A recircumscription of Goodyera (Orchidaceae), including the description of Paorchis gen. nov., and resurrection of Cion
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Abstract. Goodyerinae is a cosmopolitan terrestrial subtribe of Orchidaceae; some currently accepted Goodyerinae genera have been suggested to be non-monophyletic, including the type genus for the subtribe, Goodyera. The polyphyly of Goodyera has been supported by multiple molecular studies, with several primarily Indomalayan and Australasian species forming distinct clades more closely related to genera such as Erythrodes, Lepidogyne, and Platylepis. The potential morphological limits of these clades, however, have not been fully explored. To better reflect the evolutionary relationships of these species, I present a broadscale taxonomic recircumscription of Goodyera s.l. based on detailed morphological investigation of herbarium and living specimens. Goodyera s.s. is restricted to species bearing commonly marked, often basal leaves, and flowers with a short-footed, stout column. A new genus, Paorchis, is described, four names are transferred to Platylepis, and Cionisaccus, Eucosia, and Salacistis are resurrected, with the following new combinations: Cionisaccus procera, Paorchis thailandica, Eucosia afzelii, E. brachiorrhynchos, E. seikomontana, E. viridiflora, Platylepis flaccida, P. goudotii, P. roseum, P. scirpta, Salacistis angustifolia, S. beccarii, S. clavata, S. elmeri, S. erythrodes, S. fumata, S. lanceolata, S. rhombodoides, S. rubicunda, and S. werneri. Keywords: Convergent evolution, Madagascar, Malay Archipelago, orchid, Platylepis, Southeast Asia, taxonomy.
Goodyerinae (Orchidaceae: Orchidoideae: Cranichideae) is a terrestrial subtribe characterized by a rhizomatous habit, often petiolate, variously marked and/or darkly-pigmented and iridescent leaves, distinct stipes, and sectile pollinia. Most genera are vegetatively similar, being primarily differentiated by characteristics of the labellum and column. Although Goodyerinae is the second most species-rich Orchidoideae subtribe (after Orchidinae), with at least 740 species in 35 genera (Pridgeon et al. 2003; Chase et al., 2015), and has received recent systematic attention (e.g., Bhattacharjee & Chowdhery, 2018; Ormerod, 2005, 2007), it is the least well-known terrestrial subtribe of Orchidaceae. Indeed, Pridgeon et al. (2003) and Chase et al. (2015) discuss the likelihood that several genera may not represent natural groups, being either polyphyletic or paraphyletic. This systematic uncertainty is borne out by the
complicated taxonomic histories of many Goodyerinae taxa: most accepted genera have several synonyms, many species have been placed in three or more genera, and many species have been described multiple times. For example, Zeuxine Lindl. has eight synonyms, and Macodes petola (Blume) Lindl. has been described 11 times in five genera. Only recently have Hu et al. (2016) and Chen et al. (2019) provided much needed molecular phylogenetic context to this recalcitrant subtribe, supporting the polyphyly of several genera. Within Goodyerinae, the largest genera are Microchilus C. Presl (ca. 145 spp.), Goodyera R. Br. in W.T. Aiton (s. lato, 98
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