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Assessing Azorella (Apiaceae) and its allies: Phylogenetics and a new classification

Abstract

Phylogenetic analyses of nuclear rDNA spacers (ITS and ETS) from Azorella and five closely related genera confirm earlier plastid results indicating that Azorella, Huanaca, Mulinum, and Schizeilema are all polyphyletic, and that the monotypic genus Laretia is nested within one of the six subclades of Azorella. Only Stilbocarpa is monophyletic, but that genus is embedded within a larger clade that includes representatives of three other genera (Azorella, Huanaca, and Schizeilema). Both nuclear and plastid datasets identify the same 10 clades, but the placement of these clades remains unstable. A new classification is presented in which these six genera are reduced to a single genus (Azorella) comprising 58 species arranged in 10 sections, one of which is newly described here (Azorella sect. Ranunculus), and one lectotypified (Azorella sect. Glabratae). A total of 13 new combinations are made (A. albovaginata, A. allanii, A. boelckei, A. burkartii, A. colensoi, A. echegarayi, A. echinus, A. hallei, A. lyallii, A. polaris, A. prolifera, A. robusta, A. ulicina), along with three replacement names (A. atacamensis, A. ruizii, A. schizeilema). For each of the 58 accepted species, a full synonymy is provided along with geographic ranges (and nomenclatural notes, where useful).

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Acknowledgments

The authors thank the following institutions for providing assistance in obtaining plant samples and facilitating our herbarium studies: Allan Herbarium at Landcare Research (CHR), Australian National Botanic Garden (CBG), Australian National Herbarium (CANB), Field Museum of Natural History (F), Fundación Miguel Lillo (LIL), Instituto de Botánica Darwinion (SI), Instituto Nacional de Biodiversidad (INB), Missouri Botanical Garden (MO), Museo de Ciencias Naturales de Salta (MCNS), Museo de La Plata (LP), Muséum National d'Histoire Naturelle (P), Museum of New Zealand Te Papa Tongarewa (WELT), New York Botanical Garden (NY), Pontificia Universidad Católica del Ecuador (QCA), Royal Botanic Gardens Kew (K), Royal Botanic Gardens Victoria (MEL), Royal Botanic Gardens Sydney (NSW), Smithsonian Tropical Research Institute (STRI), United States National Herbarium (US), Universidad Autónoma de Chiriquí (UCH), Universidad de Concepción (CONC), Universidad de Magallanes (HIP), Universidad de Panamá (PMA), Universidad Nacional de Colombia (COL), Universidad Nacional del Comahue (BCRU), Universidad Nacional Mayor de San Marcos (USM), Universidad Nacional de Jujuy (JUA), University of Waikato (WAIK). We gratefully acknowledge the assistance of Pete Lowry, Roy Gereau, and Larry Kelly for their expertise and advice in interpretating several complex nomenclatural issues. We also thank Eduardo Ruiz Ponce, Oscar Toro, Asunción Cano Echevarría, Susy Castillo Ramón, Favio Gonzalez Garavito, Julián Aguirre Santoro, Julio C. Betancur, Katya Romoleroux, Fernanda Samaniego, Alejandra Vasco, Anabela Plos, Carolina Calviño, Cecelia Ezcurra, Susanna Martínez, Isler Chinchilla, and Zabdy Samudio for field and herbarium assistance, and Greg Chandler, Larry Hufford, and Jun Wen for providing leaf tissue samples. Funding for field and laboratory work was provided by the National Science Foundation (DEB 0949819), the National Geographic Society (CRE 8355-07), the Department of Biology and the Integrative Life Sciences Program at Virginia Commonwealth University, and the Cullman Program for Molecular Systematics at the New York Botanical Garden.

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Correspondence to Gregory M. Plunkett.

Appendix 1

Appendix 1

List of species, voucher information (collector and number, and herbarium of deposition), localities, and GenBank accession numbers.

Taxon name; voucher; locality; GenBank accessions: ITS; ETS.

OUTGROUP: Diplaspis nivis Van den Borre & Henwood; Donaldson 479, CBG; Victoria, Australia; KX645099; KX645022. D. cordifolia (Hook.) Hook.f.; Ratkowsky 195, MO; Tasmania, Australia; KX645100; KX645023. Dickinsia hydrocotyloides Franch.; Wen 5003, F; Sichuan, China; KX645101; KX645024.

INGROUP: Azorella acaulis (Cav.) Drude; Chandler 1125, CANB; Tierra del Fuego, Argentina; KX645082; KX645005. A. albovaginata (Gillies & Hook.) G. M. Plunkett & A. N. Nicolas; Chiapella 294, SI; Neuquén, Argentina; KX645028; KX644951. A. albovaginata (Gillies & Hook.) G. M. Plunkett & A. N. Nicolas; Werderman 613, NY; Curicó, Chile; KX645029; KX644952. A. allanii (Cheeseman) G. M. Plunkett & A. N. Nicolas; Plunkett 2525, NY; North Island, New Zealand; KX645089; KX645012. A. ameghinoi Speg.; Bonifacino 487, US; Santa Cruz, Argentina; KX645080; KX645003. A. andina (Phil.) Drude; Ward s.n., MO; Río Negro, Argentina; KX645081; KX645004. A. aretioides Willd. ex DC.; Cuatrecasas 26920, US; Nariño, Colombia; KX645073; KX644996. A. atacamensis G. M. Plunkett & A. N. Nicolas; Werderman 1009, US; Antofagasta, Chile; KX645033; KX644956. A. atacamensis G. M. Plunkett & A. N. Nicolas; Latorre 206, CONC; Chañaral, Chile; KX645032; KX644955. A. biloba (Schlecht.) Wedd.; Mendoza 620, NY; Santa Cruz, Bolivia; KX645070; KX644993. A. boelckei (Mathias & Constance) G. M. Plunkett & A. N. Nicolas; Calviño 714, BCRU; Chubut, Argentina; KX645083; KX645006. A. burkartii (Mathias & Constance) G. M. Plunkett & A. N. Nicolas; Calviño 705, BCRU; Río Negro, Argentina; KX645084; KX645007. A. compacta Phil.; Chandler 1093, CANB; Salta, Argentina; KX645063; KX644986. A. compacta Phil.; Mendoza 45, NY; Cochabamba, Bolivia; KX645064; KX644987. A. concolor Rendle; Sleumer 1179, US; Santa Cruz, Argentina; KX645025; KX644948. A. corymbosa Pers.; Balslev 3339, NY; Cotopaxi, Ecuador; KX645041; KX644964. A. crassipes Phil.; Ezcurra 3190, SI; Neuquén, Argentina; KX645049; KX644972. A. crassipes Phil.; Calviño 732, BCRU; Neuquén, Argentina; KX645048; KX644971. A. crenata (Ruiz & Pav.) Pers.; Øllgaard 8342, NY; Carchi, Ecuador; KX645074; KX644997. A. cryptantha (Clos) Reiche; Plunkett 2273, NY; Coquimbo, Chile; KX645040; KX644963. A. cryptantha (Clos) Reiche; Fortunato 5536, NY; San Juan, Argentina; KX645039; KX644962. A. cuatrecasasii Mathias & Constance; Barclay 7165, US; Cundinamarca, Colombia; KX645071; KX644994. A. diapensioides A. Gray; Solomon 14236, MO; La Paz, Bolivia; KX645075; KX644998. A. diversifolia Clos; Plunkett 2314, NY; Araucanía, Chile; KX645076; KX644999. A. echegarayi (Hieron.) G. M. Plunkett & A. N. Nicolas; Fortunato 5642, NY; San Juan, Argentina; KX645038; KX644961. A. echinus (DC.) G. M. Plunkett & A. N. Nicolas; Calviño 727, BCRU; Neuquén, Argentina; KX645027; KX644950. A. exigua (Hook.f.) Drude; Nicolas 64, NY; South Island, New Zealand; KX645094; KX645017. A. filamentosa Lam.; Chandler 1123, CANB; Tierra del Fuego, Argentina; KX645078; KX645001. A. fragosea (F. Muell.) Druce; Gray 4790, MO; New South Wales, Australia; KX645087; KX645010. A. fragosea (F. Muell.) Druce; Weber 13730, NY; New South Wales, Australia; KX645088; KX645011. A. fuegiana Speg.; Chandler 1127, CANB; Tierra del Fuego, Argentina; KX645079; KX645002. A. haastii (Hook.f.) Drude; Wardle 94/170, WAIK; South Island, New Zealand; KX645090; KX645013. A. hallei (Skottsb.) G. M. Plunkett & A. N. Nicolas; Zech 21, CONC; Santa Cruz, Argentina; KX645053; KX644976. A. hallei (Skottsb.) G. M. Plunkett & A. N. Nicolas; Raffo 139, BCRU; Santa Cruz, Argentina; KX645052; KX644975. A. hydrocotyloides (Hook.f.) Kirk; Bulloch s.n., WAIK (590); South Island, New Zealand; KX645091; KX645014. A. julianii Mathias & Constance; Gines 4634, US; Mucubají, Venezuela; KX645072; KX644995. A. lyalii (Armstr.) G. M. Plunkett & A. N. Nicolas; Walls s.n., CHR (362049A); Codfish Island, New Zealand; KX645096; KX645019. A. lycopodioides Gaud.; Chandler 1119, CANB; Tierra del Fuego, Argentina; KX645057; KX644980. A. lycopodioides Gaud.; Calviño 711, BCRU; Río Negro, Argentina; KX645056; KX644979. A. macquariensis A. E. Orchard; Jackson 69, CANB; Macquarie Island; KX645061; KX644984. A. madreporica Clos; Calviño 717, BCRU; Chubut, Argentina; KX645067; KX644990. A. microphylla (Cav.) G. M. Plunkett & A. N. Nicolas; Calviño 704, BCRU; Río Negro, Argentina; KX645055; KX644978. A. microphylla (Cav.) G. M. Plunkett & A. N. Nicolas; Cabrera 33128, SI; Chubut, Argentina; KX645054; KX644977. A. monantha Clos; Chandler 1102, CANB; Mendoza, Argentina; KX645058; KX644981. A. monantha Clos; Plunkett 2292, NY; Santiago, Chile; KX645059; KX644982. A. monantha Clos; Chandler 1124, CANB; Tierra del Fuego, Argentina; KX645060; KX644983. A. monteroi S. Martínez & Constance; Montero 11816, CONC; Malleco, Chile; KX645046; KX644969. A. monteroi S. Martínez & Constance; Brownless 1007, CONC; Antuco, Chile; KX645047; KX644970. A. multifida (Ruiz & Pav.) Pers.; Cerón 19429, MO; Bolívar, Ecuador; KX645043; KX644966. A. multifida (Ruiz & Pav.) Pers.; Hutchison 1625, NY; Piura, Peru; KX645042; KX644965. A. nivalis Phil.; Calviño 726, BCRU; Neuquén, Argentina; KX645026; KX644949. A. pallida (Kirk) Kirk; Plunkett 2533, NY; South Island, New Zealand; KX645092; KX645015. A. patagonica Speg.; Zech 20, CONC; Santa Cruz, Argentina; KX645068; KX644991. A. pedunculata (Spreng.) Mathias & Constance; Luteyn 6709, US; Napo, Ecuador; KX645044; KX644967. A. polaris (Hombr. & Jacq.) G. M. Plunkett & A. N. Nicolas; Croft 10437, CBG; Macquarie Island; KX645098; KX645021. A. prolifera (Cav.) G. M. Plunkett & A. N. Nicolas; Chandler 1099, CANB; Mendoza, Argentina; KX645034; KX644957. A. prolifera (Cav.) G. M. Plunkett & A. N. Nicolas; Plunkett 2279, NY; Santiago, Chile; KX645035; KX644958. A. pulvinata Wedd.; Solomon 16611, MO; La Paz, Bolivia; KX645045; KX644968. A. ranunculus d'Urv.; Goodall 719, P; Tierra del Fuego, Argentina; KX645085; KX645008. A. ranunculus d'Urv.; Rodriguez 3273, CONC; Tierra del Fuego, Argentina; KX645086; KX645009. A. robusta (Kirk) G. M. Plunkett & A. N. Nicolas; Johnson s.n., CHR (310924B); New Zealand; KX645097; KX645020. A. roughii (Hook.f.) Kirk; Plunkett 2562, NY; South Island, New Zealand; KX645093; KX645016. A. ruizii G. M. Plunkett & A. N. Nicolas; Teillier 2504, MO; Santiago, Chile; KX645066; KX644989. A. ruizii G. M. Plunkett & A. N. Nicolas; Plunkett 2287, NY Santiago, Chile; KX645065; KX644988. A. schizeilema G. M. Plunkett & A. N. Nicolas; Godley s.n., CHR (134249); Auckland Island, New Zealand; KX645095; KX645018. A. selago Hook.f.; Donaldson 234, CBG; Heard Island; KX645062; KX644985. A. spinosa Ruiz & Pav.; Plunkett 2307, US; Araucanía, Chile; KX645030; KX644953. A. spinosa Ruiz & Pav.; Zöllner 11823, MO; Valparaíso, Chile; KX645031; KX644954. A. trifoliolata Clos; Chandler 1115, CANB; Mendoza, Argentina; KX645069; KX644992. A. trifoliolata Clos; Calviño 719, BCRU; Río Negro, Argentina; KX645077; KX645000. A. trifurcata (Gaertn.) Pers.; Kew 379-81.04150 Cultivated, Royal Botanic Garden Kew; KX645051; KX644974. A. trifurcata (Gaertn.) Pers.; Chandler 1112, CANB; Tierra del Fuego, Argentina; KX645050; KX644973. A. ulicina (Gillies & Hook.) G. M. Plunkett & A. N. Nicolas; Chandler 1092, CANB; Jujuy, Argentina; KX645036; KX644959. A. cf. ulicina (Gillies & Hook.) G. M. Plunkett & A. N. Nicolas; Chandler 1103, CANB; Mendoza, Argentina; KX645037; KX644960.

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Plunkett, G.M., Nicolas, A.N. Assessing Azorella (Apiaceae) and its allies: Phylogenetics and a new classification. Brittonia 69, 31–61 (2017). https://doi.org/10.1007/s12228-016-9446-0

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Keywords

  • Apiaceae
  • Azorella
  • Huanaca
  • Laretia
  • Mulinum
  • Schizeilema
  • Stilbocarpa
  • phylogeny
  • taxonomy
  • megaherbs