas AGM University Research & Extension Center Texas AGM University System Dalids, TX J 3252-0599 B.S.: Texas Technological University (Agronomy and Farm Machinery) 1954-72: Junior to principal soil scientist farm superintendent, Hobilizelle Agricultural Laboratory, Texas Research Foundation, Re 1972-76: Farm manager, mins a altel Experiment Station 1976-80: Research scientist, farm manager, Texas Agricultural Eyecare Station 1980-1996: Research Scientist, Texas Agricultural Experimenc Station. Project H-8164 “Resource Efficient Plants for Texas and the South West” development of genetically superior ornamentals for Texas. This mission 1s appro ached in two ways: is concerned with the by cooperation with other scientists in testing their material and by basic researc h on new plants not now in the nursery trade. The major thrust is with superior selections of Texas native plants that will hese plants are collected in the wild (seeds or be low users of fossil energy and water. Propagules of t lapeation. While undergoing adaptation cuttings), increased to greater numbers and tested for ac Sipa 17(4): 855. 1997 856 Sipa 17(4) studies, further cuttings and seed are taken for propagation, fertility, consumptive water and other investigations. HONORS AND AWARDS The Presidenc’s Award, Texas Woman’s University Tenth Annual Texas Wildflower Day; Donovan Stewart Correll Memorial award in che field of scientific writing for the book, A Field Guide to Texas Trees. The Silver Seal Award, National C ounci] of State Garden Clubs, Inc.; Charles Woddle Wile baie Distinguished Lecturer, Texas Tech University CAREER MAJOR ACCOMPLISHMENTS Release of Plant Varieties and Other Genetic Materials: Leucophyllum frutescens, “White Cloud’ Ce Lencophyllum candidum, ‘Silver Cloud’ Cenizo. LencophyHum frutescens, ‘Green Cloud’ Cenizo. Chilopsts linearis, Dark Storm’ Desert Willow. Chilopss linearis, White Storm’ Desert Willow. Salvia regla, ‘Mount Emory’ Mount un — re ‘Renner Weeping Lovegrass. (J.C. Read, oo and L.J. Streetman Leacophyllum minus x L. frutescens, R, ain C ee Hy i Leucophylum candidum, ‘Thunder Cloud’ Cenizo. Anmorpha fruticosa var. angustifolia, Dark Lance’ False Indigo. id Cenizo. PUBLICATIONS Simpson, B.J. and B.W. Hipp. 1985. The eee and transplanting of the Texas Madrone. Proc. Texas State Hort. Soc. Conf. Hipp, B.W. and B.J. Simpson. nau Nitrogen requirements of the Texas Madrone. Proc. Texas State Hort. Soc. Conf. 1:3-5. Hipp, B.W. and B.J. Simpson. 1986. Influence of sulfur on soil pH and growth of . Environ. Hort. 4(4):142-144. Hipp, B.W., B.J. SHnpson, and P.S. Graff. | cenizo, t aos] 1988. Influence of nitrogen and phosphorus on growth and tissue N and P concentration in Salvia Breggil. J. Env. Hort. 6(2):59-61 oe B. BJs ‘&B. W. Hipp, and E.L. McWilliams. 1989. “Thunder Cloud’ cenizo. Horse 24:17 See B. BJ. = W. Hipp, and E. = i Williams. 1989. ‘White Storm’ and “Dark Storm’ dessert willow. HortSci. 24:17 Simpson B. . me W. Hipp, and E . ees 1989. ‘Dark Lance’ false indigo. HortSci. 24:713-7 Hipp, B.W., 7s Simpson, and P.S. Graff. 1989. Influence of phosphorus on nitrogen fertilizer requirements of Melampodium lencanthum (blackfoot daisy). J. Environ. Hort. 7:83-85 Read, J.C. ane B.J. Simpson. 1989. Documented plant chromosome janis 1989:2, C oo counts of three species in the genus Leacophyllum Sida 13 eras B.J. 1 ). The modern urban blacklands, from ae prairie to soccer fields. Blackland eemen Baylor University, Waco, TX. In Pr DOT, Simpson, B.J. 19 Symphoricarpos occidentalis (C Say cee ae), new to Texas. Sida 14 2-513. Read, J.C. and B.J. Simpson. 1992. Documented chromosome numbers 1992:3 mentation and notes on the distribution of Melica montezumae. Sida 15:15 1—1 52 Simpson, B.J., J.P. Karges, and J.M. oo Texas and the United States. Sida | aaa . Docu- 1992. Quercus polymorpha (Fagaceae) new to jon SIDA 17(4): 856. 1997 IN MEMORIAM Technical Articles ee B.J. 1980. Check list of the native trees of Texas. Texas Almanac 1980-1981. » Edition. eee R.L., B.J. Simpson, and A.B. Hamon. (agarito, red barberry, and Texas barberry) are attacked pe a whitefly, Tetralenrodes visorum (Homoptera: Aleyrodidae). SNA Res. Conf., Atlanta, Crocker, R.L., B.J. Simpson, and C.L. Simpson. 1981. Insect problems of native woody ornamentals under cultivation. SNA Res. Conf., Atlanta, Hipp, B.W. and B.J. Simpson. 1981. Influence of N on oor and tissue N concentration on Lencophyllum candidum (violet silverleaf, cenizo). Commun. Soil Sci. Plant. Anal. 12:205-209 Simpson, B.J. and B.W. Hipp. 1982. Influence of nitrogen on rooting of Leacophyllum candidum cuttings. Plant Propagator. 27(4):10-11. Crocker, R.L., C.L. Simpson, B.J. Simpson, and A.B. Hamon. 1982. Vulnerability of six native ornamental Sephora to mealybug, Phenacoccus sp. near : madierensis (Homoptera: ee ina cies SNA Res. Conf., Atlanta, Crocker, R.L., C.L. Simpson, B.J. Simpson, and AB. Aine 1982. Brown soft scale Coccus hesperidum and ee fruite lecanium, parthenolecaninm corni (Homoptera: Coccidae), attack native Sophora under cultivation. S onf., Atlanta, GA. Simpson, B.J. 1983. Native plants - release of new varieties and future prospects. Proc. Field, Container and Bedding Plant Seminar. Simpson, B.J. and B.W. Hipp. 1984. Drought tolerant Texas native plants for amenity plancing. P. 483-500. In M.A. Collins (ed.). Water for the 21° Century: will it be there? S.M.U., Dallas. Crocker, R.L. and B.J. Simpson. 1985. Larvae of io moth, Awtomeris io (Lepidoptera: ae), infest panicled amorpha, Amorpha ae wlata (Fabaceae), under evaluation 1981. Three native, potential ornamentals Saturniid for use in perce SNA Res. Conf., Atlanta, Crocker, R.L. a . Simpson. 1985. Spirea aphid, A Ve einai stera: Aphididae), F | | | P and anon a Clastoptera xanthocephala var. unicolor (Hlemopters: Ceti pina’), eae mexicana (Asteraceae), a native shrub under evaluation for domestication. SNA R conf., Atlanta, GA. Simpson, B.J. and B.W. Hipp. 1980. oe Texas flora with urban oP control poten- tial. Proc. XVI International Erosion Control Assoc., Dallas. Pp. 143-151. Hipp, B.W. and B.J. Simpson. 1986. Nitrogen nutrition of container grown prairie plants. I. Phus. | X North American Prairie Conf., Denton. 26.( 3 Smith, L.A. and B.J. Simpson. 1986. Amenity landsca ping in sth prairie environ- ments. Proc. X North American Prairie Conf., Denton. 26.07: pp. Simpson, B.J. and B.W. Hipp. 1986. Nicrogen nutrition of container grown prairie plants. Il. Prunus. Proc. X North American Prairie Cont., Denton. 26.08: pp. 1986. The oaks of Texas. J. Arbor. 12:302—3034. and B.J. Simpson. 1987. New pests of ornamental shrubs native to the - damianita , Meh ean meet SNA (In Press). Morgan, D.L. and B. a Simpson. 1989. New plants for western landscapes. Proc. SNA Res. onf., Atlanta, Simpson, B. J. 19 meridian. Proc. ive Ann. Meet. S. Reg. IPPS. Morgan, D.L. and B.J. Simpson. 1990. Native grasses for western landscapes. Proc. SNA Res. Conf., Atlanta, GA. : eee native Texas woody ornamentals found west of the ggih Sipa 17(4):; 857. 1997 858 Sipa 17(4) onl Hipp, B.W., M.C. Engelke and B.. faa 1991. Use of resource efficient landscape plants co reduce urban runoff. P. 183-190. In: R. Jensen, ed. How healthy is the upper Trinity River? The Texas Water Resoiarees ae College Station oe BJ. 1991. A preliminary annotated checklist of the woody flora of Lancaster Hill, Crockett Country, Texas.,, In: The Edwards Plateau and its many ecosystems. Proc, NPSOT. Kerrville. Pp. 71-81. sce B.J. and S.C. Tracy. LOOL. Rediscovery of Fendlera rupicola in the Edwards Plateau. : The Edwards Plateau and its many ecosystems. Proc. NPSOT. Kerrville. Pp. 49-58. Abstracts Hipp, B.W. and B.J. Simpson. 1979. Influence of cropping system on fertilizer response on a blackland prairie soil. Amer. Soc. Agron. Hipp, B.W. and B.J. ie 1982. Influence of cropping system on fertilizer response by grain sorghum and on chemical and physical properties of a vertisol. Amer. Soc. Argron. Giordano, C., B.W. Hipp, and B. Simpson. 1984. Water use by buffalo grass and Sc. Augustine grass in North Texas. Amer. Soc. Agron. Smith, L.A. and B.J. Simpson. 1986. Amenity landscaping in southern prairie environ- ments. Abstr. Tenth N Amer. Prairie Conf., 26.06 Hipp, B.W. and B. - Simpson. a Nitrogen oe of container grown prairie plants. I. Rhus. Abstr. Tenth N. Sm. Prairie Conf., 26.( Craeccn. B.J. and * W. Hipp. ne Nitrogen nutrition of container grown prairie plants. Il. Pranus. Abstr. Tenth N. Amer. Prairie Conf., 26.08 Colbaugh, PE, S.L. Fields, and B.J. Simpson. 1986. iy mildew resistance among native populations of cedar elm. Abstr. Amer. Phytopath. Texas Research Foundation Publications Laws, W.D. and B.J. Simpson. 1959. Grain sorghum in say systems for the Black- lands. Bull. 8. Hob. Agric. Lab., Tex. Res. Found., Renner, TX. Laws, W.D. and B.J. Simpson. 1960 . Framing systems for cotton production in the Black- lands. Bull. 9. Hob. Agric. Lab., Tex. Res. Found., Renner, TX. esac B.J. 1964. wines production in framing systems for north central Texas. Bul 19. Hob. Agric. Lab., Tex. Res. Found., Renner, TX. Simpson, B.J. 1968. Grazing and management studies on NK-37 bermuda grass. Bull. 6 aay Lab., Tex. Res. Found., Renner, TX. Laws, oe ., BJ. Siccon: and H. Brawand. 1970. Wheat eed test at Rennes 1970, Renner Res. Rep. 1(7): Hob. Agric. Lab., Tex. Res. Found., Rent Brawand, H., B.J. Simpson, and W.D. Laws. 1971. Plant ae ne dry matter odacn aa in peep Se s for the dex xas Blacklands. Renner Res. Rep. 1(8): Hob. hare Lab., Tex. Res. Found., Renne Simpson, B.J. and H. ae ee 1971. Wheat variety test at Renner Rep. 1(9):Hob Agric. Lab., . Res. Found., Renner, Simpson, B.J., J.R. Birchett, ‘LL Brawand. 1972. Wheat variety test at Renner 1972. Renner Res. Rep. 2(2): Hob. Agric. La., Tex. Res. Found., Renner, TX. 971. Renner Res. - Texas Agricultural Experiment Station Publications Whiteley, E.L., B.J. Simpson, and §.H. Whitehurst. 1976. Short-season, narrow-row cot- ton studies in the northern Blacklands. Tex. Agric. Exp. Sta. R-3382 Sipa 17(4): 858. 1997 IN MEMORIAM Bo? Simpson, B.J., D. pes and H.J. Walker. 1976. 7 sorghum hybrid performance Dallas, Texas - 1976. Tex. Agric. Exp. Sta. PR-3426- Simpson, B.J. 1977. a Bae yard garden - 1976 - 1977. Tex. Agric. Expt. Sta. RCIR —_ No. 77- Simpson, B.J., D. Pietsch, and H.J. Walker. 1978. Grain sorghum hybrid performance Dallas, Texas—1977. Ex. Agric. Exp. Sta. PR-3478-6. Niles, G.A. and B.J. ee bee Performance of cotton varieties, 1974 - 1977. Tex. Agric. Exp. Sta. RCT . 78-3 Whiteley, Bie Es... oo and S.H. Whitehurst. 1979. Narrow-row cotton in the northern Blacklands. Tex . Agric. Exp. Sta. MP-1410. Simpson, B.J., D. Pietsch, aint HJ. Walker. 1979. Grain sorghum hybrid performance Dallas, ne 1978. Tex. Agric. a Sta. PR-3540. Simpson, B.J., D. Pietsch, and H.J. Walker. 1980. Grain sorghum hybrid performance Dallas, Texas - 1979. Tex. Agric. Exp. Sta. PR-3661 Read, J.C. and B.J. Simpson. 1981. Renner Weeping Lovegrass. Tex. Agric. Exp. Sta. 1803. Simpson, B.J. 1983. ‘Mount Emory’ Mountain Sage. Tex. Agric. Exp. Sta. L-2057. Simpson, B,J. 1983. ‘Green Cloud’ Cenizo. Tex. Agric. Exp. Sta. L-2058. Simpson, B.J. 1983. ‘White Cloud’ Cenizo. Tex. Agric. Exp. Sta. L-2059. Simpson, B.J. 1983. ‘Silver Cloud’ Cenizo. Tex. Agric. Exp. Sta. L- 2060. Simpson, B.J. 1985. ‘Thunder Cloud’ Cenizo, violet silverleaf. Texas Agric. Exp. Sta. 2174 Simpson, B.J. 1986. ‘White Storm’ Desert Willow. Tex. Agric. Exp. Sta. L-2188. Simpson, B.J. 1986. ‘Dark Storm’ Desert Willow. Tex. Agric. Exp. Sta, L-2189. Simpson, B.J. 1986. ‘Rain Cloud’ Hybrid Cenizo. Tex. Agric. Exp. 2190. Simpson, B.J. 1986. ‘Dark Lance’ False Indigo. Tex. Agric. Exp. Sta. al Hipp, B.W. and B.J. Simpson. 1987. Thirty-five years of farming systems ere in the Texas Blacklands. Texas Agric. Exp. Sta. Bull. 1604 Hipp, B.W., T.C. Knowles, and B.J. Simpson. 1992. Soils of the Texas Agricultural Ex- periment Stations at Dallas and Prosper. Texas Agric. Exp. Sta. College Station. Bull 1705. Books ae B.J. 1988. A field guide to Texas trees. Texas Monthly Press, Austin. 372 pp. + xi + 113 colored plates (maps and photographs). Popular Articles Simpson, B.J. and J.C. Read. 1976 Cenizo. Texas Parks and Wildlife 34(10):24—27. Simpson, B.J. 1976. An oak from an acorn. Texas Wildflower Newsletter 1(1):1 1-12. Simpson, B.J. 1977. The Texas madrone. Texas Wildflower Newsletter 1(2):7-8. Simpson, B.J. 1977. The Renner collection. Texas Wildflower Newsletter 1(3):6—7 Simpson, B.J. 1977. The silver-le afs. Texas Wildflower Newsletter Le Simpson, B.J. 1977. The Ashe juniper. Texas Wildflower Newsletter 2( Simpson, B.J. 1978. The glorious maples of Texas. Texas Wildflower Weysie ceer ’ 2):5 Simpson, B.J. 1978. The sumacs of Texas. Texas Wildflower Newsletter 2(3):5— Simpson, B.J. 1978. The Texas pistachio. Texas Wildflower Newsletter 2 4):1 1-1 Simpson, B.J. 1978. Silverleaf mountain mahogany. Texas Wildflower ey 3( 1): 9-10. Simpson, B.J. 1979. The desert willow. Texas Wildflower New rslecrter 3(2):5 Simpson, B.J. 1979. Mesquite. Texas Wildflower Newsletter 3(3):8-9. ~ = is | rar bo Sipa 17(4): 859. 1997 860 Sipa 17(4) Simpson, B.J. 1979. Wild olive. Texas Wildflower Newsletter 3(4):7. Simpson, B.J. 1979. Redbuds of Texas. Texas Wildflower Newsletter 4(1):5—6. Simpson, B.J. 1979. Attractive Texas Natives. The Texas Nurseryman 10(3):8, Simpson, B.J. 1980. The mighty oaks of Texas. Texas Wildflower Newsletter 4(2):5—9. Simpson, B.J. 1980. Silverbells and snowbells. Texas Wildflower hes (3): 11-13. Simpson, B.J. 1980. Adaptation studies. Texas Nurseryman 1 1( Simpson, B.J. 1980. The evergreens of Texas. Texas Wildflower ne es 4(4):5-10, Moon, R.E. and B.J. Simpson. 1980. A vision of trees. Vision 3(6):13—16. Moon, R. ‘ and 7 J. Simpson. 1980. Fall color among Texas Trees. ne Wildflower News- Moon, ‘. ui . B. J. Simpson, 1980. East Texas Trees and Shrubs for Urban Forestry. Texas Forest News 50:1 2-15. Moon, R.E. and B.J. Simpson. 1981. Winter fruits, bark, provide fine sights. Texas Wildflower Newsletter 5(2):5-8. Simpson, B.J. 1981. The drought: combat it with these plants. Texas Wildflower News- etter 5(3):7—-10. Simpson, B.J. 1981. The LO best of Texas. Native Pl. Soc. Texas News 5(4):9—16. Simpson, B.J. 1981. A look at our “lose” Texas Trees and Shrubs. Native Pl, Soc, Texas pws OC) ):3—8 Simpson, = J. 1982 2, Landscaping with Texas woody legumes. Native Pl. Soc. Texas News (2):5—8, 13-14, ee oe 982. Native fruits of Texas offer opportunities. Native Pl. Soc. Texas News 6(3):4-8. Simpson, B.J. 1982. Native fruits of Texas offer new opportunities. Native Pl. Soc. Texas News 6(4):32-7. Simpson, B.J. 1982. Water-logged plants beautify compact areas. Native Pl. Soc. Texas News 7(1):2 Simpson, B.J. 3 The junipers of Texas. Native Pl. Soc. Texas News 7(2):3—7. Simpson, B,J. 1983. After decade of observation, new ornamentals introduced. Native Pl, Soc. Texas News 7(3):9. Simpson, BJ. I 983. Spectacular Texas yuccas. Native Pl. Soc. Texas News 7(4):2-5. Simpson, B.J. 1983. Native release program introduces drought-tolerant plants to Texas andscapes. Amer. Nurseryman 158 (10):90—93, Simpson, B.J. and C. Abbot. 1984. Worthy wildflowers. Native Pl. Soc. Texas News 8(1):2—4. Simpson, B.J. and C. Abborc. 1984. ee species complete list of our worthy wildflowers. Native Pl. Soc. Texas News 8(2):2—4. Simpson, see oe The very best native rene shrubs of Texas. Native Pl. Soc. Texas News 8(1): Simpson, Benny J. re Unique and useful native plants west of the 100" meridian. Nurseryman Simpson, B.J. 1992. Redbuds for spring color. Gardens 6(3):28—29. > 5 3] Simpson, B.J. 1992. Desert willow: a legacy of summer color. Gardens 6(6): 26-28. Simpson, B.J. 1992. Native color for summer. Gardens 6(7):28 Simpson, B.J. 1992. A sunburst of sunflowers. Gardens 6(9): 26-27. Simpson, B.J. 1993. The Rolling Plains in Flower. Native Pl. Soc. Texas News 11(3):1 Simpson, B.J. 1993. Remembering Carroll Abbott. Native Pl. Soc. Texas News 12(4):1 Sipa 17(4): 860. 1997 REVIEWERS FOR VOLUME 17, 1996-1997 The following individuals have kindly supported Sipa through their time and expertise in reviewing manuscripts published in volume 17. Your sup- port is vital and deeply appreciated. There are 402 domestic and 473 foreign subscriptions. Outside of the United States, Brazil has the most subscribers. SibaA, CONTRIBUTIONS TO BoTANY is now distributed in 90 plus countries. Volume 17 is the largest published to date with 873 pages and 84 manu- scripts. Subscriptions for 1998 remain the same for individuals: $25. There has been increased $5 for domestic and foreign institutions; $40 for do- mestic institutions, and $50 for foreign institutions. Page costs charged remain at $35 per printed page. We thank all authors, reviewers, subscribers, and readers for your con- tinued interest and support.—Barney Lipscomb (BRIT), Editor; John W. Thieret (NKU). Associate Editor; Félix Llamas (LEB), Contributing Spanish Edttor. Avila B., Alejandro de Ewan Joseph Miller, ac B. Smith, Gerald L. Baranski, a J. Fox, W.E. Morat, Philipp Stein, Bruce Barker, Fryxell, Paul A. Morden, C lfford W. Stevens, W.D. Barkley, 1 ae M. Gandhi, K.N. Morin, Nancy R. Stone, Benjamin C Barneby, Rupert C. Gereau, Roy Morris, Misael Me ne Strother, John L. Basile, Dominick V Guzman, Ramon Cuevas Naczi, Rober Stuckey, Ronald L. Baskin, Jerry Hall, Gus Nesom, Guy - Swink, Floyd Baxter, John W. Hatch, Stephan L. Nicolson, Dan H. Taylor, Charlotte Benz, Bruce F. Henrickson, Jim Noguchi, Junko Thierec, John W. Bogler, David Hess, William J. Novelo R., Alejandro Todzia, Carol A. Bruederle, Leo P. Hodgson, Wendy oyes, Richard D. Tucker, Gordon C. Bryson, Charles T. Holmgren, Noel H. Padgett, Donald J. Turner, B.L Burger, William Johnson, Dale E. arfitt, Bruce Ullrich Bernc Carter, Richard Jones, Stanley D. Perkins, Kent Urbatsch, Lowell E. Cayouette, Jacques Keller, Harold W. Peterson, Paul M. Vazquez, Antonio Chamberlat iger, wv, Pipoly III, John J Vincent, Michael A. Coile, Nancy C. Kincaid, Dwight T. Pringle, James S. Wall, Dennis P. Conran, John Leeuwenberg, Antony J.M. Pruski, John F. Webster, Rebee D. Cowan, Richard Lersten, Nels R. Reed, Monique, D. Whitkus, Richard Daniel, Thomas F. Les, Donald H. Reeder, John R. Wilkinson, Hazel P. Darwin, Steven P. Lewis, Deborah Q. Ricke eon, Jen M. Wilson, Hu Davidse, Gerrit Llamas, Félix Robinson, Harold Windler, aah R. Davila, Patricia D. Lorence, David H. Rodman, James E. ipff, avis, Aaron P. Loct, Emily Rossman, Amy G. Woodruff, oe dsa Dickinson, Timothy | McPherson, Gordon Rzedowski, J. Wunderlin, Richard P. Diggs, George Magrath, Lawrence K. Sanders, Roger W. Zanoni, Thomas A. Dorr, Laurence J. Martin, David L. Schnepp, John W. Zarucchi, James Douville, Judith A. Martin, Gary J. Shetler, Stanwyn G. Eddie, W.M.M. Mason, Jr., Charles T. Sheviak, C.J. Endress, Mary E. Meerow, Alan W. Smith, Edwin B. Stipa 17(4): 861. 1997 INDEX TO VOLUME 17, 1996—1997 TITLES OF ARTICLES WITH AUTHORS A new combination in Boutelona (Poaceae) by Joseph K. Wiper and Stantey D Jones 109 A new combination in Eragrostis (Poaceae: Eragrostideae) by Pau. M. Prrerson 1O5 A new combination in Thevetia (Apo- cynaceae) by JUSTIN Kirk WILLIAMS 185 A new species of Carex (Cyperaceae) from Uruguay and a new name in the genus by Geratp A. WHeeELER 471 A new species of Ewcrosia a ee from northern Peru by ALAN Mrerow and ABUNDIO SAGASTEGUI Alva 761 A new subspecies of Gentianella heterosepala (Gentianacea) from Mexico by Jost A VILLARREAL Q, 111 A new of Swertia radiata e y JAMES HENRICKSON 83 A new, disjunct variety of Spreelia gentia- notdes (1 variety ; | es (Log ) ft Bibb County, Alabama by ae comes 17 A report of four exotic Cyperms (Cyperaceae) species new to Florida, U.S.A. by Ri- CHARD Carrer, RANDy L. Mears, KATHLEEN CRADDOCK Burks, IARLES T. BRYSON 275 A revision of Erionenron and Dasyochloa (Poaceae: and Eragrostideae) by Jesus VaLbrs-ReyNna and SrepHan L. Hatcu 645 A synopsis of genus Gentlea (Myrsinaceae) and a key to the genera of Myrinaceae in Mesoamerica by JON M. RICKETSON and JOHN J. Popoty HI 697 Annotated checklist of trees and shrubs of bama by STeveN L. YOUNGHANCE and JOHN D. FREEMAN 367 Brachiaria ernciformis and Urochloa brizantha Poaceae: Paniceae) new to Texas by WittiamM E. Fox, III, SrepHan L. Harcu, and RoGer Q. LaNpers, JR. 287 SIDA 17(4): 862. 1997 Calcaratolobelia (Campanulaceae): A new genus of spurred lobelioids from Mexico and oe America by Ros- ert L. WILB Calystegia (C ae in Texas by ANIEL F. Austin, Georce M. Diccs, Jr., and Barney L. Lirscomsp 837 Capraria mexicana (Scrophulartaceae), and endangered addition to the United States E IDEKER 523 Catalpa (Catalpa ea et Bignoniaceae) and bois d’arc (Maclura pomifera, Moraceae) in early Texas records by Dr WENIGER 231 Cerastium pumilum and Stellaria pallida (Caryophyllaceae) new to Texas by RicHarD K. RABELER and ANTON A. REZNICEK 843 Chaenorrhinum minus ae rophulariaceae) new to Texas b RGE M. Diccs, Jr., CONSTANCE ES. ah. and R. JOHN TAYLOR 633 3 s flora by Asa Gray's botanical text- books as Gray and his contemporaries (1836-1887 by Ronatp L, 769 STUCKEY Conductive tissue in Ceratophyllum demersum (Ceratophyllaceae) by Epwarp SCHNEIDER and SHERWIN CaRLQUIST 437 Contributions to the flora and ecology of the northern Longleaf Pine Bele in Rankin County, Mississippi by MICHAEL Wayne Morris 615 Contributions toward a new flora of the Philippines: I. A synopsis of the genus Myrsine (Myrsinaceae) by Joun J. Pipoty THT 115 Conventions for reporting plant chromo- some cae xy JOHN L. SrroTHER ‘SOM 829 C mene nananixonit (Rosaceae Ser. [ntricatae) a new species of hawthorn from Eastern Texas by J.B. Puipps and R.J. O'KENNON 569 and CG INDEX nile Laat orale var, Stewartii a . eypsophi- nee [Re ee . RICHARD SPELLENBERG 282 oe chromosome numbers 1996: _ Miscellaneous U.S.A. and Mexican species, mostly Asteraceae by ZAIMING ZHAO 259 Documented chromosome numbers 1996: synonym of a variz Chromosome numbers in some South African Compositae by JOHN L. STROTHER, LINDA E, WATSON, and Jose L. PANERO 265 Documented chromosome numbers 1996: 4, Chromosome numbers ot Campanu- laceae. IV. Miscellaneous counts by THomas G. Lammers and Nancy HENSOLD 519 Documented chromosome numbers 1996: Carex Sec- |. Chromosome numbers in tion Ovales (Cyperaceae) from eastern North America by P.E. Rorurock and A.A. aici 251 Documented cl numbers 1997.1 Chromosome numbers in Compositae from Morocco and ae by JOHN L. STROTHER and LinpA E. Watson 629 Emanuel D. Rudolph asa a leok reviewer for Chotce by WiuiaM R. Burk 803 Emanuel D. Rudolph’s role for Biological Abstracts by WILLIAM R. Burk 243 a serving society: A case study m the Sierra de sate Biosphere fee by Bruce PF. Jupiry CEvALLos E., nai H Munoz M., and : sco SANTANA M First nea a Wolffiella lingulata (Lemna- ceae) in western Mexico by T.L. MorGan 289 Flora vascular de la Laguna de Babicora, thihuahua, México ou A. EDUARDO Estraba C., RICHAr ToutcHa LEBGUE 309. Floristic petesr of the San José del Cabo , Baja California Sur, México Jose ine LEON DE LA Luz, RAYMUNDO DoMiN- GUEZ CADENA, MiGueL DOMINGUEZ LEON, and José JUAN PEREZ NAvaRRO 599 7, and Sia 17(4): 863. 1997 863 Fuertesimalva, a new genus of neotropical Malvaceae by Paut A. FRYXELL 6S Geographic spatial autocorrelation of mor- phological characters in paaee hakuunensis eral by SOON SUK E Park, and vee GI theca laty ), new to the flora of Iowa by: 1 HOMAS G. ee RS 841 Hybridization in two distantly related Mexi- can black oaks Quercus conzatti and Quercus eduardit (Fagaceae: Quercus: Sec- tion Lobatae) by JEFFREY R. BACON and RICHARD SPELLENBERG 17 Hypochaeris glabra (Asteraceae), a new record for Texas _ Georce M. Diccs, JR., Ropert J. O'KENNON, and BarNey LipscoMB 635 Invasion and spread of Corncya monensts (Brassicaceae) in North America by Ropert F.C. Naczi and JOHN W. Tuierer 43 In Memoriam—Benny J. Simpson 850 n Memoriam—Lynn Lowrey 847 Krapovickasia physaloides (Malvaceae) and Lactuca intybacea (Asteraceae) new to exas and the United States by Larry E. BRowN 291 Las Comunidades vegetales en la Isla Socorro, México by José Luis LEON DE uz, AURORA BrEcEDA, and Rocio er Benet 2 15 Lectot of Clitoria australis Ge sbaceas) by Pau R. Fantz 637 Leaf-print analyses: An ecologically friendly methodology for plant identification by R.J. Ferry Sr., R. FOROUGHBAKHCH, L.A. Hauap, S. Contreras J.V. STAR, M.H. Babu, and H. Gamez 681 Morphological variation and synopsis of the Mubhlenber. ie repens complex (Poaceae) by Ciirrorpd W. Morden and STEPHAN L. Harcu 134 Morphology of Pennisetum orientale (Poaceae: Paniceae) by J. RAMu, S. L. Hatcu, M Hussey, and E. C. BasHaw 163 Naming a southwestern Ma/axzs (Orchi- daceae) by THOMAS K. TODsEN 639 Qu 864 Natural hybridization among three sympa- tric Baptisia (Fabaceae) species in north central Texas by MatrHew A. Kosnik, Grorce M. Jr., Peceoy A. RepsHaw, and Barney L. Lipscombp 479 New county records for Bellardia trixago (Scrophulariaceae) in Texas by L.H. Do AND W.C. Houmes, and J.R. SINGHURST 295 New geogri iphical and morp hological data Sy — Sideroxylon thorner (Sapotaceae) ae C. ANDERSON 343 New species of Ardisia (Myrsinaceae) from Ecuador and Peru by JoHN J. Pipoiy I] 445 New species of Gefssanthus (Myrsinaceae) from the Hylaea/Andean interface of Ecuador and Peru by JOHN J. Pipoiy II] 459 New species of Psychotria Subg. Hetero- psychotria (Rubiaceae: Psychotrieae) from South America by CHARLOTTE M. Taytor 709 New ide of Rhamnaceae from Yunnan, ina ee “AN GUO-SHENG and DENG L ae 67 New species of Credigales on Bignoniaceae from Brazil by Jor P. HENNEN and Heten M.P. Sorao 173 Nomenclatural notes and a synopsis of Mesoamerican Sty/ogyne (Myrsinaceae) JON M. RickrTson and JOHN J. PIpocy HI 591 Nomenclatural notes and a synopsis of the genus Myrsime (Myrsinaceae) in Mesoa- merica by JON M. Rickson and JOHN J. Prpoty HI 576 Nomenclatural notes on Neotropical Clusieae (Clusiaceae) by JOHN J. PIroLy Ill 765 Notes on Hemerocallis middendorffii (Liliaceae) in Korea by SOON SUK KANG, NEONG Gi CHUNG, and Kr Bar Park 83 Notes on Sprgelia (Loganiaceae) by ies :NRICKSON 89 Noteworthy Carex, - om Cyperus, Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) Stipa 17(4): 864. 1997 Stipa 17(4) Arkansas, Louisiana, wena ae North Carolina, Tennessee, and Texas by CHares T Bryson, JOHN R. ae RicH- AAs end STANLEY D. JONES 501 Palynological evidence supporting the iden- from Alabama, Georgia, tity of two taxa of Berberis (Berberi- ArT GARG, ARIQ Husain, and R.R. Rao $75 Passiflora cerasina, a new species of Passiflora daceae) from Tibet subgenus Passiflora (Passifloraceae) from French Guiana by HI aire ANNONAY and CHRISTIAN FEUILLET 551 Pereskia acileata (Cactaceae), in the Lower Rio Grande Valley of IDEKER 527 ~ se Texas by Jor Rare and noteworthy vascular plants from the Fort Campbell Military Reserva- tion, Kentucky and Tennessee by E Wwarb W. Cnestrer, EuGt ones LANDON E., Veer and Davip CAMPBELL 269 haces of Barbieria from Clitorta i rae Phaseoleae: Clitoriinae) by Paut R. Fant: Results of a field survey ne Cyperus grayoides (Cype taceae) in Arkansas by JOHN Hee genus from Mexico and Guatemala by THroport M. BarkLey and JouNn P. JANOVEC 77 Sideroxylon iain a new name for Bumelia anomata (Say by LORAN C. ANDERSON 565 Suggested modifications of the Salisbury stomata index devised from a study of Stanhopea (Orchidaceae) by R.J. Ferry Sr., R. ForouGHBAKHCH, L.A. Hauap H. Gamez, J.V. Star, S. CONTRERAS, d M.H. Babu 691 Saeleaene notes on Rhynchospora crinipes ; and related species in section Fuscae (Cyperaceae) by Ropert Krat 385 Taxonomic notes on the Centrosema pubescens entham complex in Central America (Leguminosae: Phaseoleae: Clitoriinae) by Paut R. Fanrz 321 INDEX Taxonomic status of Hymenocallis choctawen- sis and Hymenocallis puntagordensis (Amaryllidaceae) by GERALD L. SmitH and Mark A. GARLAND 305 Taxonomy of North American species of Zizania (Poaceae) by Epwarp E. TERRELL, PAUL M. PETERSON, ee sL REVEAL, and MELVIN R. Duvatt 533 Th sof Cypripedium ae (Orchidaceae) including a morpholog:- cal analysis of a newly discovered popu- lation in Eastern Virginia by Troy W. Wetpy, Henry T. Mitopozeniec, Lisa W attace, and MartHa A. Case 423 The ees genera of the . DC.) with key and eee taxo- SIDA 17(4): 865. 1997 865 nomic notes by JUSTIN KIRK WILLIAMS LOY The vegetation of Cucphuong National Park, Vietnam by NGuyen NGHIA THIN 719 Two new combinations in Eleocharis (C ypera- ceae) by M. SocorRO GONZALEZ— ELIzONbo and Paut M. Peterson 413 Una nueva especie de Agave ae A — (Agavaceae) de México by 7 A VILLARREAL Q. 191 Wood anatomy of Limnanthaceae and paeolaceae in relation to habic and phylogeny by SHERWIN CaRLQuistT and CHRISTOPHER JOHN DONALD 333 BOTANICAL NAMES New names are in bold face. Aaronsohnia pubescens 628 Acamptopappus sphaerocephalus 260 Agave 191, montana 191 Allamanda 205 Alnus eri ssp. jorullensis 13 Alstont fence ane Anacyclus clavatus 628, radiatus 628 Anthemis arvensis 628 Aphi ia ieur riddellit 260 Apocynaceae Apocynum 205 Arbutus xalapensis 13 Arctotis acaulis 267, undulata 267 Ardisia 445, 698, mexicana 705, premon- 146, zakii 449, Havida 45 bsterii 454 Aspidosperma 206 tana Aster adnatus 621 Astrancthium splendens 260 Athanasia bremeri 266 Baccharis pteronioides || Baileya multiradiata 260 Baptisia 479, australis 496, x bicolor 497 bracteata var. PucO pars 497, x bushi 497, sphaerocarpa 498, x variicolor 498 Barbieria 55, 64, y pinnata 64 Bellardia trixago 295 Berberis replicata var. griffichiaana 576 Brachiaria eruciformis 287 Brazoria arenaria 261 dispar 576, = a 261 Calamintha arkansana 261 Calcaratolobelia 555, 561, macrocentron 562, villaregalis 562, knoblochii 562, mevaughtii 562, cordifolia 562, tenella 563, gibbosa 563, flexuosa var. flexuosa 563, var. 563, goldmanii 563, aurita 563, pringlei 564, margarita 564 Calendula incana subsp. agarbiensis 628 Calystegia 837, macounii 837, Caesalpinea oxycarp intermedia sepium Stipa 17(4): 866. 1997 subsp. limnophila 838, subsp. angulata 838, silvatica subsp. fraterniflora 839 Cameraria 206 Campanula isophylla 521 Campanulaceae 519 Capraria biflora 261 , frucescens 261, mexi- Carex 71, alata 253, albolutescens 3, austrina 502, bicknellii var. opaca 255, cumulata 255 sstucacea 255, fissa var. aristata — gibertit 475, herceri 471, horma- thodes 255, hyalina 255, 503, longi 256, oklahomensis 503, ozarkana 256, reniformis 256, scoparia 504, silicea 256, 256, suberecta 256, tincta 256, tribuloides 257, vexans 257 Cacalpa bignonioides 231 253, cf. brevior fe straminea Celastrus pringlei Centrosema 63, pubescens 321, 330, molle 330 Cerastium pumilum 843 Ceratophyllum demersum 437 Chaenorrhinum minius 631 Chaetopappa bellioides 260, imberbis 260 Chamaemelum mixtum 628, nobile 628, var. discoidea 628 Chrysanthemum coronarium 628, segetum 328 Chrysocoma longifolia 267 Cladanthus arabicus 628 Clermontia kakeana 52] Clitoria 55, australis 635, 636, Subg. Bractearia 63 Clitoriopsis 63 Clusia caudata 7 engleriana 766, colombiana 767, avetia 866, hylaeae sect. ; Pllospenna 7606, polyandra 7 Coincya monensis - oF 2 Comarostaphylis discolor 11 Conoclinium betonicifolium var. betonici- folium 260, var. integrifolium 260 Coreopsis wrightit 260 INDEX Corydalis micrantha 261 Cotula sehen Sar 266 Crataegus nananixonii 569, pubescens 11 Cc ees 698 Cyanea hirtella 521 Cybianthus 698 Cyperus a s 505, alopecuroides 275, diffor 505, distans 276, eragrostis 507, oe 283, lancastriensis 508, lanceolatus 508 oxylepis 510, pilosus 510, prolifer 277, S11, 278, surinamensis 5 Cyphomeris gypsophiloides var. stewartii 282 ochraceus pumilus sphacelatus —_ — Cypripedium kentuckiense 423 Dasyochloa 645, 663, pulchella 664 Dyssodia pentachaeta var. pentachaeta 260, tenuiloba var. tenuiloba 260 Echites 20 Eleocharis albibracteata var. boliviana 414, macrostachya 512, quinqueflora var. bernardina 414 Eragrostis 105, pectinacea var. tracyi 106, hirsuta subsp. hirsuta 109, subsp. a 109 Erigeron tenuis 260 Erioneuron 645, 656, avenaceum 660, var. kurtzianum 661, var. pygmaeum 662, , veracruzensis 260 var. longiglume 662, nealleyi 663, pilosum 657, var. mendocinum 658, var. longearistatum 65 Eucrosia 761, calendulina 761 Fernaldia 207 Forsteronia 208 Fragaria mexicana 12 Pactesimalva 69, 70, chilensis 72, corni- ulat 5 ata 72, echinata 72, insularis 73, jacens 73, ss fies pie calyx 73, limensis 74, pen , penta- cocca 74, pen aie ve peruviana 74, sanambrosiana 75, sere 75 Gaillardia aestivalis var. aestivalis 260 odon 260, pulchella var. Simei Geissanthus 698, vanderwerffii 460 spectabilis 463, challuayacus 466 Sipa 17(4): 867. 1997 867 Gentianella heterosepala subsp. durangensis 111 Gentlea 697, 698, 699, austinsmithii 704, crenulata 702, lancifolia be latisepala 706, seen 705, penduliflora 700, standleyi 7 vatterit 702, venosissima , stevensii bey taca- nensis 703, 03 Gilia ludens 261 Glossopappus macrotus 628 Grindelia obvatifolia 260, tenella 260 , rigidula 261 Haplophyton 20¢ Havetiopsis martit 767 Hemerocallis hakuunensis 667, midden- dorfhi 833 Heterolepis aliena 267 Heterotheca latifolia 841, mucronata 260 Heterotoma 560, lobelioides var. lobelioides 561, var. glabra 5 Hippobroma longiflora 5 21 Hymenocallis erie caigen 305, 307, puntagordensis 305, Hymenoxys linearifolia 2 Hypericum galioides 622, setosum 622 Hypochaeris achyrophorus 628, glabra 633 Hyptis alata 623 Hlicitum dori danaiin G22 Tris cristata 622 Krapovickasia physaloides 291 Kyllinga brefifolioides 512 Lactuca intybacea 291 Laubertia 208 Leysera tenella 267 Limnanthaceae 333 Limnanthes douglasi 333 Lobelia appendiculata 261, erinus 521 Ludwigia linearis 623 Lupinus caballoanus 261, platamodes 261 texensis 26 Machaeranthera pinnatifida 260 Maclura pomifera 231 Malacothrix fendleri 260 Malaxis Sanne rgii 637, porphyrea 637 Mandevilla 208 Melanthium virginicum 623 Mesechites 208 Mikania cordifolia 621 868 Muhlenbergia fastigiata 358, plumbea 360, 360, utilis 363, villiflora var. villiflora 364, repens 349, richardsonis 362, var. villosa 364 Myrsine 115, 119, 579, 126, aralioides 152, arvensis 144, S80, apoensis 698, 125, calcarata amorosoana 586, coriacea 583, subsp. coriacea 583, subsp. nigrescens 584, cruciata 130, cubana 587, densiflora 132, dependens 582, fastigiata 149, glandulosa 154, juergensentt 586, medeciloae 122, mindanaensis 140, multibracteata 48, oblongibacca 129, pellucido- 585, pen 4 peregrina 136, philippinensis 14 peltata 272 punctata nibukana 14 Nymphoides Odontadenia 209 — grandiflorum 266, suffrucico- 1 266 Or cnn maritimus 628 sel carpum cubense Pallensis spinosa 625 Panax li 618 Parathesis 698 Passiflora cerasina 551 Pennisetum orientale 163 > 5 Pentalinon 2 Pereskia aculeata 527 Periandra 63 Persea palustris 62 Phacelia integrifolia 261 Phragmites australis 273 Platanthera cristata 624 Pluchea foetida 621 Plumeria 209 Polygala boykinit 624 Porophyllum gracile 260, scoparium 260 Porotenus biporus 175, bibasiporulus 177 Prestonia 209 Prospodium amapaensis 180, laevigatum 182 Psilostrophe cooperi 260 Psoralea oo 213 Psycho — 1 709, cauligera 709, conepho- ere 715, eae 716, costanensis 716, cuatrecasasii 716, fusiformis Sipa 17(4): 868. 1997 Sipa 17(4) i 712, Subg. oe ene hypochlorina 713, 716, 2; ervisei 716, paeonia 716, en, 16, transiens 716 Quercus conzatti 17, eduardii 17, rugosa 12, obtusata 12, scycophylla 13 Rapidophyllum hystrix 618 Rauvolfia 210 Rhabdadenia 210 Rhamnus daliensis 680 397, curtissil Fuscae 387, , pleranctha 389 Rhynchospora crinipes 385, 405, filifolia 407, fuscae 388, harperi 394 sect. Richardia brasiliensis 272 TC Robinsonecio 77, ie gerberifolius 7 Oo hyresthes Rubus faa 6 , pringlei Rumfordia exauriculata 260 Sabatia campanulata 622 Sageretia gongshanensis 677 Schisandra glabra 624 Scutellaria ovata var. mexicana 261 Sideroxylon alachuense 565, tenax 567 thornet 343 Silphium gracile 260, simpsonii var. wrightil 260 Smilax moranensis | 2 Solidago altissima 260, gigantea 260, velucina 260 Spigelia 89, eatin 417, var. ala- bamensis 418, loganioides 93, hedyo- tidea 99 Stanhopea 681, 691] Scellaria pallida 843 Stemmadenia 210 Stevia lucida var. oaxacana 260 Scewartia malacodendron 624 Streptanthus hyacinthoides 261 Stylogyne 591, 592, 698, darienensis 593, elamerulisor 594, hayesit 593, turbacensis subsp. turbacensis 594 subsp. laevis 595 coals QS Styrax ramirezii Swertia radiata var. maderensis 83 Symplocos citrea 10 Synardisia 698 Tabernaemontana 210 INDEX Telosiphonia 210 Tephrosia lindheimeri 261 Ternstroemia lineata 10 Thelesperma megapotamicum 26] Thenardia 211 Thevetia 185, 211, peruviana 187, pinifolia 8 / Tintinnabularia 211 Trachelium coeruleum 521 Trachelospermum 212 Tradescantia hirsutiflora 261, humilis 261 occidentalis 261, reverchonii 261, subacaulis 261 Tridax balbisioides 261 Triphora trianthophora 624 Tropaeolaceae 333 Siva 17(4): 869. 1997 869 Tropaeolum majus 333 Uredo amapaensis 179 Urocarpidium 71 Urochloa brizantha 287 Ursinia abrotanifolia 267, nudicaulis 267 pinnata 267, punctata 267 Vallesia 21. Verbesina mexicana 261, walteri 622, zaragosana 261 Wolffiella lingulata 289 Xylothamia pseudobaccharis 261, riskindit 26] Zea diploperennis 3 Zizania 533, 534, aquatica 535, var. brevis 538, paluscris 539, var. interior 542, texana 543 AUTHOR INDEX Anderson, Loran C, 65 Hensold, Nancy 519 Annonay, Hilaire a Holmes, W.C. 295 Austin, Daniel F. — Husain, Tariq 575 Bacon, Jeffrey R. Hussey, M.A. 163 Badu, M.H. 681, im Ideker, Joe 523, 527 Barkley, M. 77 Janovec, John P. 77 Bashaw, E.C Jones, Stanley D. 109, 50 Benet, ee . oria 215 Kang, Soon Suk 667, 833 Benz, Bruce F. | Kosnik, Matthew A. 479 Breceda, Aurora 215 Kral, Robert 385 Brown, Larry E. 289 Lammers, Thomas G. 519, 841 Bryson, Charles a etm S01 Landers, Roger Q. 287 Burk, William R. 803 Lebgue, Toutcha 807 Burks, Kathleen a ri ee 275 Leon de la Luz, José Luis 215, 599 Campbell, David 269 Deng, L.L. 677 Carlquist, Sherwin 333, 437 Lipscomb, Barney 633, 479, 837 Carter, ae er ee SO] Logan, John M. 283 Case, Marcha 23 MacDonald, John R. 501 See E. eee McKinney, Landon E. 269 Chester, E hee W. 269 Mears, Randy L. 27 Chung, Myong Gi 667, 833 Meerow, Alan W. 761 Contreras, S. 681, 69] Mlodozeniec, Henry T. 423 Diggs, Jr., George M. 479, 631, 633, 837 Morden, Clifford W. 349 Do, L.H. 295 Morris, Michael Wayne 615 Dominguez Cadena, Raymundo 599 Munoz M., Elizabeth 1 aa Leon, Miguel 599 Mischalel- Derek A. 289 Donald, Christopher John 333 Naczi, Robert EC. 43 Duvall —— 333 Nesom, Guy L. 829 Estrada Tas Eduardo 809 O’Kennon, R.J. 569, 633 Fan, G.S. V7 Panero, nJoee 1G . oo. Paul R. 55, 321, 635 Park, Ki Bae 667 Ferry, Sr., R.J. 681, 691 Pérez Navarro, seer S99 Feuillec, Christian 55 | Peterson, ue A 105, 413,533 Foroughbakhch, R. 681, 691 Phipps, J.B. Fox IIT, William E, 287 Pipoly = hn} 445,459,579, 591, Freeman, John D. 367 697 Fryxell, Paul A. 69 eee ed K. 843 Gamez, H. 681, 691 Ramu, J. 163 Garg, Arti 575 Rao, R.R. 575 Garland, Mark A. Redshaw, Peggy A. 479 Gonzalez-E ane 5 Socorro 413 Reveal, ae L.533 Gould, Katherine 417 Reznicek, A.A. 251, 843 Hatch, Stephan L. 163, 287, 349, 645 Ricketson, Jon - 579,591, 697 Hauad, L.A. 681, 691 Rothrock, PE Hennen, Joe F173 Sagastegui A. iis 76I Henrickson, James 89 Santana M., Francisco 1 SIDA 17(4): 870. 1997 — INDEX Schneider, Edward L. 437 Singhurst, J.R. 295 Smith, Gerald L. 305 Sorao, Helen M.P. 173 Spellenberg, Richard 17, 282, 532, 8 Star, J.V. 681, 691 Strother, John L. 265, 627, 829 Stuckey, Ronald L. 769 Taylor, Charlotte M. 709 Taylor, Constance E.S. 631 Taylor, R. John 631 Terrell, Edward E. 933 3 ea ee n, N.N. 7 19 Sipa 17(4): 871. 1997 ( 7 Todsen, Thomas K. 637 Valdés-Reyna, Jestis 645 Villarreal Q., José A. 111, 191 Wallace, Lisa E. 42: Watson, Linda E. 265, 627 Weldy, Troy W. . 23 Weniger, Del 2 Wheeler, eu n Wilbur, Robert L Williams, Justin on 185, 197 Wipff, Joseph K. 109 Wofford, B. Eugene 269 Younghance, Steven L. 367 Zhao, Zaiming 259 jon 872 Sipa 17(A4) SIDA GUIDELINES FOR CONTRIBUTORS SIDA, CONTRIBUTIONS TO BoTAaNy is an international journal of systematic botany con- taining primary research papers sensu lato, including anatomy, biogeography, chemotax- onomy, Cladistics, ecology, floristics, genetics & evolution, numerical taxonomy, paleo- botany, and palynology. Sipa is open to all auchors anywhere; coverage is not restricted to any peop i l area. All manuscripts submitted to Sipa are considered by at least two reviewers. Manu- ae may be in English or Spanish. Page costs are $35 per page but may be wavered or reduced under certain circumstances. Please contact the editor and make arrangements before publication. Manuscripts are not rejected due co lack of financial support. Submit manuscripts and editorial questions to Barney Lipscomb, Botanical Research Institute of Texas, SO9 Pecan Street, Fore Worth, TEXAS 76102-4060, U.S.A. Our electronic mail address ts sida@brit.org. Access our Sipa home page at the URL Aitp://www. brit.org/sida/ Manuscripts must be double spaced throughout and three copies sent to the editor. Upon acceptance for publication please return two copies along with a copy in your word- processing program format (with format commands) on a 3!/2” data disk. You may also send your electronic copy attached to an e-mail. Consult the latest issue of SpA for format of articles and notes regarding title, author, and address. Ranges of numbers are separated by double hyphens (6--8). References, footnotes, and legends should be typed on separate pages and inserted at end of manuscript. Abstract Every paper should include both an English and Spanish, or another major language abstract Specimen Citations Use the collector's last name and number or (s.n.) and underline. Abbreviate the months of the year by the first three letcers of the month without a period. Example: Texas. Jeff Davis Co.: | mi W of Fort Davis, 3 Jan 1972, Smith 118 (BRIT). Author Citations Use barium, Royal Botanic Gardens, Kew (1992). Use et or the e ampersand (&) between two authors; for more than two authors, restrict to the first one followed by et al. Do not the author abbreviations as given in the Author Abbreviations compiled at the Her- italicize or underline in the manuscript the terms ec, et al., ex, or in. Latin Descriptions or Diagnoses Either a Latin description or diagnosis must be included for each new taxon. This Latin is not to be underlined in the manuscript except as noted below. The ICBN (Rec. 32B.1) recommends that the main points in which a new taxon differs from its allies should be included. These main points would make up a diagnosis. Ifa full Latin description is used, those main diagnostic points must be underlined. If a Latin diagnosis is used, including just those differentiating points, do not underline. After the Latin description or diagnosis an English version is recommended, following the same underlining requirements as noted above. Sipa 17(4): 872. 1997 GUIDELINES 873 Figures A scale should be in or on the illustration or photograph; reduction will automatically reduce the scale and subject proportionately. In half-tones, sharp glossy photographs with uction. Figures are to be numbered in a single jos good contrast are necessary for good reproc sequence in the manuscript. Please mount illustrative material (half-tones, line drawings, etc.) on flexible card stock. Illustrative material may be scanned with a high speed drum scanner. You may also submit graphic materials in electronic format. Please consult the editor for proper graphics format (tif, jpeg, gif., bmp, etc.). Color figures are best if repro- duced from a color transparency. References This includes all of the literature cited in the text and may include other article cita- tions the author may deem desirable. Normal text references should be cited as follows: ‘Smith (1976) stated...’ or ‘the latest revision (Smith 1980)’ when reference is used as authority for a statement. When there are ae or more authors use only the name of the first author followed by et al.: ‘Smith et al. (1933) stated’. References at the end of the article are arranged alphabetically and Eee making use of a,b, etc. ifan author had more than one publication in a given year. Author's names are typed in lower case except for initial capitals. Only the first letter of the initial word, proper nouns, and proper adjectives of titles should be capitalized. When the author of the relevant part of a book 1s not an author of the book, the sequence, punctuation, and spacing for references to that part are as follows. SHOES) of the part. Year. Title of the part. In: Author(s) or Editor(s). Title of the book. Place of publication: Publisher. Pages of the part. For journal abbrevia- tions, use Botanico-Periodicum-Huntianum. jon Abbreviations When ay — abbreviations are used the aca is omitted. Distance: mm, dm, cm, m, km mi; directions (in caps): S, N, E, W, m : first 3 letters only, Jan, Feb, etc. es Paiie: Co.: 2 km W of Dot, 5 cin 197 i. cae = (SMU). Notes When references are few in number, cite them in the text and omit reference section. Text citations of literature are in parentheses and should state the journal abbreviation, volume, pages, and year. The citation of a book includes the last name of the author, if not mentioned previously, the full unabbreviated title of the book, and the year published in parentheses. Documented Plant Chromosome Numbers Refer to Stipa 17:627—629. 1995. Sipa 17(4): 873. 1997 Commentaries on Asa Gray's botanical textbooks by Gray and his contemporaries (1830-1887 ) Ronald L. Stuckey 709 Emanuel D. Rudolph as a book reviewer for Choice William Rk. Burk 803 Flora vascular de la Laguna de Babicora, Chihuahua, México A. Eduardo Estrada C., Richard Spellenberg, and Toutcha Lebgue SOY Conventions for reporting plant chromosome numbers Jobn L. Strother and Guy L. Nesom 82° Notes Notes on Hemerocallis middendorffii (Liliaceae) in Korea 833 Calystegia (Convolvulaceae) in Texas 337 Heterotheca latifolia (Asteraceae), new to the flora of lowa Cerastium pumilum and Stellaria pallida (Caryophyllaceae) new to Texas 843 In Memoriam—Lynn Lowrey 847 In Memoriam—Benny J. Simpson 85‘ Reviewers for volume 17, 1996-1997 86 ) Index to volume 17, 1996-1997 8¢ ~ SIDA guidelines for contributors 87 Book notices and reviews 676, 696, 708, 760 Vv | ISSN 00360-1488