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Revision of the Drosera Villosa Complex (Droseraceae) Supports 200 Year-Old Neglected Species Concepts

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Revision of the Drosera Villosa Complex (Droseraceae) Supports 200 Year-Old Neglected Species Concepts Phytotaxa 156 (1): 1–40 (2014) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ Article PHYTOTAXA Copyright © 2014 Magnolia Press ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.156.1.1 Exhuming Saint-Hilaire: revision of the Drosera villosa complex (Droseraceae) supports 200 year-old neglected species concepts PAULO MINATEL GONELLA1, FERNANDO RIVADAVIA2, PAULO TAKEO SANO3 & ANDREAS FLEISCHMANN4 1 Programa de Pós-Graduação em Botânica, Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, CEP 05508–900, São Paulo, Brasil; e-mail: [email protected] 2 1 Daniel Burnham Ct., San Francisco, 94109, USA; e-mail: [email protected] 3 Laboratório de Sistemática Vegetal, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, CEP 05508– 900, São Paulo, Brasil; e-mail: [email protected] 4 LMU Munich, Systematic Botany and Mycology, Menzinger Strasse 67, D-80638 Munich, Germany; e-mail: [email protected] Abstract The Drosera villosa complex is here reviewed and includes six species endemic to Brazil: D. villosa, here identified for the first time as a narrow endemic species native to the neighboring highlands of the Serra Negra and Serra do Ibitipoca, in southern Minas Gerais state; D. ascendens, rediscovered nearly 200 years after its description, narrowly endemic to the Diamantina Plateau, central Minas Gerais; D. graomogolensis, endemic to northern Minas Gerais, but here found to be more widespread than previously reported; D. latifolia, a highly polymorphic and widespread taxon, previously placed in synonymy of D. villosa and heretofore misidentified as D. ascendens, is here elevated to species rank; and two new species here described, D. riparia and D. chimaera. Furthermore, two new natural hybrids are reported: D. villosa × D. tomentosa var. glabrata and D. latifolia × D. tomentosa. The morphological characters distinguishing these taxa from each other and from similar species are discussed, together with habitat and ecological information, detailed illustrations and photographs, distribution maps, and a key to the species of the D. villosa complex is provided. Key words: Brazil, carnivorous plants, Chapada Diamantina, Espinhaço Range, new species Resumo O complexo Drosera villosa é aqui revisado e é composto por seis espécies endêmicas do Brasil: D. villosa, aqui identificada pela primeira vez como uma espécie endêmica das vizinhas Serra Negra e Serra do Ibitipoca, no sul de Minas Gerais; D. ascendens, redescoberta após quase 200 anos, micro-endêmica no Planalto de Diamantina, no centro de Minas Gerais; D. graomogolensis, endêmica do norte de Minas Gerais, porém aqui considerada mais amplamente distribuída do que reportado anteriormente; D. latifolia, um táxon altamente polimórfico e amplamente distribuído, anteriormente colocado em sinonímia de D. villosa e até então erroneamente identificado como D. ascendens, é aqui elevado ao status de espécie; e duas novas espécies que são aqui descritas, D. riparia e D. chimaera. Dois novos híbridos naturais são reportados: D. villosa × D. tomentosa var. glabrata e D. latifolia × D. tomentosa. As características morfológicas que distinguem esses táxons uns dos outros e de espécies similares são discutidos, juntamente com informações sobre habitat e ecologia, ilustrações detalhadas e fotografias, mapas de distribuição e uma chave para as espécies do complexo D. villosa é apresentada. Palavras-chave: Brasil, Cadeia do Espinhaço, Chapada Diamantina, espécies novas, plantas carnívoras. Accepted by Duilio Iamonico: 28 Nov. 2013; published: 3 Jan. 2014 1 Introduction The cosmopolitan sundews of the genus Drosera Linnaeus (1753: 281) (Droseraceae Salisb.) comprise nearly 200 species, with ca. 100 of these endemic to southwestern Australia (McPherson 2010). After the works of Saint- Hilaire (1824, 1826), the genus Drosera has been hitherto very poorly researched in Brazil, with most recent studies, such as Silva & Giulietti (1997) and the generic treatment in Flora Neotropica (Correa & Silva 2005), lacking substantial field work, relying mostly on herbarium specimens – which are often badly deformed due to the delicate nature of most Drosera taxa. As a result, these and most other previous studies failed to fully describe the diversity of the genus in Brazil, resulting in a small number of species currently registered for the country: only 14 are noted in the online Lista da Flora do Brasil (Silva 2013), the most extensive work on the Brazilian flora since Martius’ Flora Brasiliensis (produced between 1840 and 1906). Nonetheless, several recent studies suggest that Brazil hosts a much greater diversity of Drosera (Santos 1989, Rivadavia 2003, 2008, 2009, Fleischmann et al. 2007, Rivadavia et al. 2009, Rivadavia & Gonella 2011, Gonella et al. 2012). We estimate this number to be closer to 30 taxa, of which 18 belong to the so-called “Brazilian-tetraploid clade” (Rivadavia et al. 2003), characterized by a chromosome number of 2n=40, circinate leaf vernation, and flower scapes usually erect at the base. The species complex of the affinity of D. villosa Saint-Hilaire (1826: 267) belongs to the tetraploid clade, together with D. camporupestris Rivadavia (2003: 85), D. chrysolepis Taubert (1893: 505), D. graminifolia Saint- Hilaire (1826: 269), D. grantsaui Rivadavia (2003: 82), D. quartzicola Rivadavia & Gonella (2011: 34), D. schwackei (Diels 1906: 89) Rivadavia (2008: 39), D. spiralis Saint-Hilaire (1826: 270) and the four species that make up the D. montana Saint-Hilaire (1826: 260) complex: D. montana, D. tentaculata Rivadavia (2003: 79), D. tomentosa Saint-Hilaire (1826: 261), and a yet unpublished taxon. The history of the D. villosa complex is summarized below: — During a long trip through the interior of Brazil in the early 19th century, the French naturalist Auguste de Saint-Hilaire discovered and later published several Drosera species, including D. villosa and D. ascendens Saint-Hilaire (1826: 268); the former from the Serra Negra, southern Minas Gerais state, and the latter from the Serra de Curumatahy (currently Curimataí), near Diamantina, central Minas Gerais, southeastern Brazil (Saint- Hilaire 1826). — Eichler (1872) added a new taxon to the group, D. villosa var. latifolia Eichler (in Martius & Eichler 1872: 395), based on material collected by Glaziou at the Serra dos Órgãos, Rio de Janeiro state, southeastern Brazil, differing from type variety by the broader leaves and shorter petioles. — Diels (1906) considered both D. ascendens and D. villosa var. latifolia as synonyms of D. villosa in his generic revision, and this taxonomic opinion has been followed by all subsequent taxonomic treatments (e.g. Santos 1980, Silva 1994, Silva & Giulietti 1997, Duno de Stefano & Silva 2001, Correa & Silva 2005, Silva 2013). According to the concept of D. villosa proposed by Diels (1906), this species could be recognized by the oblanceolate-linear leaves and fusiform-curved seeds. — Buxbaum (1924) published D. villosa var. bifurca Buxbaum (1924: 119), differing from the type variety by the bifurcated scapes, and small differences in sepal shape. — Silva (1997) published a new species, D. graomogolensis Silva (1997: 85), based on material collected at Grão Mogol, northern Minas Gerais state, eastern Brazil. The author considered the species endemic to Grão Mogol and very similar to D. villosa, differing in the seed and style shapes, as well as in the length of the upright stem. — Rivadavia et al. (2003), Rivadavia (2005), Rivadavia (2008), and Rivadavia (2009) considered D. ascendens and D. villosa as separate species, but did not mentioned the distinctive characteristics or the respective distributions. — In 2007 a new taxon belonging to this complex was discovered in northern Minas Gerais, and a further very unusual taxon was uncovered in central Minas Gerais in 2010. These discoveries raised doubts about the true identity of D. ascendens, and eventually led to a surprising reversal in the taxonomic views previously held by the current authors. The aim of this study is to present a detailed morphological and ecological circumscription of the taxa of the D. villosa complex, including: I. Identify and circumscribe D. villosa, as described by Saint-Hilaire (1826); II. Define the taxa allied to D. villosa based on morphological evidence; III. Evaluate the distribution range of each taxon; 2 • Phytotaxa 156 (1) © 2014 Magnolia Press GONELLA ET AL. IV. Present a complete botanical description of all taxa recognized as belonging to this complex, including descriptions of eventual new species. Material & Methods Extensive field studies were undertaken by the authors between 1990 and 2013 across the known natural range of D. villosa complex populations throughout eastern Brazil. In addition, herbarium specimens at B, BHCB, CESJ, F, GFJP, HB, HRB, M, MBM, MO, NY, OUPR, P, R, RB, SP, SPF, UEC, and US (herbarium acronyms according to Thiers 2011) were personally examined and annotated by at least one of the authors. Type materials held at P, R, and SPF were personally examined under stereo dissecting microscope. Digitized high-resolution images of the type materials held at K, P, and WU were also examined. Descriptions are based on fresh, dried and spirit material. Distribution maps were prepared using georeferenced location data obtained from herbarium records (extrapolated or approximated for a few old or vaguely specified localities) or our own field observations and created
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