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Phylogenetic Systematics of Strophostyles (Fabaceae): a North American Temperate Genus Within a Neotropical Diversification Author(S): Erin T

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Phylogenetic Systematics of Strophostyles (Fabaceae): a North American Temperate Genus Within a Neotropical Diversification Author(S): Erin T Phylogenetic Systematics of Strophostyles (Fabaceae): A North American Temperate Genus Within a Neotropical Diversification Author(s): Erin T. Riley-Hulting, Alfonso Delgado-Salinas, Matt Lavin Source: Systematic Botany, 29(3):627-653. Published By: The American Society of Plant Taxonomists DOI: http://dx.doi.org/10.1600/0363644041744464 URL: http://www.bioone.org/doi/full/10.1600/0363644041744464 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Systematic Botany (2004), 29(3): pp. 627±653 q Copyright 2004 by the American Society of Plant Taxonomists Phylogenetic Systematics of Strophostyles (Fabaceae): A North American Temperate Genus Within a Neotropical Diversi®cation ERIN T. R ILEY-HULTING,1 ALFONSO DELGADO-SALINAS,2 and MATT LAVIN1 1Department of Plant Sciences and Plant Pathology, Montana State University, Bozeman, Montana 59717; 2Departamento de BotaÂnica, Instituto de BiologõÂa, Universidad Nacional AutoÂnoma de MeÂxico, Apartado Postal 70-233, 04510, MeÂxico, D. F., MEXICO Communicating Editor: Aaron Liston ABSTRACT. A combined parsimony analysis of cpDNA trnK, nrDNA ITS/5.8S, and morphology reveals that the genus Strophostyles is monophyletic. In contrast to the conventional view of the geographic relationships of eastern North America, Strophostyles is most closely related to neotropical genera. Its sister is the South American genus Dolichopsis, which is endemic to the Chaco, a region characterized by having an annual frost interval. Strophostyles is apomorphically diagnosed by diver- gent stipules, persistent secondary ¯oral bracts, calyces with four acute to sometimes attenuate lobes, and seed testa often with a cellular coat. The relationship with Dolichopsis is supported in part by a shared keel petal morphology involving a gibbous ventral margin proximal to the rostrum. Phylogenetic analysis of ITS/5.8S sequences and morphometric analysis of quantitative traits suggest that the three traditionally recognized species of Strophostyles can be recognized under the phylogenetic species concept. Strophostyles umbellata is the most genetically variable at the ITS locus and geographically centered in southern Appalachia. Strophostyles helvola shows the least amount of intraspeci®c genetic variation at this locus, suggesting a recent and rapid range expansion throughout eastern North America. Nucleotide sequence variation is inter- mediate in Strophostyles leiosperma, a species distributed primarily in central North America. The genus Strophostyles Elliott is classi®ed within is similar to the ¯oral morphology of Dolichopsis, Ory- the tribe Phaseoleae of the legume subfamily Papilion- xis, Oxyrhynchus, Ramirezella, and some Vigna subge- oideae. The liana habit of Strophostyles is common to nus Sigmoidotropis. All other New World Phaseolinae papilionoid genera especially of this tribe. The asym- genera have a keel beak that is abruptly hooked or metric ¯oral morphology of Strophostyles, whereby the coiled at least one-half turn to the right side of the rostrate keel petals curve to the right side of the ¯ower, ¯ower. Although MareÂchal et al. (1978) and Pelotto and is characteristic of many genera in the tribe Phaseoleae del Pero MartõÂnez (1998) have used overall similarity subtribe Phaseolinae, a group of trifoliolate-leaved li- in morphology or secondary chemistry, respectively, to anas comprising such well-known genera as Phaseolus suggest a close relationship of Strophostyles with Doli- L. and Vigna Savi. Indeed, the most recent higher level chopsis, such analyses have not been comprehensive in taxonomic treatment of Phaseoleae (Lackey 1981), in taxon sampling, or have involved only limited data. addition to phylogenetic analyses of nuclear ribosomal The suggestion that the northern temperate Strophos- 5.8S and ¯anking internal transcribed spacers (the ITS tyles is sister to Dolichopsis, which is endemic to the region; Delgado-Salinas et al. 1999) and the chloroplast Chaco, is in need of independent veri®cation with trnK locus (Delgado-Salinas et al. unpublished data), DNA sequence data. suggest that the closest relatives of the temperate Stro- At the species level, ¯oristic treatments dealing with phostyles are the primarily neotropical Dolichopsis Hass- Strophostyles (e.g., Radford 1968; Correll and Johnston ler, Macroptilium (Bentham) Urban, Mysanthus G.P. 1970; Isely 1998) have recognized at least three species, Lewis & A. Delgado, Phaseolus, Oryxis A. Delgado & although species delimitation remains uncertain. Mis- G.P. Lewis, Oxyrhynchus Brandegee, Ramirezella Rose, identi®cation is common especially in the southeastern and Vigna subgenus Sigmoidotropis (Piper) Verdcourt. USA where the distributions of the traditionally rec- Although these genera, collectively the New World ognized species broadly overlap. This dif®culty arises Phaseolinae (Lackey 1983), form a strongly supported because either the key morphologies are inadequate for clade (Wojciechowski et al. in press; Delgado-Salinas diagnosing species identity, or species delimitations et al. unpublished data), the putative monophyly and have been incorrectly drawn, or extensive introgressive intergeneric relationships of most of them, including hybridization is occurring. Strophostyles, has never been comprehensively ad- This study was designed to determine the closest dressed. relatives of Strophostyles using nucleotide sequences The New World Phaseolinae generally show a high from the cpDNA trnK and nrDNA ITS/5.8S regions, degree of elaboration of petal morphologies, particu- as well as morphological data. Such data were targeted larly in deviations from bilateral symmetry. In Stro- because they have been shown to be highly informa- phostyles, the standard, wings, and keel petals are bi- tive in legumes from the level of closely related genera laterally symmetrical except for the distal end of the down to populations within an individual species (e.g., keel, which curves to the right side of the ¯ower. This Delgado-Salinas et al. 1999; Hu et al. 2002; Lavin et al. 627 628 SYSTEMATIC BOTANY [Volume 29 TABLE 1. Morphological characters scored for discrete states that are phylogenetically informative at the species and genus level. See Appendix A for character descriptions. An ``L'' designates a multistate taxon. Phylogenetically informative characters 1234567891011121314151617 Ramirezella spp. 000L0000000000000 Oxyrhynchus spp. 000L0000000000100 Macroptilium spp. 00000111011L0000L Mysanthus uleanus 01010111011100001 Oryxis monticola 01010000001000101 Dolichopsis paraguariensis 11000000100110111 Dolichopsis ligulata 11000000100110111 Strophostyles umbellata 00101000100001101 Strophostyles helvola 00101000100001101 Strophostyles leiosperma 00101000100001101 2003; Schrire et al. 2003). The lack of a comprehensive TCGAAGG-39;and39 trnK intron reverse, TK2R: 59-CCCGGAAC- TAGTCGGATGG-39. DNA sequencing was performed on an au- taxonomic treatment of the genus at the species level tomated sequencer at Northwoods DNA (Becida, Minnesota). Phy- and below warranted a reevaluation of the constituent logenetic data are available from http://gemini.oscs.montana. taxa within Strophostyles. The goals of this study thus edu/;mlavin/data/stroph.htm and TreeBase study accession include a taxonomic monograph of the genus Strophos- number S1015. Missing entries accounted for 1.5% of the ITS se- quence data set, and 8.3% of the combined data set. This last ®gure tyles, which addresses the identity and relationships of is high because trnK sequences were not obtained for Oryxis (if species within the genus, as well as the relationships this genus is omitted, missing entries account for 2.9% of the com- of the genus to putative neotropical relatives. bined data matrix). Phylogenetic Analysis. Sequences were aligned manually with Se-Al (Rambaut 1996). Maximum parsimony analyses were per- MATERIALS AND METHODS formed with PAUP* (Swofford 2001). The combined data set was analyzed with the branch and bound search option, whereas the Taxon Sampling. Each of the three species of Strophostyles was ITS data set required a heuristic search that included 100 random sampled as exhaustively as possible for sequences from the ITS addition replicates and tree-bisection-reconnection branch swap- region, as well as for morphological variation. Field and herbarium ping. In all analyses, the maximum number of trees was set at specimens of Strophostyles and related genera were sampled for 10,000, which is suf®cient to capture all topological variation (cf. morphological characters that included the many vegetative, ¯oral, Sanderson and Doyle 1993). Bootstrap resampling for clade sup- and fruiting traits that
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