Andinia subgen. Brachycladium (Luer)Karremans&S.Vieira-Uribe, 2017

Wilson, Mark, Frank, Graham S., Jost, Lou, Pridgeon, Alec M., Vieira-Uribe, Sebastian & Karremans, Adam P., 2017, Phylogenetic analysis of Andinia (Pleurothallidinae; Orchidaceae) and a systematic re-circumscription of the genus, Phytotaxa 295 (2), pp. 101-131 : 123-126

Andinia subgen. Brachycladium (Luer)Karremans&S.Vieira-Uribe , comb.nov. Bas. Lepanthes subgen. Brachycladium

Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 15: 31. 1986. Type: Lepanthes nummularia Reichenbach f., Xenia

Orchid. 1: 142. 1856.

Syn. Lepanthes sect. Brachycladae Reichenbach f., Xenia Orchid. 1: 142, 1856. Type: Lepanthes nummularia Reichenbach f., Xenia Orchid. 1: 142. 1856. Syn. Lepanthes sect. Caulescentes Garay, Canad. J. Bot. 34: 252, 1956. Nom. nud. Syn. Brachycladium (Luer) Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 103: 307. 2005. Nom. illeg. hom. syn. Oreophilus W.E.Higgins & Archila, Selbyana 29(2): 202. 2009. Nom. illeg. superfl. syn. Neooreophilus Archila, Revista Guatemal. 12(2): 73. 2009. Syn. Penducella Luer & Thoerle, Orchid Digest 74(2): 68. 2010.

Andinia subgen. Brachycladium sect. Brachycladae (Rchb.f.) Karremans & S.Vieira-Uribe , comb. nov. Bas. Lepanthes subgen. Brachycladium sect. Brachycladae Rchb. f., Syst. Bot. Missouri Bot. Gard. 52: 3. 1994. Type: Lepanthes nummularia Reichenbach f., Xenia Orchid. 1: 142. 1856.

Syn. Lepanthes subgen Brachycladium sect. Bilamellatae Luer, Syst. Bot. Missouri Bot. Gard. 52: 3. 1994. Type: Lepanthes stalactites Luer & Hirtz, Lindleyana 2(2): 105. 1987.

Plants repent, pendent. Leaves glabrous with margins entire. Ovary glabrous. Lip at least obscurely trilobate, with the midlobe transformed into an appendix-like structure, prominently hirsute, much shorter than the column. Column foot absent. Anther apical, stigma ventral, rostellum inconspicuous. Pollinaria with a drop-like viscidium. Two species distributed through the Andean regions of Colombia, Ecuador and Peru ( Fig. 7), an elevational range of 1,690 to 3,000 m ( Fig. 8 ). An example, Andinia nummularia (Rchb.f.) Karremans & S.Vieira-Uribe , is illustrated ( Fig. 11e ).

Andinia nummularia (Rchb.f.) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes nummularia Rchb.f., Xenia Orchid. 1: 142. 1856.

Andinia stalactites (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes stalactites Luer & Hirtz, Lindleyana 2(2): 105. 1987.

Andinia subgen. Brachycladium sect. Amplectentes (Luer) Karremans & S.Vieira-Uribe , comb. nov. Bas. Lepanthes subgen. Brachycladium sect. Amplectentes Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 3. 1994. Type: Lepanthes pilosella Reichenbach f. , Flora 69: 556, 1886.

Plants repent and pendent. Leaves mostly with margins denticulate-fimbriate, sometimes hirsute. The ovary glabrous to hirsute or spiculate. The lip variously lobed but always lacks the midlobe transformed into appendix and always surrounding or embracing the column. Anther and stigma apical, rostellum elongate, antrorse, pollinaria with a drop-like viscidium. Fifty one species distributed through the Andean regions of Colombia, Ecuador, Peru and Bolivia ( Fig. 7), with an elevational range of 1,200 to 2,660 m ( Fig. 8 ). An example, Andinia montis-rotundi ( Ortiz 1997: 318) Karremans & S.Vieira-Uribe , is illustrated ( Fig. 11f ).

Andinia ariasiana (Luer & L.Jost) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes ariasiana Luer & L.Jost, Monogr. Syst. Bot. Missouri Bot. Gard. 72: 104. 1998. Andinia auriculata (Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus auriculatus Archila, Revista Guatemal. 17(1): 44. 2014.

Andinia bifida (Tobar & Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus bifidus Tobar & Archila, Revista Guatemal. 15(2): 2. 2012.

Andinia cardiocheila (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes cardiocheila Luer & R.Escobar, Orquideología 19(2): 86. 1994.

Andinia catella (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes catella Luer & R.Escobar, Orquideología 16(1): 10. 1983.

Andinia caveroi (D.E.Benn. & Christenson) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes caveroi D.E.Bennett & Christenson, Icon. Orchid. Peruv. : t. 670. 2001.

Andinia chaoae (S.Vieira-Uribe & L.Jost) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus chaoae S.Vieira-Uribe & L.Jost , Lankesteriana 15(3): 213. 2015.

Andinia chelosepala (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Oreophilus chelosepalus Luer & Hirtz, Selbyana 30(1): 16. 2009.

Andinia chilopsis (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes chilopsis Luer & Hirtz, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 88. 2002. Andinia ciliaris (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes ciliaris Luer & Hirtz, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 8. 1994.

Andinia composita (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes composita Luer & R.Escobar, Orquideología 19(2): 88. 1994.

Andinia cordilabia (Luer) S.Vieira-Uribe & Karremans , comb. nov.

Bas. Lepanthes cordilabia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 88. 2002.

Andinia dactyla (Garay) Karremans & S.Vieira-Uribe, Orquideología 33(2): 114. 2016.

Bas. Lepanthes dactyla Garay, Orquideología 6(1): 13. 1971.

Andinia dentata (Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus dentatus Archila, Revista Guatemal. , 17(1): 41. 2014.

Andinia destituta (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes destituta Luer & R.Escobar, Orquideología 17(3): 179. 1988.

Andinia erepsis (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes erepsis Luer & Hirtz, Die Orchidee 37(5): 215. 1986.

Andinia exigua (Luer & L.Jost) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes exigua Luer & L.Jost, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 89. 2002.

Andinia geminipetala (Luer & J.Portilla) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes geminipetala Luer & J.Portilla, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 89. 2002. Andinia hippocrepica (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes hippocrepica Luer & R.Escobar, Orquideología 18(1): 54. 1991.

Andinia irrasa (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes irrasa Luer & R.Escobar, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 18. 1994. Andinia lueri S.Vieira-Uribe & Karremans, Orquideología 33(2): 116. 2016.

Andinia lunaris (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes lunaris Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 21. 1994.

Andinia lunatocheila (Tobar & Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus lunatocheillus Tobar & Archila, Revista Guatemal. 15(2): 26. 2012.

Andinia lupula (Luer & Hirtz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes lupula Luer & Hirtz, Amer. Orchid Soc. Bull. 53(11): 1162. 1984.

Andinia lynniana (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes lynniana Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 91. 2002.

Andinia macrotica (Luer & Dalström) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes macrotica Luer & Dalström, Monogr. Syst. Bot. Missouri Bot. Gard. 61(4): 3. 1996. Andinia micropetala (L.O.Williams) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes micropetala L.O.Williams, Bot. Mus. Leafl. 9: 4. 1940.

Andinia mongei (Tobar & Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus mongeei Tobar & Archila, Revista Guatemal. 15(2): 23. 2012.

Andinia monilia (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes monilia Luer & R.Escobar, Orquideología 16(1): 12. 1983.

Andinia montis-rotundi (P.Ortiz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes montis-rotundi P.Ortiz, Orquideología 20(3): 318. 1997.

Andinia octocornuta (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes octocornuta Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 91. 2002.

Andinia ortiziana (S.V.Uribe & Thoerle) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus ortizianus S.V.Uribe & Thoerle, Orquideología 28(2): 135. 2011.

Andinia pendens (Garay) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes pendens Garay, Can. J. Bot. 34: 252. 1956.

Andinia persimilis (Luer & Sijm) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes persimilis Luer & Sijm, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 92. 2002. Andinia phallica (Tobar & Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus phalicus Tobar & Archila, Revista Guatemal. 15(2): 20. 2012.

Andinia pholeter (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes pholeter Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 33. 1994.

Andinia pilosella (Rchb.f.) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes pilosella Reichenbach f. , Flora 69: 556. 1886.

Andinia platysepala (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes platysepala Luer & R.Escobar, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 36. 1994. Andinia pseudocaulescens (L.B.Sm. & S.K.Harris) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes pseudocaulescens L.B.Sm. & S.K.Harris, Bot. Mus. Leafl. 2(3): 33. 1934.

Andinia ricii (Luer & R.Vásquez) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes ricii Luer & R.Vásquez, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 40. 1994. Andinia rosea (Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus roseus Archila, Revista Guatemal. 17(1): 43. 2014.

Andinia rotunda (Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus rotundus Archila, Revista Guatemal. 17(1): 42. 2014.

Andinia sibundoyensis (Kolan.) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus sibundoyensis Kolan., Ann. Bot. Fennici 50: 169. 2014

Andinia sudamericana (Archila) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus sudamericanus Archila, Revista Guatemal. , 17(1): 40. 2014.

Andinia triangularis (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes triangularis Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 43. 1994.

Andinia tridactyla (Luer) S.Vieira-Uribe & Karremans , comb. nov.

Bas. Lepanthes tridactyla Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 92. 2002.

Andinia ursula (Luer & R.Escobar) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes ursula Luer & R.Escobar, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 46. 1994. Andinia viebrockiana (Luer & L.Jost) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes viebrockiana Luer & L.Jost, Monogr. Syst. Bot. Missouri Bot. Gard. 72: 108. 1998. Andinia vieira-pereziana (P.Ortiz) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Neooreophilus vieira-perezianus P.Ortiz, Orquideología 28(1): 7. 2011.

Andinia villosa (Løjtnant) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes villosa Løjtnant, Bot. Not. 130(4): 419. 1977.

Andinia werneri (Luer) Karremans & S.Vieira-Uribe , comb. nov.

Bas. Lepanthes werneri Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 88: 93. 2002.

Andinia subgen. Masdevalliantha (Luer) Karremans & Mark Wilson, comb. nov. Bas. Pleurothallis subgen. Masdevalliantha Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 20: 44. 1986. Type. Pleurothallis masdevalliopsis Luer, Phytologia 44(3): 170. 1979.

Syn. Pleurothallis subgen. Xenion Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 20: 96. 1986.

Syn. Masdevalliantha (Luer) Szlachetko & Margonska., Polish Bot. J. 46(2): 117. 2001.

Syn. Xenosia Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 95: 265. 2004.

Plant repent, ascending or caespitose. Inflorescence single-flowered. Lip three-lobed, with a midlobe prominent and obtuse, not transformed into appendix. Lip winged, not involving the relatively much shorter column. Column foot present. The subgenus includes the species previously assigned to the genera Masdevalliantha and Xenosia . Four species distributed through the Andean regions of Colombia, Ecuador and Peru ( Fig. 6 ), with an elevational range of 2,400 to 3,825 m ( Fig. 8 ). The three most similar species are illustrated to aid identification ( Fig. 9 ) and a photograph of an example, Andinia longiserpens (C.Schweinf.) Karremans & Mark Wilson , is provided ( Fig. 11a ).

There has been some confusion regarding the generic affinity of the species Humboltia spiralis Ruiz & Pavón (1798: 237) and its relationship to Pleurothallis macrorhiza Lindley (1834: 9) . Persoon (1807) erroneously transferred H. spiralis to Stelis . Stelis spiralis (Ruiz & Pav.) Persoon (1807: 524) was subsequently declared a synonym of Stelis purpurea (Ruiz & Pav.) Willdenow (1805 : pl. 4, 140). Hence, Humboltia spiralis , Pleurothallis spiralis , Stelis spiralis and Xenosia spiralis are all currently listed as synonyms of Stelis purpurea in the World Checklist of Selected Plant Families ( WCSP 2016). To further compound the confusion, Luer (2006) transferred P. macrorhiza to Xenosia , as Xenosia macrorhiza (Lindl.) Luer (2006: 233) , listing the previously described X. spiralis as a synonym. And in the World Checklist of Selected Plant Families Pleurothallis macrorhiza , Humboltia macrorhiza (Lindl.) Kuntze (1891: 667) and Xenosia macrorhiza are also listed as synonyms of S. purpurea . We have determined, however, that the type of H. spiralis is an Andinia , not a Stelis , and that H. spiralis and P. macrorhiza are likely the same species. Because H. spiralis Ruiz & Pavón (1798) has taxonomic priority over P. macrorhiza Lindley (1834) , the appropriate combination under Andinia is Andinia spiralis (Ruiz & Pav.) Karremans & Mark Wilson. All of these names, except S. purpurea , are therefore reduced to synonymy with A. spiralis .

Andinia longiserpens (C.Schweinf.) Karremans & Mark Wilson , comb. nov.

Bas. Pleurothallis longiserpens C.Schweinf., Bot. Mus. Leafl. 10: 183. 1942.

Andinia masdevalliopsis (Luer) Karremans & Mark Wilson, comb. nov.

Bas. Pleurothallis masdevalliopsis Luer, Phytologia 44(3): 170. 1979.

Andinia spiralis (Ruiz & Pav.) Karremans & Mark Wilson, comb. nov.

Bas. Humboltia spiralis Ruiz & Pav., Syst. Veg. Fl. Peruv. Chil. 1: 237. 1798.

Andinia xenion (Luer & R.Escobar) Karremans & Mark Wilson , comb. nov.

Bas. Pleurothallis xenion Luer & R.Escobar, Orquideología 16(1): 38. 1983.

Andinia subgen. Minuscula Karremans & Mark Wilson , subgen. nov. Type. Pleurothallis trimytera Luer & R. Escobar, Orquideología 16: 34, 1983.

Plant repent-ascending, leaves coriaceous, inflorescence loose, successively flowered, ovary papillose, lip three-lobed, with lobes triangular, conspicuous, apical, anther and stigma apical, column foot absent.

Etymology: The name refers to the minuscule size of the plants and flowers of this subgenus compared to the other subgenera.

Three species distributed through Andean regions of Ecuador and Colombia ( Fig. 6 ), with an elevational range of 1,800 to 2,590 m ( Fig. 8 ). The three species are illustrated ( Fig. 10 ) and a photograph of an example, Andinia trimytera (Luer & R.Escobar) Pridgeon & M.W.Chase , is provided ( Fig. 11d ).

Andinia hystricosa (Luer) Pridgeon & M.W.Chase, Lindleyana 16(4): 251. 2001.

Bas. Pleurothallis hystricosa Luer, Monogr. Syst. Bot. Missouri Bot. Gard. 52: 54. 1994.

Andinia panica (Luer & Dalström) Pridgeon & M.W.Chase, Lindleyana 16(4): 251. 2001.

Bas. Pleurothallis panica Luer & Dalström, Monogr. Syst. Bot. Missouri Bot. Gard. 61(3): 6. 1996.

Andinia trimytera (Luer & R.Escobar) Pridgeon & M.W.Chase, Lindleyana 16(4): 252. 2001.

Bas. Pleurothallis trimytera Luer & R.Escobar, Orquideología 16: 34. 1983.

Figures

FIGURE 6. Distributions of clades or proposed subgenera Aenigma, Andinia, Masdevalliantha and Minuscula.

FIGURE 8. Mean elevations and elevational ranges of collections from Tropicos for proposed subgenera Andinia; Aenigma; Masdevalliantha; Minuscula; all four subgenera combined (i.e. excluding Brachycladium); and Brachycladium. Blue triangles represent mean collection elevation (m) and black bar represents elevational range of collections.

FIGURE 9. Drawings of Andinia longiserpens (from Luer 2006) and A. masdevalliopsis (from Luer 1986b) and A. xenion (from Luer 1986). Courtesy of Missouri Botanical Garden Press.

FIGURE 10. Drawings of Andinia hystricosa and A. trimytera from Luer (1994) and A. panica from Luer (1996). Courtesy of Missouri Botanical Garden Press.

FIGURE 11. Representatives of the proposed infrageneric taxa: (a) subgenus Masdevalliantha - Andinia longiserpens (photo credit: Ron Parsons); (b) subgenus Aenigma - Andinia schizopogon; (c) subgenus Andinia - Andinia lappacea; (d) subgenus Minuscula - Andinia trimytera; (e) subgenus Brachycladium section Brachycladae - Andinia nummularia;and (f) subgenus Brachycladiumsection Amplectentes - Andinia montis-rotundi.