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Phylogenetic Analysis of Microlicieae (Melastomataceae), with Emphasis

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Phylogenetic Analysis of Microlicieae (Melastomataceae), with Emphasis applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Botanical Journal of the Linnean Society, 2021, 197, 35–60. With 6 figures. Phylogenetic analysis of Microlicieae (Melastomataceae), with emphasis on the re-circumscription of the large genus Microlicia Keywords=Keywords=Keywords_First=Keywords 1, 2 3 HeadA=HeadB=HeadA=HeadB/HeadA ANA FLÁVIA ALVES VERSIANE *, ROSANA ROMERO , MARCELO REGINATO , Downloaded from https://academic.oup.com/botlinnean/article/197/1/35/6188938 by guest on 01 October 2021 2 4 HeadB=HeadC=HeadB=HeadC/HeadB CASSIANO A. DORNELES WELKER , FABIÁN A. MICHELANGELI and RENATO GOLDENBERG5 HeadC=HeadD=HeadC=HeadD/HeadC Extract3=HeadA=Extract1=HeadA 1Programa de Pós-Graduação em Biologia Vegetal, Departamento de Biologia Vegetal, Universidade REV_HeadA=REV_HeadB=REV_HeadA=REV_HeadB/HeadA Estadual de Campinas, Rua Monteiro Lobato 255, Campinas, São Paulo, 13083–862, Brazil 2 REV_HeadB=REV_HeadC=REV_HeadB=REV_HeadC/HeadB Instituto de Biologia, Universidade Federal de Uberlândia, Rua Ceará s/n, Uberlândia, Minas Gerais, 38400–902, Brazil REV_HeadC=REV_HeadD=REV_HeadC=REV_HeadD/HeadC 3Departamento de Botânica, Universidade Federal do Rio Grande do Sul, Avenida Bento Gonçalves REV_Extract3=REV_HeadA=REV_Extract1=REV_HeadA 9500, Porto Alegre, Rio Grande do Sul, 91509–900, Brazil BOR_HeadA=BOR_HeadB=BOR_HeadA=BOR_HeadB/HeadA 4Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458–5126, USA BOR_HeadB=BOR_HeadC=BOR_HeadB=BOR_HeadC/HeadB 5Departamento de Botânica, Universidade Federal do Paraná, Avenida Coronel Francisco H. dos Santos BOR_HeadC=BOR_HeadD=BOR_HeadC=BOR_HeadD/HeadC 100, Curitiba, Paraná, 81531–970, Brazil BOR_Extract3=BOR_HeadA=BOR_Extract1=BOR_HeadA Received 8 April 2020; revised 3 August 2020; accepted for publication 14 January 2021 EDI_HeadA=EDI_HeadB=EDI_HeadA=EDI_HeadB/HeadA EDI_HeadB=EDI_HeadC=EDI_HeadB=EDI_HeadC/HeadB EDI_HeadC=EDI_HeadD=EDI_HeadC=EDI_HeadD/HeadC Microlicieae are a monophyletic tribe comprising seven genera: Chaetostoma, Lavoisiera, Microlicia s.s., Poteranthera, EDI_Extract3=EDI_HeadA=EDI_Extract1=EDI_HeadA Rhynchanthera, Stenodon and Trembleya. Microlicia s.s. includes 172 species predominantly distributed in the campo rupestre of Brazil. Its delimitation is complex because the generic boundaries, mostly with Lavoisiera and CORI_HeadA=CORI_HeadB=CORI_HeadA=CORI_HeadB/HeadA Trembleya, are unclear. Here we present a phylogenetic analysis for Microlicieae focusing on Microlicia s.s., with the CORI_HeadB=CORI_HeadC=CORI_HeadB=CORI_HeadC/HeadB specific goals: (1) to test the monophyly of Microlicia s.s.; (2) to investigate morphological characters that could help CORI_HeadC=CORI_HeadD=CORI_HeadC=CORI_HeadD/HeadC in circumscribing clades and/or genera in the tribe and (3) to provide an appropriate classification for Microlicia CORI_Extract3=CORI_HeadA=CORI_Extract1=CORI_HeadA s.s. and related genera. This study was based on plastid (atpF-atpH, trnS-trnG), nuclear ribosomal (nrITS, nrETS) and nuclear low-copy (waxy) DNA sequences, through maximum likelihood and Bayesian inference analyses. ERR_HeadA=ERR_HeadB=ERR_HeadA=ERR_HeadB/HeadA The history of 12 morphological characters was estimated based on ancestral state reconstruction analyses. Our ERR_HeadB=ERR_HeadC=ERR_HeadB=ERR_HeadC/HeadB analysis shows Microlicia s.s. to be paraphyletic with Chaetostoma, Lavoisiera, Stenodon and Trembleya nested in ERR_HeadC=ERR_HeadD=ERR_HeadC=ERR_HeadD/HeadC it. Most characters traditionally used to diagnose these genera are homoplastic. We propose the inclusion of these four genera in a broadly circumscribed Microlicia s.l., and provide new combinations and names for their species. ERR_Extract3=ERR_HeadA=ERR_Extract1=ERR_HeadA As here defined, Microlicieae has three genera, Rhynchanthera, Poteranthera and Microlicia s.l., Microlicia s.l. INRE_HeadA=INRE_HeadB=INRE_HeadA=INRE_HeadB/HeadA being the fourth richest genus in Melastomataceae with c. 245 species. INRE_HeadB=INRE_HeadC=INRE_HeadB=INRE_HeadC/HeadB ADDITIONAL KEYWORDS: campo rupestre – Chaetostoma – Lavoisiera – paraphyletic – phylogenetic – INRE_HeadC=INRE_HeadD=INRE_HeadC=INRE_HeadD/HeadC Poteranthera – Rhynchanthera – Stenodon – treespace – Trembleya. INRE_Extract3=INRE_HeadA=INRE_Extract1=INRE_HeadA App_Head=App_HeadA=App_Head=App_HeadA/App_Head BList1=SubBList1=BList1=SubBList INTRODUCTION phylogenetic relationships among species that can then be used to construct classifications consistent with BList1=SubBList3=BList1=SubBList2 Taxonomic classifications based on classical evolutionary processes and taxa that share common morphological approaches do not always reflect the SubBList1=SubSubBList3=SubBList1=SubSubBList2 ancestors; this phylogenetic approach based on molecular natural relationships between organisms (Egan, SubSubBList3=SubBList=SubSubBList=SubBList data has been effective in informing the classification Vatanparast & Cagle, 2016). Molecular sequences of large and complex groups such as Myrtaceae (Lucas SubSubBList2=SubBList=SubSubBList=SubBList provide an independent source of data for understanding SubBList2=BList=SubBList=BList et al., 2007, 2018; Mazine et al., 2014; Bünger et al., 2016; Amorim et al., 2019), Orchidaceae (Chase et al., 2015; *Corresponding author. E-mail: [email protected] Tang et al., 2015; D’haijère et al., 2019) and Poaceae © 2021 The Linnean Society of London, Botanical Journal of the Linnean Society, 2021, 197, 35–60 35 36 A. F. A. VERSIANE ET AL. (Soreng et al., 2015). In Melastomataceae, it has been flowers solitary or in inflorescences, number of petals, applied successfully to recent advances and taxonomic number of locules in the ovary, capsule dehiscence realignments across different tribes (e.g. Penneys et al., (basipetal or acropetal) and the persistence of the 2010, 2020; Michelangeli et al., 2011, 2013, 2016, 2018; columella (Almeda & Martins, 2001; Fritsch et al., Rocha et al. 2016a; Veranso-Libalah et al., 2017; Bacci, 2004; Fidanza et al., 2013; Martins & Almeda, 2017). Michelangeli & Goldenberg, 2019; Bochorny et al., 2019; The morphological delimitation of the remaining Guimarães et al., 2019). genera, Chaetostoma, Poteranthera, Rhynchanthera Melastomataceae are among the largest tropical and Stenodon, is clearer than between Microlicia plant families worldwide with 177 genera and 5750 s.s., Lavoisiera and Trembleya. Chaetostoma can be species (Michelangeli et al., 2021), and Microlicieae recognized by the crown of bristles on the hypanthium are one of the major tribes in the family. Currently, apex (Koschnitzke & Martins, 2006; Silva et al., 2018). Downloaded from https://academic.oup.com/botlinnean/article/197/1/35/6188938 by guest on 01 October 2021 Microlicieae include 265 species in seven genera: Poteranthera are small annual herbs with glandular Chaetostoma DC. (12 accepted species), Lavoisiera hairs on the leaf margins (Kriebel, 2012; Rocha et al., DC. (41 species), Microlicia D.Don (172 species), 2016b; Almeda & Pacifico, 2018). Rhynchanthera can Poteranthera Bong. (five species), Rhynchanthera DC. be easily distinguished by its haplostemonous flowers (nom. cons.) (15 species), Stenodon Naudin (two species) with staminodia (Renner, 1990). Finally, Stenodon and Trembleya DC. (18 species) (Renner, 1990; Rocha has thick, woody, decorticating branches and et al., 2016b; Martins & Almeda, 2017; Silva et al., stamens with an inconspicuous ventral appendage 2018; Pacifico et al., 2019; Pacifico & Almeda, 2020; (Fritsch et al., 2004). Pacifico & Fidanza, 2021; Romero et al., 2021) (Fig. 1). Due to its high diversity, Microlicia s.s. shows non- Microlicieae are monophyletic (Clausing & Renner, exclusive and/or polymorphic characters that are 2001; Fritsch et al., 2004; Michelangeli et al., 2013; usually not informative for its recognition, particularly Rocha et al., 2016a, b), with a recent diversification when compared to Lavoisiera and Trembleya. At and a distribution nearly endemic to Brazil (Fritsch present, Microlicia s.s. can be recognized by its et al., 2004; Simon et al., 2009). solitary flowers that are usually pentamerous (seldom Microlicia s.s. is the largest genus of Microlicieae, hexamerous or octamerous), the hypanthium apex with > 170 species and a predominant distribution in lacking bristles, isomorphic to dimorphic stamens, the campo rupestre and cerrado of Brazil, both in the tetrasporangiate or polysporangiate anthers, three Brazilian Cerrado domain (Romero et al., 2021). Ten locular ovary with a glabrous apex, and fruits with species can be found in the Andes and Guiana Shield basipetal dehiscence and a deciduous columella areas; these occur in Bolivia (Microlicia arenariifolia (Almeda & Martins, 2001; Romero, 2003). DC., M. weddellii Naudin and M. woodii R.B.Pacifico, Most of our current phylogenetic understanding Almeda & Fidanza), Colombia (M. colombiana on the tribe stems from Fritsch et al. (2004). In this Humberto-Mend. & R.Romero), Peru (M. sphagnicola work, Rhynchanthera was recovered as sister to the Gleason), Venezuela (M. guanayana Wurdack), Bolivia remaining genera, and Lavoisiera and Trembleya and Peru (M. peruviana Cogn.), Bolivia and Brazil (M. were recovered as well-supported clades. The insignis Schtl. and M. windischii Versiane, D.Nunes position of the remaining genera, Chaetostoma, & R.Romero), and Brazil, Guyana and Venezuela (M. Microlicia s.s. and Stenodon, were weakly supported. benthamiana Triana) (Wurdack, 1958; Romero, 2003; This same study also showed that more samples Mendoza-Cifuentes et al., 2019;
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