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(2782) Proposal to conserve Calanthe , nom. cons., against the additional names Phaius , Cyanorkis , and Gastorkis ( Orchidaceae , Collabieae )

2020; Wiley; Volume: 69; Issue: 6 Linguagem: Inglês

10.1002/tax.12396

1996-8175

Mark W. Chase, Maarten J. M. Christenhusz, André Schuiteman,

Botany, Ecology, and Taxonomy Studies

(2782) Calanthe R. Br. in Bot. Reg. 7: ad t. 573 ('578'). 1 Nov 1821 [Orchid.], nom. cons. Typus: C. veratrifolia Ker Gawl. (in Bot. Reg.: t. 720. 1 Jul 1823), nom. illeg. (Orchis triplicata Willemet, C. triplicata (Willemet) Ames). (=) Phaius Lour., Fl. Cochinch.: 517, 529. Sep 1790, nom. rej. prop. Typus: P. grandifolius Lour. (=) Alismorkis Thouars in Nouv. Bull. Sci. Soc. Philom. Paris 1: 318. Apr 1809, nom. rej. Typus (hic designatus): A. sylvalismis Thouars (Hist. Orchid.: t. 35, 36. 1822) (Centrosis sylvatica Thouars). (=) Cyanorkis Thouars in Nouv. Bull. Sci. Soc. Philom. Paris 1: 317. Apr 1809, nom. rej. prop. Typus (hic designatus): C. cyanorchis Thouars (Hist. Orchid.: t. 33, 34. 1822) (Epidendrum tetragonum Thouars). (=) Gastorkis Thouars in Nouv. Bull. Sci. Soc. Philom. Paris 1: 317. Apr 1809, nom. rej. prop. Typus: Limodorum tankervilleae Banks. Phylogenetic analyses of the orchid tribe Collabieae (Xiang & al. in PLoS One 9: e87625. 2014; Zhai & al. in Molec. Phylogen. Evol. 77: 216–222. 2014) have clearly demonstrated that species of the genera Calanthe and Phaius as currently circumscribed are intermixed in the phylogenetic tree (non-monophyletic) with the smaller genera Gastrorchis Schltr. (1921) and Cephalantheropsis Guillaumin (1960) embedded among the various clades formed by the first two. Before these last two genera were described, their species were included in Phaius and Calanthe, respectively. Calanthe is a relatively large genus (c. 220 species; Clayton & Cribb, Gen. Calanthe: 1–411. 2013; Chase & al. in Bot. J. Linn. Soc. 177: 151–174. 2015) and is important in many Asian floras and global horticulture, whereas Phaius has c. 41 species (Stone & Cribb, Lady Tankerville's Legacy: 1–278. 2017), Gastrorchis 8 and Cephalantheropsis only 5. To avoid the need to transfer the more numerous species of Calanthe to Phaius, which has priority, Yukawa & Cribb (in Bull. Natl. Mus. Nat. Sci., Ser. B, 40: 145–151. 2014) proposed to split up Calanthe, leading to the resurrection of Preptanthe Rchb. f. (1853) and Styloglossum Breda (1829) to maintain a monophyletic Calanthe s.str. This is unsatisfactory from a morphological perspective because there are no significant morphological characters to distinguish Preptanthe and Styloglossum from Calanthe s.str. Likewise, there are no reliable morphological characters distinguishing the other genera, including Phaius and Gastrorchis, from Calanthe. Traditionally, Calanthe and Phaius were distinguished by the lip being adnate to or almost free from the column, respectively, but all intermediate conditions exist. Zhai & al. (l.c.) proposed Paraphaius J.W. Zhai & F.W. Xing to accommodate a group of former Phaius species that clustered with a clade composed of Calanthe species and moved another species of Phaius into a newly proposed section of Calanthe. Zhai & al. (l.c.) included only 9 (mostly Chinese) of the c. 41 species of Phaius in their analyses and none of Gastrorchis, which was shown by Xiang & al. (l.c.) to also be embedded in this clade, perhaps as sister to Calanthe s.str. It is unclear how the remainder of the species in these genera will fit into the generic scheme proposed by Zhai & al. (l.c.), but it is likely that more genera would need to be established in order to maintain a classification that attempts to retain some form of Calanthe s.str. Phaius precedes Calanthe by 31 years, and both names are still in common use. We propose here to combine all genera in the Calanthe-Phaius alliance and to reject the name Phaius and its synonyms Cyanorkis and Gastorkis in favour of that of the largest and horticulturally most important genus, Calanthe, which will give taxonomic stability to this group. Recognition of both Phaius and Calanthe would require acceptance of a still undetermined number of additional genera, resulting in a classification with little if any morphological support. If the name Phaius is maintained because of its priority over Calanthe, a much larger number of new combinations will be needed when the two genera are combined. Several Calanthe s.l. species and a large number of artificial hybrids are ubiquitous in horticulture, whereas only three or four species of Phaius are at all common in cultivation, and few hybrids have been made. In the interest of nomenclatural stability, we propose the rejection of Phaius against the already conserved genus Calanthe (conserved against Alismorkis Thouars, nom. rej.; Wiersema & al., ICN Appendices I–VII, https://naturalhistory2.si.edu/botany/codes-proposals/). The names proposed by Thouars (l.c. 1809, and later) have caused much confusion because in addition to using the nomenclature broadly in use at that time he also proposed an alternative naming system (Friis & Rasmussen in Taxon 24: 307–318. 1975). He published the three genera Alismorkis, Cyanorkis and Gastorkis in 1809 with a short diagnosis for each, but without listing the species that he considered to belong to them. Of Alismorkis and Cyanorkis he noted that they contained a single species. In the case of Gastorkis he mentioned that Limodorum tankervilleae Banks would belong to it, among others, but the diagnosis exclusively applies to some species that were later included in Schlechter's genus Gastrorchis. According to the Code (Art. 10.2 and Art. 40.3), L. tankervilleae must be considered the type of Gastorkis, so that the latter cannot have priority over Gastrorchis, since L. tankervilleae cannot be included in Gastrorchis (it is a synonym of the type of Phaius). In a later publication, Thouars (Hist. Orchid. 1822) changed the spelling of -orkis to -orchis and provided illustrations of the four species concerned, giving names to them both in his alternative naming system and according to "traditional" nomenclature: Gastorkis ('Gastorchis') tuberogastris ('Tuberogastris'), with traditional name Limodorum tuberculosum; Gastorkis ('Gastorchis') villosagastris ('Villosagastris'), with traditional name Limodorum villosum; Cyanorkis ('Cyanorchis') cyanorchis ('Cyanorchis'), with traditional name Epidendrum tetragonum; and Alismorkis ('Alismorchis') sylvalismis ('Sylvalismis'), with traditional name Centrosis sylvatica. The last mentioned is listed in two columns of the "Premier Tableau des Espèces d'Orchidées" in the front matter of the book as "Centrosis? Sw." "Plantaginea", where the generic name can be taken as a validly published, although superfluous, replacement name for Alismorkis. Since Thouars (l.c. 1822) only associated one specific epithet with these generic names, we designate (above) Alismorkis sylvalismis Thouars (l.c. 1822: tt. 35 & 36) and Cyanorkis cyanorchis Thouars (l.c. 1822: tt. 33 & 34) as types of Alismorkis and Cyanorkis, respectively. Beginning with Sprengel (Syst. Veg. 3: 1–936. 1826), Richard (in Mém. Soc. Hist. Nat. Paris 4: 1–74. 1828) and Lindley (Gen. Sp. Orchid. Pl.: 123. 1831), authors have largely misunderstood and ignored Thouars's alternative names such as Gastorkis villosagastris and used the traditional names as basionyms, so that Limodorum villosum became the basionym of Bletia villosa (Thouars) Spreng., Phaius villosus (Thouars) Schltr. and Gastrorchis villosus (Thouars) J.V. Stone & P.J. Cribb. However, as first argued by Kuntze (in Bull. Herb. Boissier 2: 457–464. 1894) and supported by Friis & Rasmussen (l.c.), for the purpose of nomenclature Thouars's alternative names must be accepted as validly published, and they will occasionally have priority when new combinations are needed. In summary, conserving Calanthe against Phaius, Cyanorkis and Gastorkis would allow for a much smaller number of new combinations and new names. A broadly circumscribed Calanthe is the most taxonomically and nomenclaturally stable solution; it would better reflect morphology and be compatible with likely results of future molecular phylogenetic studies. MWC, https://orcid.org/0000-0002-9927-4938 MJMC, https://orcid.org/0000-0003-1398-8743 AS, https://orcid.org/0000-0001-6701-8158 We thank editors John Wiersema and John McNeill for additional comments and advice on the nomenclature of some of the names.