Systematics of Dasyphyllum (Asteraceae)
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Instituto de Pesquisas Jardim Botânico do Rio de Janeiro Escola Nacional de Botânica Tropical Programa de Pós-Graduação em Botânica Systematics of Dasyphyllum (Asteraceae) Mariana M. Saavedra Elsie F. Guimarães e Rafaela C. Forzza Barnadesioideae • Synapomorphies: – axillary spines – barnadesioids trichomes • Restricted distribution in South America • 9 genus and 85 species – Dasyphyllum (33 spp.) – Chuquiraga (23 spp.) – Barnadesia (19 spp.) – Arnaldoa (3 spp.) – Doniophyton (2 spp.) – Fulcaldea (2 spp.) – Duseniella, Huarpea e Dasyphyllum vagans Schlechtendalia (1 spp.) Funk et al. 2009 Dasyphyllum Kunth Kunth (1820) – D. argenteum Cabrera (1959) – Dasyphyllum • 36 species (34 new combinations) Infrageneric classification sensu Cabrera • Trees Dasyphyllum subg. Archydasyphyllum • Pinnate leaves • Anthers’ apical appendage • 2 species obtuse or emarginate • D. diacanthoides (Less.) Cabrera • D. excelsum (D.Don) Cabrera) D. diacanthoides Stuessy et al. 1996 Dasyphyllum sensu Cabrera (1959) • Trees or shrubs Dasyphyllum subg. Dasyphyllum • Trinerved leaves • Anthers’ apical appendages bilobed Section Microcephala (=Dasyphyllum) Section Macrocephala • Small and numerous heads • Large and solitaries or few heads • 27 species, 6 varieties • 11 species, 2 varieties D. sprengelianum Stuessy et al. 1996 Stuessy et al. 1996 D. spinescens Morphological phylogeny Urtubey & Stuessy 2001 • 52 of 88 species of the subfamily • 29 spp. Dasyphyllum • 31 morphological characters • 13 characters are informative to Dasyphyllum Infrageneric classification of Dasyphyllum: * = subg. Dasyphyllum sect. Dasyphyllum ** = subg. Dasyphyllum sect. Macrocephala *** = subg. Archydasyphyllum Molecular phylogeny Gustafsson et al. (2001) Dasyphyllum is paraphyletic Both subgenera are monophyletic Corroborated by Gruenstaeudl et al. (2009) Molecular phylogeny Gustafsson et al. (2001) Dasyphyllum subg. Dasyphyllum Dasyphyllum hystrix Dasyphyllum sect. Macrocephala is monophyletic Objectives • To propose a hypothesis of phylogenetic relationship among the genus’ species as a tool for testing the infrageneric classification • Taxonomic treatment of Dasyphyllum – Revision of Dasyphyllum sect. Macrocephala – Synopsis of Dasyphyllum sect. Dasyphyllum (=Microcephala). D. spinescens D. brasiliense Phylogeny • Morphological 39 taxa 67 characters 215 trees CI = 0,29 RI = 0,53 Dasyphyllum sect. Macrocephala Dasyphyllum sect. Dasyphyllum Dasyphyllum diacanthoides/ D. excelsum Macrocephala Strict consensus Majority consensus Phylogeny • Combined (ITS1, 5.8S e ITS2 - Gustafsson et al. 2001) 21 taxa 733 characters 36 trees CI = 0,60 RI = 0,68 Heuristic search – strict consensus branch and bound Taxonomic treatment of Dasyphyllum 33 species, with 5 new species 2 sections, 3 species of uncertain position 153 binomials, 72 basionyms and 95 typi 29 new synonyms proposed 28 lectotypi 3 neotypi 2 epitypi Dasyphyllum D. vagans D. reticulatum D. spinescens D. spinescens D. vagans Dasyphyllum D. globosum D. trichophyllum D. diacanthoides Acrodromous basal Acrodromous suprabasal Pinnate Dasyphyllum D. littorale D. diamantinense D. sprengelianum D. donianum D. leptacanthum Dasyphyllum style anthers Dasyphyllum Monoecious Gynodioecious Revision of Dasyphyllum sect. Macrocephala 6 species typifications: 10 lectotypi 1 epitypus Stuessy et al. 1996 Morphology Monoecious plants D. cryptocephalum (Baker) Cabrera D. donianum (Gardner) Cabrera Involucre >25mm = D. infundibulare (Baker) Cabrera Solitary heads = D. latifolium (Gardner) Cabrera Leaves 3 or 5 primary D. fodinarum (Gardner) Cabrera ribs D. reticulatum (DC.) Cabrera = D. reticulatum var. robustum Domke = D. lanosum Cabrera D. sprengelianum (Gardner) Cabrera = D. candolleanum (Gardner) Cabrera = D. sprengelianum var. inermis (Gardner) Cabrera = D. velutinum (Baker) Cabrera D. trichophyllum (Baker) Cabrera D. cryptocephalum D. fodinarum D. trichophyllum Botumirim (MG) Serra do Caraça (MG) D. reticulatum D. donianum D. sprengelianum Gouveia (MG) Correntina (BA) Itapanhoacanga (MG) Synopsis of D. brasiliense (Spreng.) Cabrera = D. tomentosum (Spreng.) Cabrera Dasyphyllum sect. Dasyphyllum = D. orthacanthum (DC.) Cabrera D. flagellare (Casar.) Cabrera D. floribundum (Gardner) Cabrera 24 species D. lanceolatum (Less.) Cabrera 14 spp. in Brazil (5 sp. nov.) = D. synacanthum (Baker) Cabrera D. leptacanthum (Gardner) Cabrera 10 spp. in Andes D. retinens (S.Moore) Cabrera D. spinescens (Less.) Cabrera typifications D. vagans (Gardner) Cabrera 15 lectotypi D. varians (Gardner) Cabrera 3 neotypi D. argenteum Kunth 1 epitypus = D. lehmannii (Hieron.) Cabrera Morphology D. armatum (Koster) Cabrera D. brevispinum Sagást. & M.O. Dillon Ginodioecious plants D. colombianum (Cuatr.) Cabrera D. ferox (Wedd.) Cabrera Involucre <20mm D. inerme (Rusby) Cabrera Numerous heads D. leiocephalum (Wedd.) Cabrera = D. horridum (Muschl.) Cabrera Leaves 3 primary ribs D. maria-lianae Zardini & Soria D. popayanense (Hieron.) Cabrera = D. cabrerae Sagást. = D. weberbaueri (Tovar) Cabrera D. vepreculatum (D.Don) Cabrera Stuessy et al. 1996 Ligulate corolla Internal pubescence in the basal part of the corolla tube (ex. D. capixaba) Anthers apical appendages bilobed D. capixaba Saavedra sp. nov. ined. D. spinescens Scandent plants Cerrado, Floresta Atlântica, Chaco Leaves with pubescent abaxial surface Ligulate corolla, short lobes Internal pubescence throughout the entire surface of the corolla tube Anthers apical appendages bilobed D. globosum Saavedra D. minutiflorum Saavedra sp. nov. ined. sp. nov. ined. Cerrado, Chaco e Floresta Atlântica Scandent plants Ligulate corolla, long or short lobes Internal pubescence throughout the entire surface of the corolla tube Anthers apical appendages emarginated D. lanceolatum D. floribundum D. vepreculatum Erect plants Cilindric heads Corolla generally tubular, long lobes Internal pubescence throughout the entire surface of the corolla tube Anthers apical appendages emarginated Andes, Cerrado/Caatinga, Floresta Atlântica D. littorale Saavedra sp. nov. ined. D. diamantinense Saavedra & M.Monge sp. nov ined. Incertae sedis Dasyphyllum diacanthoides (Less.) Cabrera Dasyphyllum excelsum (D.Don) Cabrera Dasyphyllum hystrix (Wedd.) Cabrera Conclusions • Geographic distribution: South America, 0-4300m alt. – Atlantic Forest – Cerrado – Caatinga – Chaco – Andes • Species diversity: – Brazil (20) – Bolivia (7) – Argentina (5) – Paraguay (5) – Peru (5) – Chile (2) – Colombia (2) – Equador (2) – Venezuela (1) • Ginodioecious – Dasyphyllum sect. Dasyphyllum (except D. capixaba), D. diacanthoides, D. excelsum e D. hystrix • Monoecious – Dasyphyllum sect. Macrocephala and the other genus of Barnadesioideae. Conclusions • Monophyly of Dasyphyllum sect. Macrocephala • Molecular data of D. capixaba lacking morphological combined D. capixaba Conclusions • Monophyly of Dasyphyllum sect. Dasyphyllum excluding D. hystrix • Morphological data of Arnaldoa e Fulcaldea lacking • Combined analysis with more species can contribute to solve of Incertae sedis taxa morphological combined D. hystrix D. hystrix Acknowledgments .