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Psba-Trnh and Trnl-F) DNA Sequences See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/233559424 A comprehensive phylogenetic analysis of Eriocaulaceae: Evidence from nuclear (ITS) and plastid (psbA-trnH and trnL-F) DNA sequences Article in Taxon · April 2010 DOI: 10.2307/25677597 CITATIONS READS 61 514 7 authors, including: Maria José Gomes de Andrade Ana Maria Giulietti Universidade Estadual da Bahia 173 PUBLICATIONS 3,523 CITATIONS 26 PUBLICATIONS 564 CITATIONS SEE PROFILE SEE PROFILE Alessandro Rapini Luciano Queiroz Universidade Estadual de Feira de Santana Universidade Estadual de Feira de Santana 115 PUBLICATIONS 2,088 CITATIONS 214 PUBLICATIONS 6,862 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: Phylogeography of orchids of subtribe Laeliinae View project Plant phylogeography and the origins of vegetation diversity in the "campos rupestres" (rocky outcrops) in the Chapada Diamantina View project All content following this page was uploaded by Cassio van den Berg on 22 May 2014. The user has requested enhancement of the downloaded file. TAXON 59 (2) • April 2010: 379–388 Andrade & al. • Phylogenetics of Eriocaulaceae A comprehensive phylogenetic analysis of Eriocaulaceae: Evidence from nuclear (ITS) and plastid (psbA-trnH and trnL-F) DNA sequences Maria José Gomes de Andrade,1 Ana Maria Giulietti,1 Alessandro Rapini,1 Luciano Paganucci de Queiroz,1 Adilva de Souza Conceição,1,2 Paulo Ricardo Machado de Almeida1 & Cássio van den Berg1 1 Programa de Pós-Graduação em Botânica and Laboratório de Sistemática Molecular de Plantas, Departamento de Ciências Biológicas, Universidade Estadual de Feira de Santana, Av. Transnordestina s/n. Novo Horizonte, 44.036-900, Feira de Santana, Bahia, Brazil 2 Departamento de Ciências, Centro de Ensino Superior de Paulo Afonso, Universidade do Estado da Bahia, Rua do Gangorra 503, CHESF, 48.620-410, Paulo Afonso, Bahia, Brazil Author for correspondence: Maria José Gomes de Andrade, [email protected] Abstract The Eriocaulaceae are easily recognized because of the small unisexual flowers in long-pedunculate heads and spiraperturate pollen grains. Their monophyly has never been disputed but internal relationships within the family have not been broadly explored and genera are typically distinguished by few floral characters. Here, we present the first comprehensive phylogenetic study of Eriocaulaceae based on individual and combined molecular datasets, including the plastid psbA-trnH and trnL-F and the nuclear ITS. Results are largely congruent among DNA regions and support the internal dichotomy between the two subfamilies: Eriocauloideae and Paepalanthoideae. Eriocaulon and Leiothrix are strongly supported as monophyletic, whereas Paepalanthus, Blastocaulon, and Syngonanthus are not monophyletic. The phylogenetic nature of Actinocephalus is not resolved and Lachnocaulon, Mesanthemum, Philodice, Rondonanthus, and Tonina (monospecific) are represented in our analyses by a single species each. Based on our results, we suggest two principal generic realignments in the family. The first is the division of Syngonanthus in two genera: Syngonanthus s.str., including Philodice (conserving the former name against the latter), and Comanthera, which is being reinstated to include two sections segregated from Syngonanthus: S. sect. Eul- epis and S. sect. Thysanocephalus. The second realignment suggested is the amalgamation of Actinocephalus, Blastocaulon, Lachnocaulon, and Tonina into Paepalanthus s.l. Three ‘stat. nov.’ within Paepalanthus are published (P. ser. Leptocephali, P. ser. Rosulati, and P. ser. Dimeri). Keywords molecular phylogeny; Paepalanthus; Syngonanthus; systematics; taxonomy INTRODUCTION with isostemonous flowers and eglandular petals, including: Blastocaulon Ruhland, Lachnocaulon Kunth, Leiothrix Ruh- The Eriocaulaceae are a monophyletic family classified in land, Paepalanthus Mart., Philodice Mart., Syngonanthus Ruh- Poales (Chase & al., 1995; Linder & Kellogg, 1995; Stevenson land, and Tonina Aubl. Six other genera have been proposed & Loconte, 1995; Givnish & al., 1999; Giulietti & al., 2000; after Ruhland’s treatment: Moldenkeanthus P. Morat (sy nony- Bremer, 2002; APG III, 2009). They are easily characterized mized to Paepalanthus ; Stützel, 1987), Wurdackia Moldenke by graminoid leaves, small unisexual flowers grouped in dense (synonymized to Rondonanthus Herzog; Hensold & Giulietti, capitula, a 3- or 2-locular ovary (with a single pendulous ovule 1991), Comanthera L.B. Sm. and Carptotepala Moldenke per locule), and having spiraperturate p ollen grains (Giulietti & (synonymized to Syngonanthus; Giulietti & Hensold, 1991), al., 1995, 2000). The family includes eleven genera and about Rondonanthus and Actinocephalus (Körn.) Sano. 1400 species occurring primarily in the Neotropics, especially Phylogenetic studies in Eriocaulaceae have been previ- in mountain areas or high plateaus. The Eriocaulaceae are more ously carried out based on morphological (Giulietti & al., diverse in savanna and “campos rupestres” vegetation in Brazil 1995, 2000) and unpublished molecular data (Unwin, 2004; and in “tepuis” in Guayana shield of Venezuela and Guyana. Andrade, 2007). Giulietti & al. (1995) undertook a preliminary The inter- and infrageneric relationships in Eriocaulaceae cladistic analysis based on 29 morphological characters and are not well resolved and the traditional generic circumscription eleven taxa representing eight genera, which was broadened remains based on few floral characters. These include: (1) de- to 49 morpho-anatomical characters and 25 taxa in Giulietti gree of petal union in pistillate flowers, (2) presence of glands & al. (2000). Unwin (2004) performed a phylogenetic analy- on petals, (3) number of stamens, and (4) number of pollen-sacs sis of Eriocaulaceae using sequences of trnL intron (58 taxa), per anther (Ruhland, 1903; Hensold, 1988; Giulietti & al., 1995, trnL-F spacer (58), and the gene ndhF (21), but Mesanthemum, 2000; Stützel, 1998). The last complete revision of the family Rondonanthus, and Philodice were not sampled and sampling (Ruhland, 1903) recognized two subfamilies: Eriocauloideae, within more diverse genera, such as Paepalanthus and Syngo- with diplostemonous flowers and glandular petals, including nanthus, was thin. Andrade (2007) performed the phylogenetic Eriocaulon L. and Mesanthemum Körn.; and Paepalanthoideae, analysis of Neotropical genera of the family. Therefore, the 379 Andrade & al. • Phylogenetics of Eriocaulaceae TAXON 59 (2) • April 2010: 379–388 present study aims at re-evaluating Eriocaulaceae classification were purified by enzymatic treatment using the EXOSAP-IT based on comprehensive molecular phylogenetic analyses using kit (GE Healthcare, USB, Cleveland, Ohio, U.S.A.). Sequenc- the nuclear ITS (ITS 1, 5.8S rRNA and ITS 2) and the plastid ing reactions were performed using the Big Dye Terminator regions trnL-F (trnL intron and trnL-trnF intergenic spacer) kit, version 3.1 (Applied Biosystems, Foster City, California, and psbA-trnH spacer. U.S.A.). Samples were sequenced in both directions using a SpectruMedix SCE2410 automatic sequencer according to the protocols supplied by the manufacturer. MATERIALS AND METHODS Sequence alignment and matrices. — Forward and re- verse electropherograms were superimposed and edited using Taxa sampled. — A total of 81 species (82 samples) was the STADEN Package software (Staden & al., 1998). The se- included in the analyses; all were sequenced for this study. The quences of psbA-trnH and trnL-F were subsequently aligned outgroup was composed of Abolboda pulchella Bonpl., Xyris using CLUSTAL X v.1.81 (Thompson & al., 1997). The initial anceps Lam., X. jupicai L.C. Rich., and X. lagoinhae Kral & alignment was then corrected manually according to the crite- L.B. Sm. (Xyridaceae). The 77 species of the ingroup represent ria of Kelchner (2000) for plastid regions. ITS alignment was all recognized genera of Eriocaulaceae and a great part of the in- performed exclusively by eye. Indels were considered missing frageneric taxa, according to Ruhland (1903). The rate of species data and regions with dubious alignment were excluded from sampled per genus was: Actinocephalus (6 species sampled/28 analyses. total species), Blastocaulon (3/5), Eriocaulon (5/479), Lachno- Phylogenetic analyses. — Maximum parsimony (MP) caulon (1/7), Leiothrix (9/64), Mesanthemum (1/15), Paepalan- analyses were performed using PAUP* v.4.0b10 (Swofford, thus (38/420), Philodice (1/2), Rondonanthus (1/6), Syngonanthus 2002) with Fitch parsimony (equal weights, unordered; Fitch, (11/199), and Tonina (1/1) (total species number according Go- 1971) as the optimality criterion. We performed individual vaerts, 2006) (Appendix; see this material for taxa authorship (psbA-trnH, trnL-F, ITS) and combined analyses (combined and three new series names in Paepalanthus). plastid and combined plastid + nuclear). For MP analyses, a Extraction, amplification, and sequencing of DNA. — heuristic search with 1000 replicates using random taxon-ad- DNA was extracted from silica-gel–dried leaves (5–15 mg) dition and tree bisection reconnection (TBR) swapping were or, in a few cases, fresh leaves or herbarium material using a performed, saving no more than 15 trees per replicate to pre- modified version of 2× CTAB (Doyle & Doyle, 1987) protocol. vent extensive swapping on large suboptimal islands. Trees The region psbA-trnH was amplified in a final
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