Molecular Phylogeny of Stryphnodendron (Mimosoideae, Leguminosae) and Generic Delimitations in the Piptadenia Group Author(S): Marcelo F
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Molecular Phylogeny of Stryphnodendron (Mimosoideae, Leguminosae) and Generic Delimitations in the Piptadenia Group Author(s): Marcelo F. Simon, José Floriano B. Pastore, Adriana F. Souza, Leonardo M. Borges, Viviane R. Scalon, Pétala G. Ribeiro, Juliana Santos-Silva, Vinícius C. Souza and Luciano P. Queiroz, Source: International Journal of Plant Sciences, (-Not available-), p. 000 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/684077 . Accessed: 24/11/2015 08:42 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. http://www.jstor.org This content downloaded from 23.235.32.0 on Tue, 24 Nov 2015 08:42:38 AM All use subject to JSTOR Terms and Conditions Int. J. Plant Sci. 177(1):000–000. 2016. SYMPOSIUM PAPER q 2015 by The University of Chicago. All rights reserved. 1058-5893/2016/17701-00XX$15.00 DOI: 10.1086/684077 MOLECULAR PHYLOGENY OF STRYPHNODENDRON (MIMOSOIDEAE, LEGUMINOSAE) AND GENERIC DELIMITATIONS IN THE PIPTADENIA GROUP Marcelo F. Simon,1,* José Floriano B. Pastore,*,† Adriana F. Souza,* Leonardo M. Borges,‡ Viviane R. Scalon,§ Pétala G. Ribeiro,∥ Juliana Santos-Silva,# Vinícius C. Souza,** and Luciano P. Queiroz∥ *Empresa Brasileira de Pesquisa Agopecuária (Embrapa) Recursos Genéticos e Biotecnologia, Parque Estação Biológica, Caixa Postal 02372, Brasília, Distrito Federal, 70770-917, Brazil; †Universidade Federal de Santa Catarina, Campus Universitário de Curitibanos, Caixa Postal 101, rodovia Ulysses Gaboardi km 3, Curitibanos, Santa Catarina, 89520-000, Brazil; ‡Departamento de Botânica, InstitutodeBiociências,Universidadede São Paulo, Rua do Matão 277, Cidade Universitária, São Paulo, São Paulo, 05508-090, Brazil; and William and Lynda Steer Herbarium, New York Botanical Garden, 2900 Southern Boulevard, Bronx, New York 10458-5126, USA; §Departamento de Biodiversidade, Evolução e Meio Ambiente, Instituto de Ciências Exatas e Biológicas, Universidade Federal de Ouro Preto, Ouro Preto, Minas Gerais, 35400-000, Brazil; ∥Universidade Estadual de Feira de Santana, km 03, BR-116, Campus, Feira de Santana, Bahia, 44031-460, Brazil; #Departamento de Ciências Humanas–DCH/Campus VI, Universidade do Estado da Bahia, Avenida Contorno s.n., Bairro São José, Caetité, Bahia, 46400-000, Brazil; and **Escola Superior de Agricultura “Luiz de Queiroz,” Universidade de São Paulo, Avenida Pádua Dias 11, Piracicaba, São Paulo, 13418-900, Brazil Editors: Gwilym P. Lewis and Patrick S. Herendeen Premise of research. Despite recent progress on the systematics of the Leguminosae, the relationships be- tween some genera, as well as their circumscriptions, remain uncertain. Here we investigate the relationships between Stryphnodendron and closely related genera in the Piptadenia group, a clade of problematic taxon- omy for which a robust phylogenetic hypothesis is still lacking. Methodology. Taxonomic sampling included 23 species of Stryphnodendron (77% of the genus) and rep- resentatives of genera of the Piptadenia group. DNA (matK/trnK, trnD-trnT, trnL-trnF, and ITS) sequences con- stituting a data set with 6798 bp and 94 terminals were used in parsimony and Bayesian analyses. A set of 17 mor- phological characters were optimized on the estimated phylogeny to evaluate clade synapomorphies and character evolution. Pivotal results. Our increased molecular and taxonomic sampling improved the resolution of a previously largely unresolved clade that included Microlobius, Parapiptadenia, Pityrocarpa, Pseudopiptadenia, and Stryph- nodendron. Major lineages within this clade are an early-diverging clade containing the sister genera Parapip- tadenia and Pseudopiptadenia; the genus Pityrocarpa, with Pseudopiptadenia brenanii nested within it; and a Stryphnodendron clade including the monospecific genus Microlobius. Samples of Stryphnodendron grouped into three strongly supported clades: a group of seven species bearing large leaflets and relatively few pairs of pinnae, sister to Microlobius foetidus; a clade containing samples of Stryphnodendron duckeanum; and a large and poorly resolved clade containing the remaining species of Stryphnodendron, including species with small leaflets and higher number of pinnae and highly specialized dwarf subshrubs adapted to savanna fire regimes. Lack of resolu- tion within this clade suggests rapid and recent diversification in both savannas and rain forests in the Neotropics. Conclusions. Our findings highlight the importance of densely sampled phylogenies to rigorously test the monophyly of genera. Key morphological characters are proposed to assist generic delimitations in the Piptadenia group. New taxonomic rearrangements are needed in order to accommodate the results presented here. Keywords: Amazon, Cerrado, diagnosability, diversification, monophyly, Neotropics. Online enhancement: appendix figures. Introduction 2012; LPWG 2013), the monophyly of many genera and rela- tionships between a number of clades remain uncertain. Among Despite recent progress on the molecular systematics of the the mimosoid legumes, one group where generic relationships Leguminosae (Lavin et al. 2005; Lewis et al. 2005; Cardoso et al. are problematic is the Piptadenia group, an informal assemblage that includes the following genera: Anadenanthera Speg., 1 Author for correspondence; e-mail: [email protected]. Microlobius C. Presl, Mimosa L., Parapiptadenia Brenan, Pip- Manuscript received May 2015; revised manuscript received August 2015; tadenia Benth.,PseudopiptadeniaRauschert, and Stryphnoden- electronically published November 20, 2015. dron Mart. The group was defined by Lewis and Elias (1981) 000 This content downloaded from 23.235.32.0 on Tue, 24 Nov 2015 08:42:38 AM All use subject to JSTOR Terms and Conditions 000 INTERNATIONAL JOURNAL OF PLANT SCIENCES on the basis of the shared presence of compound pollen (polyads) Ecological preference and habit in the genus range from rain and a small porate stigma at the end of a narrowing style. Those forest canopy trees (e.g., S. excelsum, S. moricolor, and S. pani- features are not exclusive to the Piptadenia group, but other culatum) to smaller trees that colonize disturbed areas (e.g., Mimosoideae taxa where they occur have very dissimilar flower S. foreroi and S. pulcherrimum), savanna trees (e.g., S. adstrin- morphology (Lewis and Elias 1981). gens and S. coriaceum), and dwarf geoxylic subshrubs (e.g., The taxonomy within the Piptadenia group has been rather S. gracile, S. heringeri, and S. pumilum) that grow in fire-prone unstable because of a series of nomenclatural changes at the savannas of the Cerrado in central Brazil (Rizzini and Heringer generic level, most notably within the genus Piptadenia, which 1987; fig. 1). Medicinal qualities of the bark of S. adstringens has been subject to a number of generic realignments based on have been described in the literature (Audi et al. 1999; Her- morphological studies (Brenan 1955, 1963, 1986; Lima and nandes et al. 2010), and the high concentration of tannins Lima 1984; Lewis 1991a, 1991b; Lewis and Lima 1991), lead- provides the basis of anti-inflammatory, analgesic, and healing ing to the recognition of segregate genera such as Parapipta- properties. An ointment based on the bark extract of S. ad- denia and Pseudopiptadenia. More recently, a molecular-based stringens, a species that has been extensively used in the tradi- phylogenetic study focused on Piptadenia found the genus to tional medicine in Brazil, has been developed for treatment of be polyphyletic, resulting in a reclassification of the genus that skin wounds (Minatel et al. 2010) and is now being commercial- rendered a monophyletic Piptadenia sensu stricto with 23 spe- ized by a Brazilian pharmaceutical company. cies, the resurrection of Pityrocarpa (Benth.) Britton & Rose, The most comprehensive phylogenetic study that focused on currently with three species, and the recognition of Piptadenia the Piptadenia group (Jobson and Luckow 2007) presented a viridiflora as an isolated lineage for which a new genus name is substantial contribution to the understanding of relationships still pending (Jobson and Luckow 2007). Such rearrangements within the group. However, there are still open questions re- are supported by a combination of morphological character garding generic delimitations, mainly because of a lack of reso- states, namely, the occurrence of stipular spines, inflorescence lution in the phylogeny and limited taxon sampling. The aim organization, and flower morphology (Jobson and Luckow of this study was to produce a phylogenetic hypothesis for the 2007). Further insights into the relationships and circumscrip- Piptadenia group based on plastid and nuclear loci, as well as to tion of genera within the Piptadenia group were provided by test the monophyly of genera previously undersampled in phy- molecular phylogenetic studies based on plastid loci (Luckow logenetic studies, with emphasis on the genus Stryphnodendron. et al. 2003; Jobson