Phytotaxa 166 (2): 101–113 www.mapress.com / phytotaxa / ISSN 1179-3155 (print edition) Article Copyright © 2014 Magnolia Press PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.166.2.1 Nomenclatural changes in Bulbophyllum (Orchidaceae; Epidendroideae) JAAP J. VERMEULEN1, ANDRÉ SCHUITEMAN2 & EDUARD F. DE VOGEL3 1 Jk.art and science, Lauwerbes 8, 2318 AT Leiden, The Netherlands; E-mail: jk.artandscience@gmail.com Herbarium, Royal Botanic Gardens, Kew, TW9 3AE, UK; E-mail: a.schuiteman@kew.org 3 Naturalis Biodiversity Center, Darwinweg 2, 2333 CR Leiden, The Netherlands; E-mail: eduard.devogel@naturalis.nl 2 Abstract New combinations in Bulbophyllum (Orchidaceae) are made to correspond to the revised classification of the genus in Genera orchidacearum, vol. 6. Several new sections introduced in that publication are formally described and validated here. Key words: Bulbophyllinae, Dendrobieae, infrageneric classification, large genera Introduction Opinions about the number of genera within the former subtribe Bulbophyllinae (now subsumed in the tribe Dendrobieae) vary between extremes. Some recognized only a few small satellite genera in addition to the megagenus Bulbophyllum, e.g. Vermeulen (1996), whereas others, e.g. Clements & Jones (2002) and Szlachetko & Margońska (2001), divided the genus into numerous smaller genera. Over the years, more than 50 generic names have been proposed in addition to Bulbophyllum. In Genera orchidacearum (Pridgeon et al. 2014), supported by unpublished DNA studies by Gravendeel and coworkers, only a single genus is recognized within the former Bulbophyllinae. However, that publication presents only the outline of a revised classification of Bulbophyllum. A more detailed account of the Asian and Australian sections of Bulbophyllum is in preparation by the authors. New infrageneric nomenclature for Genera orchidacearum, as far as it is relevant to our forthcoming account, is validated below, along with the many new combinations made here. We hope that this will bring stability to the nomenclature of Bulbophyllum. Among the better-known genera of which the species are here formally transferred to Bulbophyllum are Drymoda, Pedilochilus, Sunipia and Trias. The necessary new combinations for species in the former genus Monosepalum were recently made by Shaw (2014). Taxonomy SECTION ACROCHAENE Bulbophyllum section Acrochaene (Lindl.) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Acrochaene Lindley (1853: Acrochaene 1). SECTION BLEPHARISTES Bulbophyllum section Blepharistes J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum blepharistes Reichenbach (1872: 278). Epiphytic herbs with creeping or straggling rhizomes, shoots developing from nodes below the pseudobulbs, not fused to these; rhizome scales approx. glabrous. Roots spreading, approx. glabrous to rugose. Pseudobulbs conspicuous compared to the Accepted by Mark Chase: 31 Mar 2014; published: 23 Apr. 2014 101 size of the plant, 2-leafed, minutely papillose. Leaves persistent, veins inconspicuous. Inflorescences developing from the basal node of pseudobulbs, solitary, a many-flowered, subumbellate raceme with spirally arranged flowers. Peduncle scales 8–9. Rachis somewhat thickened. Flowers per inflorescence 1–few open simultaneously; floral bracts not amplexicaul; pedicel with the basal node less than 0.5 times the diameter of the pedicel above the attachment of the floral bract, distinctly longer than the ovary. Median sepal free, 15–20 mm long, approx. triangular, acute, margins entire; glabrous, 7–9-veined; lateral sepals as the median but 1.1–1.2 times as long, adnate along their lower margins. Petals 3–4 mm long, elliptic to obovate, almost circular, retuse to rounded, distal margin erose and fimbriate, margins glabrous, adaxial surface distally finely papillose, 5-veined. Lip mobile on a thin ligament, undivided, 4.5–5.0 mm long, ovatecordate (without spreading), margins entire; glabrous; adaxially with 2 flattened, retrorse, calli close to the base. Column with the rostellum front almost level with the stelidia apices, stigma proximally not protruding, column foot tapering towards the apex, with a small, flat, median tooth near the apex. Stelidia shorter than 1/2 of the length of the column with stelidia, deltoid, acute, upper margin slightly erose, lower margin with an inconspicuous, rounded wing. Anther abaxially with a low, rounded crest; connective hardly concave, margin not drawn-out. Pollinia 4, the inner almost as long as the outer; without appendages. This section differs from other Asiatic sections with 2-leafed pseudobulbs by having persistent leaves; it differs from extralimital 2-leafed sections by having a subumbellate inflorescence. SECTION CODONOSIPHON Bulbophyllum section Codonosiphon Schlechter (1911: 178). Bulbophyllum campanuliflorum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Codonosiphon campanulatum Schlechter (1913 [1911–1914]: 894). Not Bulbophyllum campanulatum Rolfe (1909: 62). Bulbophyllum saccoglossum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Saccoglossum papuanum Schlechter (1912 [1911–1914]: 684). Not Bulbophyllum papuanum (Schltr.) J.J.Verm., Schuit. & de Vogel; see section Pedilochilus. Bulbophyllum smileiphyllum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Saccoglossum lanceolatum Williams (1943: 297). Not Bulbophyllum lanceolatum Ames & Schweinfurth (1920 : 180). Bulbophyllum takeuchii (Howcroft) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Saccoglossum takeuchii Howcroft (2001: 519). Bulbophyllum verrucosum (L.O.Williams) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Saccoglossum verrucosum Williams (1943: 297). SECTION DRYMODA Bulbophyllum section Drymoda (Lindl.) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Drymoda Lindley (1838 [1837–1841]: pl. 8c). Bulbophyllum ayuthayense J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Drymoda siamensis Schlechter (1906: 170). Not Bulbophyllum siamense Reichenbach (1867 : 572). Bulbophyllum drymoda J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Drymoda picta Lindley (1838 [1837–1841]: pl. 8c). Not Bulbophyllum pictum Reichenbach (1874: 150) Not Bulbophyllum pictum Schlechter (1913 [1911–1914]: 758). 102 • Phytotaxa 166 (2) © 2014 Magnolia Press VERMEULEN ET AL. SECTION EUBLEPHARON Bulbophyllum section Eublepharon J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum eublepharum Reichenbach (1861: 252). Epiphytic herbs with creeping, sometimes partly straggling rhizomes; shoots developing from nodes below the pseudobulbs, not fused to these; rhizome scales approx. glabrous to densely and finely papillose. Roots glabrous, shortly papillosehirsute in places. Pseudobulbs distinct, wrinkling with age along longitudinal lines, 1-leafed. Leaves persistent, veins inconspicuous to distinct. Inflorescences developing from below the pseudobulbs; solitary, a 2–many-flowered, elongated to subumbellate raceme with spirally arranged flowers. Peduncle scales 3–5. Rachis not thickened. Flowers per inflorescence open simultaneously; floral bracts not amplexicaul; pedicel with the basal node less than 1 time its diameter above the attachment of the floral bract, distinctly longer than the ovary. Median sepal free, 2.7–9.0 mm long, elliptic to ovate, rounded to acuminate, margins entire to erose or denticulate, glabrous to somewhat papillose, surface glabrous, rarely irregularly papillose abaxially; 3–5-veined; lateral sepals as the median but 1–2 times as long, free or adnate along the lower margin, 3–15 mm long, elliptic to ovate-triangular, obtuse to long-acuminate, margins entire, rarely erose, 3–5veined. Petals 1.8–5.0 mm long, elliptic-obovate to ovate, rounded to acuminate, margins slightly erose to denticulate to fimbriate, (minutely) papillose to ciliate, surface glabrous; 1–3-veined. Lip mobile on a thin ligament, undivided, 1.8–3.6 mm long, elliptic to triangular (without spreading), margins entire; glabrous to finely papillose in places; adaxially without ridges, or with 2 inconspicuous to distinct ridges. Column with the rostellum front receding in between the stelidia, stigma proximally not protruding, column foot widening or thickening towards its apex, with patent teeth and/or a transverse ridge near the apex. Stelidia 1/2 of the length of the column with stelidia or shorter; approx. deltoid to subulate, without teeth or with an inconspicuous tooth along the upper margin, or upper margin denticulate, without teeth or with a deltoid, obtuse wing along the lower margin. Anther abaxially with a small, rounded crest; connective not concave, margin drawn-out or not. Pollinia 4, the inner 0.7–0.9 times as long as the outer; appendages absent. Several species of section Eublepharon have ciliate petals, a character shared with section Hirtula. The two differ by the pseudobulbs that wrinkle with age along longitudinal lines in section Eublepharon rather than in random patterns as in section Hirtula. Additionally, section Eublepharon never has auricles at the base of the lip, flanking the ligament. Species of section Eublepharon with a congested or subumbellate raceme resemble section Recurvae and section Rhytionanthos, but differ by having the lateral sepals not curved inwards proximally. SECTION IMITATORES Bulbophyllum section Imitatores J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum imitator Vermeulen (1992: 7). Epiphytic herbs with patent to pendulous rhizomes; shoots developing from nodes below the pseudobulbs, not fused to these; rhizome scales glabrous to colliculate or transversely ribbed. Roots spreading, approx. glabrous to finely hirsute in places. Pseudobulbs distinct to minute compared to the size of the plant, 1-leafed. Leaves persistent, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs, also from nodes scattered along the rhizome, solitary or fewfasciculate, 1-flowered, or a 2–many-flowered, subumbellate or a congested raceme with spirally arranged flowers. Peduncle scales c. 3. Rachis, if present, not thickened. Flowers usually approx. all open simultaneously per inflorescence if racemose; floral bracts not amplexicaul, or (the upper) amplexicaul if inflorescence 1–2-flowered; pedicel with the basal node 0.3—1.5 times the diameter of the pedicel above the floral bract attachment, shorter to distinctly longer than the ovary. Sepals free, more or less equal, 3–7(25) mm long, elliptic to triangular, obtuse to acuminate, margins entire; glabrous, 3(5)-veined. Petals 1.2–2.7(5) mm long, elliptic to (ob-)ovate, obtuse to acuminate, margins entire to slightly erose; glabrous, 1(3)-veined. Lip mobile on a thin ligament, undivided, 1.0—1.8 (7.0) mm long, ovate (without spreading), margins entire; glabrous; adaxially usually with 2 ridges, rarely without ridges. Column with the rostellum front approx. level with the stelidia apices or somewhat receding in between these, stigma proximally not protruding, column foot tapering or slightly widening towards its apex, without teeth. Stelidia shorter to longer than 1/2 of the length of the column with stelidia; short but not vestigial, triangular, margins without teeth or with a tooth along the upper and lower margin. Anther abaxially with a low, rounded crest; connective not concave, margin not drawn-out, or with a small, triangular beak. Pollinia 2, or 4 with the inner 0.5—0.6 times as long as the outer; without appendages. This section is most similar to section Macrouris, but the proximal edge of the stigma does not protrude from the column face. NOMENCLATURAL CHANGES IN BULBOPHYLLUM (ORCHIDACEAE) Phytotaxa 166 (2) © 2014 Magnolia Press • 103 SECTION IONE Bulbophyllum section Ione (Lindl.) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Ione Lindley (1853: Ione 1). Synonym: Section Sunipia auct. J.J.Verm. in Pridgeon et al. (2014: 39). Not Sunipia Buchanan-Hamilton ex Smith in Rees (1816, Art. Stelis nos. 11 & 13). Notes: Typification of Ione with B. paleaceum (Lindley 1853: Ione 2) Hooker (1890 [1886-1890]: 769) in Pridgeon et al. (2014: 5) is superfluous because Senghas (1991: 1467) designated Ione bicolor Lindl. as type species. Bulbophyllum angustipetalum (Seidenf.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Sunipia angustipetala Seidenfaden (1980: 7). Bulbophyllum arunachalense (A.N.Rao) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione arunachalense Rao (1997:48). Bulbophyllum australe (Seidenf.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione australis Seidenfaden (1969: 234). Bulbophyllum bifurcatoflorens (Fukuyama) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione bifurcatoflorens Fukuyama (1935: 440). Bulbophyllum cumberlegei (Seidenf.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione cumberlegei Seidenfaden (1969: 217). Bulbophyllum interpositum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione intermedia King & Pantling (1896: 120). Not Bulbophyllum intermedium Bailey (1896: 12). Not Bulbophyllum intermedium De Wildeman (1916 [1916–1919]: 176). Bulbophyllum jainii (Hynn. & Malhotra) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Sunipia jainii Hynniewta & Malhotra (1978: 31). Bulbophyllum kachinense (Seidenf.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Sunipia kachinensis Seidenfaden (1981: 211). Bulbophyllum kipgenii (Kishor, Chowlu & Vij) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione kipgenii Kishor, Chowlu & Vij (2012: 517). Bulbophyllum lomsakense J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione thailandica Seidenfaden & Smitinand (1965 [1958–1965]: 813). Not Bulbophyllum thailandicum Seidenfaden & Smitinand (1965 [1958–1965]: 797). Bulbophyllum lopalanthum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione grandiflora Rolfe (1908: 413). Not Bulbophyllum grandiflorum Blume (1850: 42). Bulbophyllum medioximum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione annamensis Ridley (1921: 115). Not Bulbophyllum annamense (Garay 1999: 306) Sieder & Kiehn in Sieder et al. (2009: 406). Bulbophyllum minutior J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione minor Seidenfaden (1969: 218). Not Bulbophyllum minus De Wildeman (1921: 94). 104 • Phytotaxa 166 (2) © 2014 Magnolia Press VERMEULEN ET AL. Bulbophyllum raskotii J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Sunipia nepalensis Raskoti & Ale (2011: 55). Not Bulbophyllum nepalense Raskoti & Ale (2013: 381). Bulbophyllum nigricans (Aver.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Sunipia nigricans Averyanov (2007: 302). Bulbophyllum pallidulum (Aver.) J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione pallida Averyanov (1997: 143). Not Bulbophyllum pallidum Seidenfaden (1979: 206). Bulbophyllum purpureofuscum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione fuscopurpurea Lindley (1853: Ione 2). Not Bulbophyllum fuscopurpureum Wight 1852: t. 1651). Bulbophyllum sunipia J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Sunipia scariosa Lindley (1833 [1830–1840]: 179). Not Bulbophyllum scariosum Summerhayes (1953: 145). Bulbophyllum rimannii (Rchb.f.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Acrochaene rimannii Reichenbach (1882: 796). Bulbophyllum roseopictum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione bicolor Lindley (1853: Ione 3). Not Bulbophyllum bicolor Lindley (1830 [1830–1840]: 49). Bulbophyllum rutiliflorum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione rutilans Seidenfaden (1995: 54). Not Bulbophyllum rutilans Vermeulen & Lamb (2008: 68). Bulbophyllum sasakii (Hayata) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione sasakii Hayata (1912: 139). Bulbophyllum soidaoense (Seidenf.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Ione soidaoensis Seidenfaden (1969: 220). Bulbophyllum tsii J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Sunipia hainanensis Z.H.Tsi in Tsi, Chen & Ding (1995: 590) Not Bulbophyllum hainanense Z.H.Tsi (1982: 118). Bulbophyllum xanthochloron J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Ione viridis Seidenfaden (1969: 222). Not Bulbophyllum viride Rolfe (1893: 170). SECTION LEOPARDINAE Bulbophyllum section Leopardinae Bentham & Hooker (1883 [1880–1883]: 502). Bulbophyyllum yunnanense Rolfe (1903: 14). Heterotypic synonym: Trias verrucosa Liu et al. (2007: 4461). NOMENCLATURAL CHANGES IN BULBOPHYLLUM (ORCHIDACEAE) Phytotaxa 166 (2) © 2014 Magnolia Press • 105 SECTION LEPANTHANTHE Bulbophyllum section Lepanthanthe Schlechter (1913 [1911–1914]: 704, 875. Bulbophyllum paulianum J.J.Verm., Schuit. & de Vogel, spec. nov. Type:PAPUA NEW GUINEA. Eastern Highland Province, Goroka, Gahavisuka National Park, cult. Jongejan 1342 (L!). Bulbophyllum nephropetalum auct. Vermeulen (1993: 81). Notes: The interpretation and neotypification of B. nephropetalum Schltr., in Vermeulen (1993) has to be rejected. Various parts of Schlechter’s original description show that B. nephropetalum is synonymous with B. erinaceum Schltr., such as the leaf width, the shape of the appendage of the median sepal and the shape of the lip. Here, B. nephropetalum auct. Vermeulen (1993 : 81) is renamed in honour of Paul Ormerod, who drew the first author’s attention to this and other mistakes in his earlier work. SECTION MACROCAULIA Bulbophyllum section Macrocaulia (Blume 1825: 318) Averyanov (1994: 279). Bulbophyllum jejosephii J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Trias pusilla Joseph & Deka (1973: 378). Not Bulbophyllum pusillum Petit Thouars (1822: t. 102). Not Bulbophyllum pusillum De Wildeman (1921: 95). SECTION MONOMERIA Bulbophyllum section Monomeria (Lindl.) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Monomeria Lindley (1830 [1830–1840]: 61). Bulbophyllum crabro (E.C.Parish & Rchb.f.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Monomeria crabro Reichenbach (1874: 143). Heterotypic synonym: Bulbophyllum monomeria J.J.Verm. in Pridgeon et al. (2014: 33), nom. inval. Bulbophyllum fengianum (Ormerod) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Monomeria fengiana Ormerod (2011: 44). SECTION PEDILOCHILUS Bulbophyllum section Pedilochilus (Schltr.) J.J.Verm. & O'Byrne (2011: 199). Bulbophyllum alpinum (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus alpinum Van Royen (1979: 251). Bulbophyllum araiophyllum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus angustifolius Schlechter (1919b: 120) Not Bulbophyllum angustifolium (Blume 1825: 310) Lindley (1830 [1830–1840] : 57). Not Pedilochilus angustifolium Schlechter (1912 [1911–1914]: 687). Bulbophyllum balgooiji J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus subalpinum Van Royen (1979: 267). Not Bulbophyllum subalpinum Van Royen (1979: 180). 106 • Phytotaxa 166 (2) © 2014 Magnolia Press VERMEULEN ET AL. Bulbophyllum brachiatum (Schltr.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus brachiatus Schlechter (1919: 47). Bulbophyllum brachypus (Schltr.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus brachypus Schlechter (1912 [1911–1914]: 692). Bulbophyllum clemensiorum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus clemensiae Williams (1946: 162). Not Bulbophyllum clemensiae Ames (1912: 129). Bulbophyllum cyatheicola (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus cyatheicola Van Royen (1979: 256). Bulbophyllum dolichopus J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus longipes Schlechter (1912 [1911–1914]: 691). Not Bulbophyllum longipes Reichenbach (1861: 253). Bulbophyllum erythrokyle J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus stictanthum Schlechter (1912 [1911–1914]: 693). Not Bulbophyllum stictanthum Schlechter (1913 [1911–1914]: 726). Bulbophyllum hermonii (Cribb & B.Lewis) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus hermonii Cribb & Lewis (1989: 135). Bulbophyllum kermesinostriatum (J.J.Sm.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus kermesinostriatum Smith (1915: 16). Bulbophyllum lagaropetalum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus parvulum Schlechter (1912 [1911–1914): 686). Not Bulbophyllum parvulum Lindley (1830 [1830–1840]: 49). Bulbophyllum lagarophyllum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus angustifolium Schlechter (1912 [1911–1914]: 687). Not Bulbophyllum angustifolium (Blume 1825: 310.) Lindley (1830 [1830–1840]: 57). Not Pedilochilus angustifolius Schlechter (1919b: 120). Bulbophyllum leptotriche J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus ciliolatum Schlechter (1912 [1911–1914]: 688). Not Bulbophyllum ciliolatum Schlechter (1913 [1911–1914]: 809). Bulbophyllum macrorrhinum (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus macrorrhinum Van Royen (1979: 265). Bulbophyllum montisdischorense J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus dischorense Schlechter (1912 [1911–1914]: 689). Not Bulbophyllum dischorense Schlechter (1913 [1911–1914]: 775). Bulbophyllum obovatum (J.J.Sm.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus obovatum Smith (1929: 441). Bulbophyllum oreomene J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus montana Ridley (1916: 190). Not Bulbophyllum montanum Schlechter (1913 [1911–1914]: 801). Bulbophyllum papuanum (Schltr.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus papuanum Schlechter (1905: 219). NOMENCLATURAL CHANGES IN BULBOPHYLLUM (ORCHIDACEAE) Phytotaxa 166 (2) © 2014 Magnolia Press • 107 Bulbophyllum pedilochilus J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus guttulatum Schlechter (1912 [1911–1914]: 689). Not Bulbophyllum guttulatum (Hooker 1890 [1886–1890]: 776.) Balakrishnan (1970: 66). Bulbophyllum petioliferum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus petiolatum Schlechter (1912 [1911–1914): 690). Not Bulbophyllum petiolatum Smith (1910: 58). Bulbophyllum petrophilum (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus petrophilum Van Royen (1979: 248). Bulbophyllum piundaundense (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus piundaundense Van Royen (1979: 270). Bulbophyllum psychrophilum (F.Muell.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Dendrobium psychrophilum Von Mueller (1889: 34). Bulbophyllum spaniostagon J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus pusillum Schlechter (1912 [1911–1914): 686). Not Bulbophyllum pusillum Petit Thouars (1822: t. 102). Not Bulbophyllum pusillum De Wildeman (1921: 95). Bulbophyllum sarawaketense (P.Royen) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus sarawaketensis Van Royen (1979: 275). Bulbophyllum teinophyllum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus grandifolium Van Royen (1979: 273). Not Bulbophyllum grandifolium Schlechter (1913 [1911–1914]: 743). Bulbophyllum tapeinophyton J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus humile Smith (1929: 440). Not Bulbophyllum humile Schlechter (1913 [1911–1914]: 730). Bulbophyllum terrestre (J.J.Sm.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Pedilochilus terrestre Smith (1929: 442). Bulbophyllum theiochroma J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus sulphureum Smith (1914: 72). Not Bulbophyllum sulfureum Schlechter (1925 : 225) Bulbophyllum vanroyenii J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus perpusillum Van Royen (1979: 241). Not Bulbophyllum perpusillum Ridley (1908a: 130). Bulbophyllum zophyranthum J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Pedilochilus flavum Schlechter (1912 [1911–1914]: 691). Not Bulbophyllum flavum Schlechter (1913 [1911–1914]: 853). SECTION PLANIBULBUS Bulbophyllum section Planibulbus J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum planibulbe (Ridley 1893: 364) Ridley (1908 :79). Epiphytic herbs with long-creeping or straggling rhizomes; shoots usually developing from near the apex of the pseudobulbs; rhizome scales approx. glabrous. Roots spreading, glabrous, shortly papillose-hirsute in places. Pseudobulbs distinct, 1leafed. Leaves persistent, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs or from nodes 108 • Phytotaxa 166 (2) © 2014 Magnolia Press VERMEULEN ET AL. scattered along the rhizome; solitary, a 2–many-flowered, congested or subumbellate raceme with spirally arranged flowers. Peduncle scales 3–4. Rachis not thickened. Flowers per inflorescence open simultaneously; floral bracts amplexicaul or not, not tubular; pedicel with the basal node level with the attachment of the floral bract, distinctly longer than the ovary. Sepals free, the median 4.5–13.0 mm long, ovate, acuminate, margins entire; glabrous, 3-veined; lateral sepals as the median but 1.5–2.2 times as long, 8.5–25.0 mm long, triangular. Petals 2.0–5.5 mm long, elliptic to ovate, obtuse to acuminate, margins entire or distally erose; glabrous, 1–3-veined. Lip mobile on a thin ligament, undivided, 1.4–3.8 mm long, approx. ovate to triangular (without spreading), margins entire, papillose; adaxially without ridges or with 3 ridges converging into a distal callus, surface papillose; abaxial surface glabrous to papillose. Column with the rostellum front receding in between the stelidia, stigma proximally not protruding, column foot not widening towards its apex. Stelidia shorter than 1/2 of the length of the column including stelidia; triangular to subulate, with a slight, rounded wing along the lower margin. Anther abaxially virtually without crest; connective not concave, margin not drawn-out. Pollinia 4, the inner 0.7–0.8 times as long as the outer; appendages absent. The young shoots pierce the pseudobulbs and arise from their apex, similar to many species of section Macrocaulia. Plants in section Planibulbus are larger, however, with long-creeping rhizomes that are wiry and not brittle and have a racemose inflorescence. SECTION PLUMATA Bulbophyllum section Plumata J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum plumatum Ames (1915: 184). Epiphytic herbs with creeping rhizomes, shoots developing from nodes below pseudobulbs, not fused to these; rhizome scales densely and finely colliculate to papillose. Roots glabrous, shortly papillose-hirsute locally. Pseudobulbs distinct, 1-leafed. Leaves persistent, veins inconspicuous. Inflorescences developing from nodes below pseudobulbs; solitary, 1-flowered or a 2–4-flowered, subumbellate raceme with spirally arranged flowers. Peduncle scales 3–5. Rachis slightly thickened or not. Flowers per inflorescence open simultaneously if inflorescence racemose; floral bracts not amplexicaul; pedicel with the basal node 0.3–1.0 times its diameter above the attachment of the floral bract, distinctly longer than the ovary. Median sepal free, 6–19 mm long, elliptic to ovate, acute to acuminate, margins entire, glabrous to minutely papillose-ciliolate, surface glabrous; 3–17-veined; lateral sepals as the median but 2.5–16.0 times as long, adnate along (the proximal part of) the upper and lower margin, or along the upper margin only, 24–94 mm long, elliptic to triangular to linear, acute to caudate, margins entire, glabrous, surface glabrous, or abaxially finely papillose; 5–11-veined. Petals 1–2 mm long excluding the appendages, elliptic to oblong to transversely ovate, rounded, apical margin with 12–32 appendages, glabrous, surface glabrous; 1–3-veined; appendages 0.8–5.5 mm long, narrowly attached or stalked, elliptic/ellipsoid to obovate/obovoid, obtuse to acute, thin and flat to approx. orbicular in section, glabrous to vesiculate. Lip mobile on a thin ligament, undivided, 1.8–3.5 mm long, triangular (somewhat narrowed towards the apex) (without spreading), margins entire; glabrous to finely papillose; adaxially without ridges, or with 2 weak ridges. Column with the rostellum front receding in between the stelidia, stigma proximally not protruding, column face furrowed longitudinally, column foot not widened towards its apex. Stelidia shorter than 1/2 of the length of the column with stelidia; approx. triangular to subulate, with a deltoid, obtuse tooth along the upper margin, or upper margin denticulate, with a deltoid to triangular, rounded to obtuse wing along the lower margin. Anther abaxially with a rounded crest; connective deeply concave, margin not drawnout. Pollinia 4, the inner 0.7–1.0 times as long as the outer; appendages absent. Among the sections with the upper margin of the lateral sepals approaching to adnate, section Plumata is distinguished by the longitudinally furrowed column face. SECTION REPTANTIA Bulbophyllum section Reptantia J.J.Verm. in Pearce et al. (2001: 121). Synonym: Sunipia Buchanan-Hamilton ex Smith in Rees (1816, Art. Stelis nos. 11 & 13). Notes: Senghas (1991: 1467) indicated Sunipia racemosa (Buch.-Ham. ex Sm.) Tang & Wang (1951: 90) as lectotype of Sunipia. Garay (1994: 612) observed that the only extant type material of Sunipia racemosa is a plate in LINN (collection Buchanan-Hamilton, t. 20, fig. 2, marked "Sunipia racemosa", and "Stelis"). This plate represents B. reptans Lindley (1830 [1830–1840]: 51)), type species of section Reptantia. NOMENCLATURAL CHANGES IN BULBOPHYLLUM (ORCHIDACEAE) Phytotaxa 166 (2) © 2014 Magnolia Press • 109 SECTION RHINANTHERA Bulbophyllum section Rhinanthera J.J.Verm., Schuit. & de Vogel, sect. nov. Type species: Bulbophyllum wrayi J.D.Hooker (1890 [1886 –1890]: 766). Epiphytic herbs with creeping rhizomes; shoots developing from nodes below pseudobulbs, not fused to these; rhizome scales minutely papillose. Roots spreading, glabrous to minutely hirsute locally. Pseudobulbs distinct, 1-leafed. Leaves persistent, veins inconspicuous. Inflorescences developing from nodes below pseudobulb, solitary, an elongate raceme with spirally arranged flowers. Peduncle scales 4–10. Rachis not thickened. Flowers per inflorescence more or less open simultaneously; floral bracts not amplexicaul; pedicel with the basal node 0.5–1.5 times its diameter above the floral bract attachment, distinctly longer than the ovary. Sepals free, the median 6–10 mm long, obovate to ovate-triangular, rounded to acuminate, margins entire; glabrous, 3-veined; lateral sepals approx. equally long or shorter, 4.5–9.0 mm long, obliquely ovate to triangular. Petals 2.5–5.5 mm long, (sub-)spathulate to elliptic to oblong to triangular, rounded to subacute, margins entire; glabrous, 1-veined. Lip mobile on a thin ligament, undivided or sub-3-lobed, 1.8–3.0 mm long; oblong to ovate, or divided in a wide, ovate basal part and a drawn-out, narrow, oblong to ovate apical part (without spreading), margins entire; glabrous; adaxially with 2 knob-like ridges close to the base or with 2 distinct ridges more distally positioned. Column with the rostellum front almost level with the stelidia apices or receding in between, stigma proximally with a protruding tooth, column foot not widened towards the apex, but with a median tooth at the apex. Stelidia approx. 1/2 of the length of the column with stelidia rarely slightly longer, triangular to subulate, with or without a rounded to triangular wing along the upper margin, with or without a deltoid to triangular, approx. obtuse wing along the lower. Anther abaxially with a rounded crest; connective concave, margin distinctly drawn-out into an elliptic to triangular beak. Pollinia 4, the inner 0.8—1.0 times as long as the outer, appendages absent. This section is most similar to racemose species of B. section Beccariana; it generally differs by having 3-veined sepals and 1-veined petals. A few species of section Beccariana have the same set of characters, but these differ in having cylindrical pseudobulbs. It is also similar to sections Altisceptrum and Phreatiopsis; the drawn-out connective front of the anther is distinctive. SECTION RHYTIONANTHOS Bulbophyllum section Rhytionanthos (Garay, Hamer & Siegerist) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Rhytionanthos Garay, Hamer & Siegerist (1994: 637). SECTION TRIAS Bulbophyllum section Trias (Lindl.) J.J.Verm., Schuit. & de Vogel, stat. nov. Basionym: Trias Lindley (1830 [1830–1840]: 60). Bulbophyllum [unranked, "§"] Trias (Lindl.) Reichenbach (1861: 249) ("Bolbophyllum"). Bulbophyllum antheae (J.J.Verm. & Lamb) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Trias antheae Vermeulen (1994: 347). Bulbophyllum bonaccordense (C.S.Kumar) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Trias bonaccordensis Kumar (1989: 105). Bulbophyllum cambodianum (Christenson) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Trias cambodiana Christenson (2003: 542). Heterotypic synonym: Trias nummularia Averyanov & Averyanova (2006: 32). Not Bulbophyllum nummularia Rolfe (1897: 35). Bulbophyllum capnophyton J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Trias nana Seidenfaden (1976: 24). Not Bulbophyllum nanum De Wildeman (1903 [1903–1905]: 122). 110 • Phytotaxa 166 (2) © 2014 Magnolia Press VERMEULEN ET AL. Bulbophyllum leion J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Trias mollis Seidenfaden (1976: 23). Not Bulbophyllum molle Vermeulen & O’Byrne (2008: 84). Bulbophyllum meson J.J.Verm., Schuit. & de Vogel, nom. nov. Replaced name: Trias intermedia Seidenfaden & Smitinand (1965 [1958–1965]: 809). Not Bulbophyllum intermedium Bailey (1896: 12). Not Bulbophyllum intermedium De Wildeman (1916 [1916–1919]: 176). Bulbophyllum stocksii (Hook.f.) J.J.Verm., Schuit. & de Vogel, comb. nov. Basionym: Trias stocksii Hooker (1890 [1886–1890]: 781). References Ames, O. 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