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Floral Convergence in Oncidiinae (Cymbidieae; Orchidaceae): an Expanded Concept of Gomesa and a New Genus Nohawilliamsia

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Floral Convergence in Oncidiinae (Cymbidieae; Orchidaceae): an Expanded Concept of Gomesa and a New Genus Nohawilliamsia Annals of Botany 104: 387–402, 2009 doi:10.1093/aob/mcp067, available online at www.aob.oxfordjournals.org Floral convergence in Oncidiinae (Cymbidieae; Orchidaceae): an expanded concept of Gomesa and a new genus Nohawilliamsia Mark W. Chase1,*, Norris H. Williams2, Aparacida Donisete de Faria3, Kurt M. Neubig2, Maria do Carmo E. Amaral4 and W. Mark Whitten2 1Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, UK, 2Department of Natural History, Florida 3 Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, USA, Instituto Nacional de Pesquisas da Downloaded from https://academic.oup.com/aob/article-abstract/104/3/387/227539 by guest on 03 June 2020 Amazoˆnia, Coordenac¸a˜o de Pesquisas em Botaˆnica, Caixa Postal 478, 69054-335, Manaus, AM, Brazil and 4Departamento do Botaˆnica, Instituto de Biologia, Caixa Postal 6109, Universidade Estadual de Campinas – UNICAMP, 13083-970 Campinas, SP, Brazil Received: 20 October 2008 Returned for revision: 26 November 2008 Accepted: 27 February 2009 Published electronically: 4 April 2009 † Background Floral morphology, particularly the angle of lip attachment to the column, has historically been the fundamental character used in establishing generic limits in subtribe Oncidiinae (Orchidaceae), but it has also been long recognized that reliance on this character alone has produced a highly artificial set of genera. In essence, lip/column relationships reflect syndromes associated with pollinator preferences; most genera of Oncidiinae as previously defined have consisted of a single floral type. Here, the degree to which this has influ- enced generic delimitation in Brazilian members of the largest genus of Oncidiinae, Oncidium, which previous molecular (DNA) studies have demonstrated to be polyphyletic, is evaluated. † Methods Phylogenetic analyses of the following multiple DNA regions were used: the plastid psbA-trnH inter- genic spacer, matK exon and two regions of ycf1 exon and nuclear ribosomal DNA, comprised of the two internal transcribed spacers, ITS1 and ITS2, and the 5.8S gene. Results from all regions analysed separately indicated highly similar relationships, so a combined matrix was analysed. † Key Results Nearly all species groups of Brazilian Oncidium are only distantly related to the type species of the genus, O. altissimum, from the Caribbean. There are two exceptions to this geographical rule: O. baueri is related to the type group and O. orthostates, an isolated species that lacks the defining tabula infrastigmata of Oncidium, is not exclusively related to any previously described genus in the subtribe. Several well-supported subclades can be observed in these results, but they do not correspond well to sections of Oncidium as previously circumscribed or to segregate genera as defined by several recent authors. In spite of their floral differences, these groups of Oncidium, formerly treated as O. sections Barbata, Concoloria pro parte, Crispa, Ranifera, Rhinocerotes, Rostrata (only O. venustum), Synsepala, Verrucituberculata pro parte and Waluewa, form a well-supported clade with Gomesa (including Rodrigueziella and Rodrigueziopsis) embedded in it. Two often recognized seg- regate genera, Baptistonia and Ornithophora, and the recently described Carriella are also embedded within the Brazilian clade. The level of variation within major subclades of the Gomesa clade is low and similar to that observed within other genera of Oncidiinae. † Conclusions Convergence on a stereotypical syndrome of floral traits associated with pollination by oil- collecting bees has resulted in these characters not being reliable for producing monophyletic taxa, and the genus Oncidium, defined by these characters, is grossly polyphyletic. Vegetative and a few floral/ inflorescence characters link these taxa with a mainly Brazilian distribution, and they were all transferred to Gomesa on this basis rather than separated from Gomesa based on their floral differences, which we hypothesize to be simple shifts in pollination strategies. Other authors have described a large number of new genera for these former members of Oncidium, but most of these are not supported by the results pre- sented here (i.e. they are not monophyletic). A new genus, Nohawilliamsia, is described for O. orthostates because it does not fit in any currently recognized genus and is only distantly related to any other member of Oncidiinae. Key words: Baptistonia, Brazilian orchids, Carriella, deceit pollination, Gomesa, ITS, matK, oil-collecting bees, Oncidium, Oncidiinae, Orchidaceae. INTRODUCTION commonly called lluvia de oro (golden rain) in Spanish or ‘dancing ladies’ in English. They are mostly epiphytic and fre- Oncidium Sw. (Oncidiinae; Cymbidieae; Orchidaceae) is one quently seen in cultivation. In its broadest circumscription, of the most conspicuous genera of neotropical orchids. Their Oncidium has over 400 species. Delimitation of Oncidium mostly bright-yellow flowers often marked with brown are and related genera has been the subject of much controversy and inconsistency (Dressler and Williams, 1975; Garay and * For correspondence. E-mail: [email protected] # The Author 2009. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: [email protected] 388 Chase et al. — Phylogenetics of Gomesa (Oncidiinae) Stacy, 1974). In simplest terms, Oncidium has always been a N.H.Williams & M.W.Chase and Zelenkoa M.W.Chase & genus of convenience, but if a set of circumscribing characters N.H.Williams, resurrection of Cyrtochilum Kunth, and expan- were to be proposed these would include: a spurless (non- sion of Caucaea Schltr., Miltonia Lindl., Otoglossum Garay & rewarding) flower with the lip (labellum) attached at a 908 or Dunsterville and Trichocentrum Poeppig & Endl. We have greater angle to the column and a complex tuberculate callus also amalgamated into Oncidium the other genera of and tabula infrastigmatica (a fleshy structure on the base of Oncidiinae that are embedded in that clade; these include the column; Fig. 1). Given the clear arbitrariness of the Cochlioda, Odontoglossum and Symphyglossum (Chase primary character, that of the angle of lip/column attachment et al., 2008). This is the fourth installment of a series of and heterogeneity of vegetative traits and chromosome papers recircumscribing genera in Oncidiinae. Here the focus numbers in Oncidium when circumscribed using this character is on the Brazilian species that have previously been included (Chase, 1986, 1987), no one was at all surprised that the early in Oncidium. restriction endonuclease studies of plastid DNA showed that In their synopsis of Oncidium, Garay and Stacy (1974) Downloaded from https://academic.oup.com/aob/article-abstract/104/3/387/227539 by guest on 03 June 2020 Oncidium was polyphyletic (Chase and Palmer, 1992). recognized several sections of O. subgenus Oncidium that Subsequent studies of plastid and nuclear DNA sequences were either endemic to Brazil (often most diverse in the (Williams et al., 2001a, b) have further detailed the extent Mata Atlantica vegetation) or had their greatest species of problems with delimitation of Oncidium, and the process diversity there. These include nearly all sections of the of generic recircumscription to achieve monophyly has been genus with a synsepal (fused lateral sepals): O. sections started, resulting in the description of Cyrtochiloides Barbata Lindl., Concoloria Kraenzl. pro parte, Crispa Rchb.f., Rhinocerotes Garay & Stacy, Synsepala Pfitzer and Waluewa (Regel) Schltr. In addition, some species of O. section Paucituberculata Lindl., which some authors have A dorsal sepal recognized as O. section Ranifera Kraenzl., also have a synse- pal. Garay and Stacy (1974) also included in O. section Paucituberculata species with thickened leaves and reduced anther cap pseudobulbs, such as O. hians Lindl., that are morphologically similar to the species of O. section Pulvinata, the Brazilian mule-ear species of Oncidium (named for their long, flat, lateral petal thickened leaves and reduced pseudobulbs; these species are now placed in Grandiphyllum Docha Neto) that have entirely stigmatic cavity free lateral sepals. Species of O. section Ranifera (e.g. tabula infrastigmatica O. raniferum Lindl., O. paranaense Kraenzl. and O. hookeri Rolfe) have larger pseudobulbs (relative to their size) and lip callus thin leaves compared with O. hians. Some species of O. section Verrucituberculata Lindl., such as O. batemanianum Parm ex Knowl. & Westc., have a synse- lip pal and are principally distributed in Brazil, whereas others lack the synsepal and are found outside of Brazil (e.g. lateral sepal O. auriferum Rchb.f. from Colombia). The key character of O. section Verrucituberculata (sensu Garay and Stacy) is the B anther cap presence of a few tubercles isolated on the lip away from the main callus, but these species are otherwise similar to species in other sections of the genus. Oncidium section Concoloria also contains some non-Brazilian species, such viscidium as O. hyphaematicum Rchb.f. and O. brachyandrum Lindl., that lack a synsepal but do have the section-defining lip without lateral lobes and an even number of parallel tubercles. stigmatic cavity Another ‘odd-ball’ Brazilian species, O. venustum Drapiez (synonym O. trulliferum Lindl.), is vegetatively similar to the members of O. section Waluewa, but it lacks the floral traits typical of this group (a pilose column and synsepal).
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