Molecular Phylogeny, Origin and Taxonomic Implications of the Tribe Cacteae (Cactaceae)
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This article was downloaded by: [UNAM Ciudad Universitaria] On: 04 April 2013, At: 17:45 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Systematics and Biodiversity Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tsab20 Molecular phylogeny, origin and taxonomic implications of the tribe Cacteae (Cactaceae) Monserrat Vázquez-Sánchez a , Teresa Terrazas a , Salvador Arias a & Helga Ochoterena a a Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70–233, México, 04510, DF, México Version of record first published: 03 Apr 2013. To cite this article: Monserrat Vázquez-Sánchez , Teresa Terrazas , Salvador Arias & Helga Ochoterena (2013): Molecular phylogeny, origin and taxonomic implications of the tribe Cacteae (Cactaceae), Systematics and Biodiversity, 11:1, 103-116 To link to this article: http://dx.doi.org/10.1080/14772000.2013.775191 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Systematics and Biodiversity (2013), 11(1): 103–116 Research Article Molecular phylogeny, origin and taxonomic implications of the tribe Cacteae (Cactaceae) MONSERRAT VAZQUEZ-S´ ANCHEZ,´ TERESA TERRAZAS, SALVADOR ARIAS & HELGA OCHOTERENA Instituto de Biolog´ıa, Universidad Nacional Autonoma´ de Mexico,´ Apartado Postal 70–233, Mexico´ 04510, DF, Mexico´ (Received 13 October 2012; revised 25 January 2013; accepted 4 February 2013) This study aimed to test the phylogenetic relationships of the tribe Cacteae, the generic circumscription within the tribe, in particular, the monophyly of the genus Ferocactus, and to provide a biogeographical hypothesis about the origin of Cacteae. The analysis included 135 species from all of the 27 accepted genera and four outgroup species. Five chloroplast regions were sequenced, aligned, and coded postulating gaps, simple sequence repeats (SSRs), and inversions as potential synapomorphies, and their contributions to phylogenetic reconstruction were evaluated. The phylogenetic analyses recovered 63% of the genera as monophyletic. The contribution of rpl16, trnL-F and psbA to the phylogenetic signal was higher than in the two more slowly evolving genes (rbcL, matK), but the gaps and SSRs supported some of the genera. This result differs from those of previous phylogenetic studies in which less than 35% of the genera were recovered as monophyletic. In this work, Astrophytum and Echinocactus were re-circumscribed with five and four species, respectively. Turbinicarpus was found to be polyphyletic; 11 species correspond to Turbinicarpus s.str., whereas a highly supported clade corresponded to Rapicactus, and three species need further study. Contrary to its current circumscription, Ferocactus was not supported as monophyletic because it is polyphyletic concerning Glandulicactus, Leuchtenbergia, Stenocactus and Thelocactus. We recognize this group of genera as the Ferocactus clade in which the species share the presence of scales in the pericarpel and ribbed stems, whether tuberculated or not. The Cacteae seem to have originated in the Sierra Madre Oriental and then dispersed to the Mexican Plateau, where radiation and diversification occurred at the boundaries of the Miocene–Pliocene Epoch. The development of the Mexican Plateau and the Trans-Mexican Volcanic Belt may have favoured the isolation of the Cacteae. A taxonomic diagnosis is presented for the tribe Cacteae and 18 genera that we now recognize. Key words: alignment coding, Ferocactus, Rapicactus, rpl16, taxonomy, Turbinicarpus Introduction at the stem tip and on the pericarpel and floral tube. The The recognition of the tribe Cacteae as a monophyletic second, Mammillaria, represents one of the largest genera of North American cacti (c. 150–180 spp.); they are low group is currently indisputable (Barcenas´ et al., 2011; growing, usually globose, and have distinctly tuberculate Downloaded by [UNAM Ciudad Universitaria] at 17:45 04 April 2013 Hernandez-Hern´ andez´ et al., 2011); however, the limits between genera, the number of species within each genus, stems. Both of these genera were artificially delimited in and their relationships to one another remain matters of the main classification systems of the 19th century. For ex- debate (Barthlott & Hunt, 1993; Butterworth et al., 2002; ample, Schumann (1898) recognized only five genera in the present tribe Cacteae. These five were split into smaller Barcenas´ et al., 2011). For example, during the past decade, 24 to 27 genera (including 314 to 384 spp.) have been recog- genera in subsequent years based on the combination of structural stem characters (e.g. simple vs. shrubby; prune- nized (Table 1; Anderson, 2001; Guzman´ et al., 2003; Hunt et al., 2006). The Cacteae was based on two major genera ribs vs. prune-tubercles), flowers (e.g. presence or absence according to the taxonomic revision by Butterworth et al. of areoles, scales and/or trichomes), fruits (e.g. fleshy vs. (2002). The first, Echinocactus, has ribbed stems and rep- dry; type of dehiscence), and seeds (e.g. shape, colour, size, resents the most typical cacti of the deserts of Mexico and microrelief), until molecular methods were integrated with the USA. It is characterized by dense masses of trichomes phylogenetic studies (Metzing & Kiesling, 2008). Butterworth et al. (2002) provided the first phylogenetic Correspondence to: Monserrat Vazquez-S´ anchez.´ E-mail: analysis to test the monophyly of the tribe, including 26 [email protected] genera and 62 species, of which 15 belong in the genus ISSN 1477-2000 print / 1478-0933 online C 2013 The Natural History Museum http://dx.doi.org/10.1080/14772000.2013.775191 104 M. Vazquez-S´ anchez´ et al. Table 1. The number of genera recognized within the tribe Cacteae in the most recent classifications and this paper. Anderson, 2001 (26 genera) Guzman´ et al., 2003 (28) Hunt et al., 2006 (24) This paper (18) Acharagma Acharagma Acharagma Acharagma (2 spp.) Ariocarpus Ariocarpus Ariocarpus Ariocarpus (6) Astrophytum Astrophytum Astrophytum Astrophytum (5) Aztekium Aztekium Aztekium Aztekium (2) Cochemiea Cochemiea Cochemiea (3) Coryphantha Coryphantha Coryphantha Cumarinia Cumarinia Cumarinia (1) Digitostigma Echinocactus Echinocactus Echinocactus Echinocactus (4) Echinomastus Echinomastus Epithelantha Epithelantha Epithelantha Epithelantha (1) Escobaria Escobaria Escobaria Ferocactus Ferocactus Ferocactus Geohintonia Geohintonia Geohintonia Geohintonia (1) Glandulicactus Leuchtenbergia Leuchtenbergia Leuchtenbergia Leuchtenbergia (1) Lophophora Lophophora Lophophora Lophophora (2) Mammillaria Mammillaria Mammillaria Mammillaria Mammilloydia Mammilloydia Neolloydia Neolloydia Neolloydia Obregonia Obregonia Obregonia Obregonia (1) Ortegocactus Ortegocactus Ortegocactus Ortegocactus (1) Pediocactus Pediocactus Pelecyphora Pelecyphora Pelecyphora Pelecyphora (2) Sclerocactus Sclerocactus Sclerocactus Rapicactus (5) Stenocactus Stenocactus Stenocactus Strombocactus Strombocactus Strombocactus Strombocactus (2) Thelocactus Thelocactus Thelocactus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus (2) Mammillaria. They used only the intron rpl16 and coded 2002; Hernandez-Hern´ andez´ et al., 2011) or were based insertions and deletions that were considered phylogeneti- on only a single molecular marker (Barcenas´ et al., 2011). cally informative. Nevertheless, they obtained a highly un- Therefore, a comprehensive study that includes many taxa resolved topology, most likely because of low character and and more than one molecular marker that aims to test the taxon sampling. With the exception of Mammillaria, which generic circumscription within the Cacteae, with emphasis has been the subject of phylogenetic studies (Butterworth on Ferocactus, is still lacking. & Wallace, 2004; Harpke & Peterson, 2006), the remain- The aim of this study was to test the monophyly of the der of the Cacteae genera with greater taxonomic diversity, genera within the tribe Cacteae, particularly emphasizing Downloaded by [UNAM Ciudad Universitaria] at 17:45 04 April 2013 such as Coryphantha (c. 43 spp.), Escobaria (c. 20 spp.), the study of Ferocactus. For gaining a higher phylogenetic Ferocactus (c. 28 spp.), Stenocactus (c. 18 spp.), Thelo- resolution, we included a greater representation of each cactus (c. 12 spp.) and Turbinicarpus (c. 21 spp.), have not taxon from each genus within the Cacteae, and we used been studied well enough to establish phylogenetic relation- five cp-DNA molecular markers (matK, rbcL, psbA-trnH, ships. Phylogenetic analyses have only been conducted in rpl16 and trnL-F) coding gaps, and