A Chondrorhyncha by any other (correct) name (Orchidaceae: Zygopetalinae)

Abstract

The genus Chondrorhyncha was originally described to accommodate a species "allied to Helcia and Trichopilia", and it was essentially defined by the oblique insertion of the sepals and its long, car-tilaginous, bristle-like rostellum, a condition likely plesiomorphic within the genera of the subtribe Zygopetalinae close to Huntleya. Since then, the genus served as a poorly defined basket to embrace a vast array of different taxa, collectively named the "Chondrorhyncha complex". Most of the genera of the Huntleya grade are easy to identify on the basis of their characteristic floral features, but generic delimitation in the Chondrorhyncha complex has been traditionally difficult, and many species have complex nomenclatural histories. This taxonomic confusion reflects repeated evolutionary changes in pollination mechanisms that produced homoplasious floral morphologies. Many species in this complex produce gullet flowers that appear to be nectar-deceit flowers for long-tongued visitors, with swept back and revolute lateral sepals that form a tubular false spur enclosing the notch on either side of the lip that permits the passage of a bee's tongue. Recent molecular work indicates that species with this deceit morphology are scattered among various clades with other floral mechanisms, and therefore genera based on gross floral morphology are polyphyletic. Most notably, the traditional concept of Chondrorhyncha is highly polyphyletic, with its members falling into eight strongly supported clades. However, unique sets of vegetative, floral, and pollinaria features allow in most of the cases a proper generic placement of the taxa belonging to the "Chondrorhyncha complex." The paper highlights the taxonomic history of the genus Chondrorhyncha and discusses the phylogenetic position of Chondro-rhyncha s.s. as well as its relationships with closely allied genera, reviewing the new generic circumscrip-tions within the Chondrorhyncha complex and the genera recently split from Chondrorhyncha, and discussing the improvements in the systematics of the group based on molecular data.

Full text

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Pupulin F (2009) A Chondrorhyncha by any other (correct) name (Orchidaceae: Zygopetalinae).
In: Pridgeon A M, Suarez JP (eds) Proceedings of the Second Scientic Conference on Andean Orchids.
Universidad Técnica Particular de Loja, Loja, Ecuador, pp 145-160.
A CHONDRORHYNCHA BY ANY OTHER (CORRECT) NAME
(ORCHIDACEAE: ZYGOPETALINAE)
Franco Pupulin
Jardín Botánico Lankester, Universidad de Costa Rica, P.O. Box 1031-7050, Cartago, Costa Rica;
Harvard University Herbaria, Cambridge, Massachusetts, USA; Marie Selby Botanical Gardens, Sarasota,
Florida, USA;
Ángel Andreetta Research Center on Andean Orchids, Universidad Alfredo Pérez Guerrero, Gualaceo, Ecua-
dor; E-mail: fpupulin@cariari.ucr.ac.cr
ABSTRACT
e genus Chondrorhyncha was originally described to accommodate a species “allied to Helcia
and Trichopilia”, and it was essentially dened by the oblique insertion of the sepals and its long, car-
tilaginous, bristle-like rostellum, a condition likely plesiomorphic within the genera of the subtribe
Zygopetalinae close to Huntleya. Since then, the genus served as a poorly dened basket to embrace a
vast array of dierent taxa, collectively named the “Chondrorhyncha complex”. Most of the genera of
the Huntleya grade are easy to identify on the basis of their characteristic oral features, but generic
delimitation in the Chondrorhyncha complex has been traditionally dicult, and many species have
complex nomenclatural histories. is taxonomic confusion reects repeated evolutionary changes
in pollination mechanisms that produced homoplasious oral morphologies. Many species in this
complex produce gullet owers that appear to be nectar-deceit owers for long-tongued visitors, with
swept back and revolute lateral sepals that form a tubular false spur enclosing the notch on either side
of the lip that permits the passage of a bee’s tongue. Recent molecular work indicates that species with
this deceit morphology are scattered among various clades with other oral mechanisms, and therefore
genera based on gross oral morphology are polyphyletic. Most notably, the traditional concept of
Chondrorhyncha is highly polyphyletic, with its members falling into eight strongly supported clades.
However, unique sets of vegetative, oral, and pollinaria features allow in most of the cases a proper
generic placement of the taxa belonging to the “Chondrorhyncha complex.” e paper highlights the
taxonomic history of the genus Chondrorhyncha and discusses the phylogenetic position of Chondro-
rhyncha s.s. as well as its relationships with closely allied genera, reviewing the new generic circumscrip-
tions within the Chondrorhyncha complex and the genera recently split from Chondrorhyncha, and
discussing the improvements in the systematics of the group based on molecular data.
e rst plant of what we consider today a true species of Chondrorhyncha was collected some
50 years before Lindley’s formal description of the genus, at the times of José Celestino Mutis’ Royal
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Figure 1. “Zygophilum.” Plate 464 of Iconograa Mutisiana. Courtesy of the Real Jardín Botánico, Madrid.

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Botanical Expedition of the Nuevo Reyno de Granada, which started in 1783 and concluded in the
rst decade of the 19th century. One of the almost 7000 botanical plates produced by the artists of the
expedition, now conserved at the Royal Botanic Garden of Madrid, depicts in full detail a Colombian
plant annotated as “Zygophilum” (Iconografía Mutisiana, pl. 464) and represents the rst documented
record of a Chondrorhyncha by western botanists (Fig. 1). e exact origin of this rst Chondrorhyncha
species was not recorded, but it was likely collected during the residence of the botanical expedition
at Mariquita (1783-1791), in northern Tolima province (Colombia), or in some of the other template
valleys of the Magdalena River (Pérez Arbeláez, 1995). Apparently, no description of this orchid was
conserved among the more than 800 descriptions and observations compiled by Mutis and the other
botanists of the expedition (Amaya, 1999), and no herbarium material of this plant is kept in the
Mutis collection at Madrid (MA-MUT) according to Pérez Arbeláez (1995).
It was not until 1846 when John Lindley, compiling the catalogue of a large orchid collection
assembled by Jean Linden during an expedition to Venezuela, Colombia, and Cuba, created the genus
Chondrorhyncha to accommodate a plant originally collected at “Jajá” [Jaji] (Lindley, 1846), along the
northern slopes of the Sierra Nevada. Among the diagnostic features of the new genus, Lindley (1846)
noted as most remarkable “the extremely oblique insertion of the sepals,” together with the bristle-like,
cartilaginous rostellum (hence the generic name, from the Greek chondros, cartilage, and rhynchos,
beak), and the four, “concavo-plana” (dorsiventrally complanate), overlapped pollinia in two pairs of
dierent size (Fig. 2). Nevertheless, in suggesting a phylogenetic placement of his new genus, Lindley
considered Chondrorhyncha to be allied to Helcia Lindl. and Trichopilia Lindl. (now considered con-
generic), two distantly related genera of subtribe Oncidiinae.
It is noteworthy that Lindley could not recognize the anities of Chondrorhyncha. At the time
of the original description of his new genus, Lindley had already published two genera closely related
to Chondrorhyncha, namely Stenia Lindl. (based on S. pallida Lindl., 1837) and Huntleya Batem. ex
Lindl. (typied by H. meleagris Lindl., 1837), and he had also described two other non-pseudobul-
bous species of Zygopetalinae as Zygopetalum cochleare [1837 = Cochleanthes abelliformis (Sw.) R.E.
Schult. & Garay] and Huntleya violacea [1839 = Pescatorea violacea (Lindl.) Dressler]. It is true that
the narrow, ligular pollinia of Stenia pallida are highly anomalous among the species of Stenia, a genus
which exhibits an unusual variation in pollinarium shape [and it is interesting that only two years later
Ranesque (1838) published the same taxon with the generic name of Stenopollen, alluding to the
same diagnostic character], but its rostellum is otherwise similar to that of Chondrorhyncha. Huntleya
meleagris, Cochleanthes abelliformis, and Pescatorea violacea, on the other hand, have both a bristle-
like rostellum and four, dorsiventrally superposed, complanate pollinia, coinciding with the characters
stated in the protologue of Chondrorhyncha. According to Lindley (1846), the only useful character
to distinguish Chondrorhyncha from its Huntleyoid relatives was the oblique insertion of the lateral
sepals, which became strongly reexed with age, a single feature that gave rise to what has been called
the “Chondrorhyncha complex.”
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Reichenbach (1852) segregated from a broad concept of Zygopetalum Hook. other “chondro-
rhynchous” genera based on diagnostic characters (apomorphies), including Chaubardia Rchb.f.,
Kefersteinia Rchb.f., Pescatorea Rchb.f., Bollea Rchb.f. (now included in Pescatorea, see Whitten et al.,
2005), and Warczewiczella Rchb.f. ese segregates have been accepted by most botanists with some
exceptions (i.e., Allen, 1949; Garay, 1969). More genera were split from Chondrorhyncha or created for
Chondrorhyncha-like species in the second half of the 20th century [i.e. Chaubardiella Garay (1969),
Dodsonia Ackerman (1979), Benzingia Dodson (1989), Ackermania Dodson & R.Escobar (1993), and
Chondroscaphe (Dressler) Senghas & G.Gerlach (1993)]. Nevertheless, by the end of the century the
list of names in the genus Chondrorhyncha had reached the respectful gure of almost 70 binomials
(Table 1).
Figure 2. Detail from the holotype of Chondrorhyncha rosea. Courtesy of the Orchid Herbarium, Royal Botanic Gar-
dens, Kew.

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Chondrorhyncha alba (Schltr.) L.O.Williams, Ceiba 5: 191. 1956 = Kefersteinia alba Schltr.
Chondrorhyncha albicans Rolfe, Bull. Misc. Inform. Kew 1898: 195. 1898 = Daiotyla albicans (Rolfe) Dressler
Chondrorhyncha amabilis Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7: 162. 1920 = Chondroscaphe amabilis (Schltr.)
Senghas & G.Gerlach
Chondrorhyncha amazonica (Rchb.f.) A.D.Hawkes, Encycl. Cult. Orchids, 110. 1965 = Warczewiczella amazonica Rchb.
f. & Warsz.
Chondrorhyncha anatona (Dressler) Senghas, Orchidee (Hamburg) 41: 91. 1990 = Euryblema anatonum (Dressler)
Dressler
Chondrorhyncha andreae P.Ortiz, Orquideología 19: 13. 1994 = Euryblema andreae (P. Ortiz) Dressler
Chondrorhyncha andreettae Jenny, Orchidee (Hamburg) 40: 92. 1989 = Aetheorhyncha andreettae (Jenny) Dressler
Chondrorhyncha antonii P.Ortiz, Orquideología 19: 14. 1994 = Echinorhyncha antonii (P.Ortiz) Dressler
Chondrorhyncha aromatica (Rchb.f.) P.H.Allen, Ann. Missouri Bot. Gard. 36: 85. 1949 = Cochleanthes aromatica (Rchb.
f.) R.E.Schult. & Garay
Chondrorhyncha aurantiaca Senghas & G.Gerlach, Orchidee (Hamburg) 42: 282. 1991 = Ixyophora aurantiaca
(Senghas & G.Gerlach) Dressler
Chondrorhyncha bicolor Rolfe, Bull. Misc. Inform. Kew 1894: 393. 1894 = Chondroscaphe bicolor (Rolfe) Dressler
Chondrorhyncha caloglossa (Schltr.) P.H.Allen, Ann. Missouri Bot. Gard. 36: 85. 1949 = Stenotyla picta (Rchb.f.) Dressler
Chondrorhyncha caquetae Fowlie, Orch. Digest, 32: 145. 1968.
Chondrorhyncha carinata P.Ortiz, Orquideología 19: 18. 1994 = Ixyophora carinata (P.Ortiz) Dressler
Chondrorhyncha caudata Ackerman, Selbyana 5: 299. 1981 = Benzingia caudata (Ackerman) Dressler
Chondrorhyncha chestertoni Rchb.f., Gard. Chron. 1879: 648. 1879 = Chondroscaphe chestertonii (Rchb.f) Senghas &
G.Gerlach
Chondrorhyncha cornuta Garay, Orquideología 5: 20. 1970 = Benzingia cornuta (Garay) Dressler
Chondrorhyncha costaricensis (Schltr.) P.H.Allen, Ann. Missouri Bot. Gard. 36: 86. 1949 = Kefersteinia costaricensis
Schltr.
Chondrorhyncha crassa Dressler, Orchidee (Hamburg) 34: 222. 1983 = Daiotyla crassa (Dressler) Dressler
Chondrorhyncha discolor (Lindl.) P.H.Allen, Ann. Missouri Bot. Gard. 36: 87. 1949 = Warczewiczella discolor (Lindl.)
Rchb.f.
Chondrorhyncha eburnea Dressler, Orchidee (Hamburg) 34: 224. 1983 = Chondroscaphe eburnea (Dressler) Dressler
Chondrorhyncha ecuadorensis Dodson, Icon. Pl. Trop. Ser. 2, 5: t. 415. 1989 = Echinorhyncha ecaudorensis (Dodson)
Dressler
Chondrorhyncha embreei Dodson & Neudecker, Orquideología 19: 82. 1993 = Chondroscaphe embreei (Dodson &
Neudecker) C.Rungius ex C.Rungius
Chondrorhyncha endresii Schltr., Repert. Spec. Nov. Regni Veg. 17: 14. 1921 = Chondroscaphe endresii (Schltr.) Dressler
Chondrorhyncha escobariana Dodson & Neudecker, Orquideología 19: 47. 1993 = Chondroscaphe escobariana
(Dodson & Neudecker) C. Rungius ex C. Rungius
Chondrorhyncha estradae Dodson, Icon. Pl. Trop. 1: t. 22. 1980 = Benzingia estradae (Dodson) Dressler
Chondrorhyncha estrellensis Ames, Sched. Orchid. 4: 54. 1923 = Stenotyla picta (Rchb.f.) Dressler
Chondrorhyncha mbriata Linden & Rchb.f., Refug. Bot. (Saunders) t. 107. 1878 = Chondroscaphe mbriata (Linden &
Rchb.f.) Dressler
Chondrorhyncha abelliformis (Sw.) Alain, Phytologia 8: 369. 1962 = Cochleanthes abelliformis (Sw.) R.E.Schult. &
Garay
Table 1. Published binomials in the genus Chondrorhyncha.
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Chondrorhyncha aveola (Linden & Rchb.f. ex Rchb.f.) Garay, Bot. Mus. Lea. 21: 256. 1967 = Chondroscaphe aveola
(Linden & Rchb.f. ex Rchb.f.) Senghas & G.Gerlach
Chondrorhyncha fosterae Dodson, Selbyana 7: 357. 1984 [likely not a true Chondrorhyncha]
Chondrorhyncha gentryi Dodson & Neudecker, Orquideología 19: 49. 1993 = Chondroscaphe gentryi (Dodson &
Neudecker) C.Rungius ex C.Rungius
Chondrorhyncha guttata (Rchb.f.) Garay, Bot. Mus. Lea. 26: 27. 1978 = Stenia guttata Rchb.f.
Chondrorhyncha helleri Fowlie, Orch. Digest 35: 170. 1971 = Stenotyla helleri (Fowlie) Pupulin, comb. ined.
Chondrorhyncha hirtzii Dodson, Icon. Pl. Trop. Ser. 2, 5: t. 416. 1989 = Benzingia hirtzii (Dodson) Dressler
Chondrorhyncha inedita Dressler & Dalström, Orquideología 23: 80. 2004.
Chondrorhyncha lactea (Rchb.f.) L.O.Williams, Caldasia 5: 16. 1942. = Kefersteinia lactea (Rchb.f.) Schltr.
Chondrorhyncha lankesteriana Pupulin, Lindleyana 15: 21. 2000 = Stenotyla lankesteriana (Pupulin) Drssler
Chondrorhyncha lendyana Rchb.f., Gard. Chron. n.s. 26: 103. 1886 = Stenotyla lendyana (Rchb.f.) Dressler
Chondrorhyncha lipscombiae Rolfe, Bull. Misc. Inform. Kew 1912: 133. 1912 = Warczewiczella lipscombiae (Rolfe)
Fowlie
Chondrorhyncha litensis Dodson, Icon. Pl. Trop. Ser. 2, 5: t. 417. 1989 = Echinorhyncha litensis (Dodson) Dressler
Chondrorhyncha lojae (Schltr.) C.Schweinf., Bot. Mus. Lea. 11: 216. 1944. = Kefersteinia lojae Schltr.
Chondrorhyncha luerorum R.Vásquez Ch. & Dodson, Rev. Soc. Boliv. Bot. 2: 1. 1998.
Chondrorhyncha macronyx Kraenzl., Notizbl. Bot. Gart. Berlin-Dahlem 7: 414. 1920.
Chondrorhyncha maculata Garay, Orquideología 4: 161. 1969 = Daiotyla maculata (Garay) Dressler
Chondrorhyncha manzurii P.Ortiz, Revista Acad. Colomb. Ci. Exact. 24: 57. 2000.
Chondrorhyncha marginata (Rchb.f.) P.H.Allen, Ann. Missouri Bot. Gard. 36: 88. 1949 = Warczewiczella marginata
Rchb.f.
Chondrorhyncha merana Dodson & Neudecker, Orquideología 19: 83. 1993 = Chondroscaphe merana (Dodson &
Neudecker) Dressler
Chondrorhyncha microcharis (Schltr.) L.O.Williams, Ceiba 5: 194. 1956 = Kefersteinia microcharis Schltr.
Chondrorhyncha palorae (Dodson & Hirtz) Senghas & G.Gerlach, Schlechter Orchideen, 1/B(26): 1631. 1992 = Benzingia
palorae (Dodson & Hirtz) Dressler
Chondrorhyncha parvilabris (Schltr.) L.O.Williams, Ceiba 5: 195. 1956 = Kefersteinia parvilabris Schltr.
Chondrorhyncha picta (Rchb.f.) Senghas, Orchidee (Hamburg) 41: 94. 1990 = Stenotyla picta (Rchb.f.) Dressler
Chondrorhyncha plicata D.E.Benn. & Christenson, Brittonia 46: 24. 1994 = Chondroscaphe plicata (D.E.Bnn. &
Christenson) Dressler
Chondrorhyncha pusilla C.Schweinf., Amer. Orchid Soc. Bull. 12: 384, g. 1944 = Kefersteinia lojae Schltr.
Chondrorhyncha reichenbachiana Schltr., Repert. Spec. Nov. Regni Veg. 17: 15. 1921 = Benzingia reichenbachiana
(Schltr.) Dodson
Chondrorhyncha rhombilabium (C.Schweinf.) Fowlie, Orch. Dig., 48: 225. 1984 = Chaubardia heteroclita (Poepp. &
Endl.) Dodson & D.E.Benn.
Chondrorhyncha rosea Lindl., Orchid. Linden. 13. 1846.
Chondrorhyncha stapelioides (Rchb.f.) L.O.Williams, Ceiba 5: 195. 1956 = Kefersteinia stapelioides Rchb.f.
Chondrorhyncha stenioides Garay, Bot. Mus. Lea. 26: 26. 1978 = Stenia stenioides (Garay) Dodson & R. Escobar
Chondrorhyncha suarezii Dodson, Icon. Pl. Trop. Ser. 2, 5: t. 418. 1989.
Chondrorhyncha subquadrata (Schltr.) L.O.Williams, Ceiba 5: 195. 1956 = Chaubardiella subquadrata (Schltr.) Garay
Chondrorhyncha thienii (Dodson) Dodson, Selbyana 7: 354. 1984 [doubtfully a true Chondrorhyncha]
Chondrorhyncha velastiguii Dodson, Icon. Pl. Trop. Ser. 2, 5: t. 419. 1989 = Ixyophora velastiguii (Dodson) Pupulin,
comb. ined.

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Although many genera of Zygopetalinae are well known in horticulture and are cultivated on
large scale for commercial purposes (e.g., Zygopetalum), and some are relatively easy to identify on the
basis of their characteristic oral features (e.g., Pescatorea/Bollea, Huntleya, Kefersteinia, among oth-
ers), the nomenclatural ination of Chondrorhyncha reects the traditional diculty in delimitating
the remaining genera of the Huntleya clade. Attempts to revise critically orchid groups belonging to
the so-called Chondrorhyncha complex were made by Garay (1969, 1970), Fowlie (1969), and Dressler
(2000), the latter study based at least in part on the preliminary results of molecular work carried out
by W. M. Whitten and collaborators at the University of Florida. However, it was not until ve years
later that a new phylogenetic arrangement of the entire subtribe Zygopetalinae was proposed on the
basis of analyses of four DNA regions data sets (Whitten et al., 2005).
According to the Whitten et al. (2005) study, the subtribe, which includes more than 400
species and ranges from Mexico, through the West Indies, to Brazil, Bolivia and Argentina in South
America, is composed of three main groups. e basal grade includes all the genera with pseudobulbs
composed of several nodes (homoblastic) and plicate leaves related to Warrea Lindl., consecutively sis-
ter to the grade mostly composed by the genera related to Zygopetalum, with single-node pseudobulbs,
convolute leaves, and many-owered inorescences; and to the usually pseudobulbless, conduplicate-
leaved, one-owered genera of the Huntleya clade (including the two morphologically anomalous
genera Cryptarrhena R.Br. and Dichaea Lindl.). Within the better-sampled clade of Huntleya and the
allied genera, the phylogenetic trees clearly showed that Chondrorhyncha was grossly polyphyletic, with
its members being scattered in at least eight dierent clades (Fig. 3).
e results of the molecular analyses were not surprising in light of the pollination mechanisms
already known in the group. In most of the documented records (species of Benzingia Dodson ex
Dodson, Chaubardia, Chaubardiella, Cochleanthes, Dichaea, Huntleya, Kefersteinia, Pescatorea, Warcze-
wiczella, and Zygosepalum Rchb.f.), the owers of Zygopetalinae attract male euglossine bees searching
for perfume compounds. However, Ackerman (1983) also recorded visits to Cochleanthes (=Warcze-
wiczella) lipscombiae in Panama by female euglossine looking for food resources, which extend their
tongue into the backswept lateral sepals in search of nectar. ese strongly reexed lateral sepals with
the margins infolded at the base mimic nectariferous spurs. Species with this deceit morphology are
scattered in the Chondrorhyncha complex among various groups with other oral mechanisms. As
demonstrated in other advanced orchid groups such as Maxillariinae and the Oncidiinae in which
Chondrorhyncha viridisepala Senghas, Orchidee (Hamburg) 40: 181. 1989 = Ixyophora viridisepala (Senghas) Dressler
Chondrorhyncha vollesii G.Gerlach, Neudecker & Seeger, Orchidee (Hamburg) 40: 131. 1989 = Echinorhyncha vollesi
(Lafontaine, G.Gerlach & Senghas) Dressler
Chondrorhyncha wailesiana (Lindl.) A.D.Hawkes, Encycl. Cult. Orchids 111. 1965 = Warczewiczella wailesiana (Rchb.f.)
E. Morren
Chondrorhyncha wercklei (Schltr.) C.Schweinf., Amer. Orchid Soc. Bull. 12: 386. 1944 = Kefersteinia wercklei Schltr.
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a mix of dierent pollination syndromes are present among closely related taxa, the resulting genera
mainly circumscribed on the basis of gross ower morphology are frequently polyphyletic (Whitten et
al., 2000; Williams et al., 2001; Ojeda et al., 2003; Whitten et al., 2007). In this case, the taxonomic
confusion that plagued the history of Chondrorhyncha reects repeated evolutionary changes in pol-
lination mechanisms, with mixtures of fragrance reward and deceit pollination.
Figure 3. Randomly chosen, most-parsimonious tree resulting from cladistic analyses of combined matK/trnL-F/ITS
rDNA data set for the mainly pseudobulbless groups of the Zygopetalinae tree. Modied from Whitten et al. (2005).

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As a nomenclatural result of the phylogenetic study, many genera in Zygopetalinae required
recircumscription, and new genera were created to reect the molecular evidence better (Whitten et
al., 2005). Most of the newly dened genera have clear vegetative and/or oral morphological synapo-
morphies. Neudecker and Gerlach (2000) anticipated the merge of Dodsonia into Stenia in describing
the morphologically intermediate Stenia glatzii Neudecker & G.Gerlach. Dressler (2001) argued for
the inclusion of the Chondrorhyncha bicolor group into Chondroscaphe (Fig. 4, A). e need to distin-
guish Warczewiczella from Cochleanthes [so reducing the latter genus to the actual gure of two species
(Pupulin 2006)] was already suggested by Fowlie almost 40 years ago (Fowlie, 1969) (Fig. 4, B—C).
e merging of Bollea into Pescatorea easily explains the relative frequency of so-called “intergeneric”
natural hybrids among these two supposed genera and shows that the main distinguishing characters
among the two groups (i.e., the width of the column wings and basal callus of the lip) are taxonomi-
cally trivial (Fig. 4, D—E). e case of Benzingia deserved special mention; this clade includes diverse
oral morphologies, but most species possess a pendent growth habit and gray-green foliage with a mi-
nutely papillose surface. In the phylogenetic tree (Whitten et al., 2005), the type species of Benzingia,
B. hirtzii Dodson, is in a moderately supported clade together with Chondrorhyncha reichenbachiana
Schltr., species of Ackermania, and Benzingia estradae Dodson (Fig. 4, F—I). Surprisingly, the sister
species of B. hirtzii is C. reichenbachiana (90/100% support), and the only other species transferred
to Benzingia before the phylogenetic analysis, B. estradae, is weakly supported as sister to Ackermania
cornuta (Garay) Dodson & R.Escobar, described as Chondrorhyncha (Garay 1970). Ackermania cau-
data (Ackerman) Dodson & R.Escobar and A. hajeckii D.E.Benn. & Christenson are strongly sup-
ported (96/100%) as sister taxa. e results of the combined molecular analyses leave some room for
taxonomic interpretation, but the recircumscription of Benzingia to include Ackermania and Chondro-
rhyncha reichenbachiana is by far the most rational, although the resulting genus is dicult to dene
in terms of vegetative and morphological features. e alternative would have been to recognize a
Benzingia s.s. for B. hirtzii and Chondrorhyncha reichenbachiana (but excluding B. estradae), creating
a new genus for Benzingia estradae + Ackermania cornuta, and another genus (the name Ackermania is
likely predated by the fungal genus Ackermannia Pat.) for A. caudata + A. hajeckii. With the exception
of Aetherorhyncha Dressler (Fig. 4, J), which is weakly supported as sister to the genus now treated
as Ixyophora Dressler, all the other new genera split from Chondrorhyncha (namely Daiotyla Dressler,
Echinorhyncha Dressler, Euryblema Dressler, Ixyophora, and Stenotyla Dressler) are on strongly sup-
ported branches (Fig. 4, K—L; 5, A—C) and easy to characterize morphologically.
It is perhaps legitimate to ask how the new generic alignments produced by the use of molecu-
lar techniques improve our understanding of the Chondrorhyncha complex with respect to previous
systematic approaches. I will address this question by four main points.
First, morphological characterization is needed to explain evolutionary processes and how they
act on certain features of the organisms. However, if morphology is to be maintained as a tool to
interpret life’s diversity and propose generic and specic anities, accurate observations are essential.
A few cases may perhaps illustrate this point. Among the species of the Chondrorhyncha complex,
Aetheorhyncha andreettae (Jenny) Dressler is the only one to produce almost erect owers on short and
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Figure 4. A. Chondroscaphe yamilethiae (Pupulin 4701); B. Warczewiczella marginata (Pupulin 6611); C. Cochleanthes
abelliformis (Pupulin 5946); D. Pescatorea lamellosa (CIOA-00322); E. Pescatorea ecuadorana (Pupulin 6957); F. Benz-
ingia hirtzii (Pupulin 6996); G. Benzingia reichenbachiana (Pupulin 3228); H. Benzingia estradae (Pupulin 6900); I. Benz-
ingia cornuta (Pupulin 5879); J. Aetheorhyncha andreettae (Pupulin 6560); K. Echinorhyncha antonii (Pupulin 6556); L.
Daiotyla albicans (Pupulin 3510).

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stout inorescences (Fig. 4, J). e entrance to the lip chamber is ornamented with short, at and sti
trichomes, and it is unusually narrow, so that we have to presume that only a relatively small bee may
gain access to the chamber. Inside the chamber, the column of A. andreettae is twisted longitudinally
(Fig. 6), indicating that the pollinarium is probably placed on one side behind the head or on the back
of the pollinator. It is interesting that these morphological features, which play a substantial role in
the pollination biology of this species, were not previously emphasized. e species actually assigned
to the genus Stenotyla are invariably provided with small but distinct pseudobulbs. e callus of the
Figure 5. A. Ixyophora viridisepala (Pupulin 5941); B. Stenotyla picta (Pupulin 3027); C. Euryblema anatonum (Pupulin
6282); D. Zygopetalinae species, probably belonging to a undescribed genus (Pupulin 6605); E. “Chondrorhyncha”
thienii (Pupulin 6281); F. Chondrorhyncha hirtzii (Pupulin s.n.); G. Chondrorhyncha inedita, watercolor of the type ow-
er, Schlim 34 (Courtesy of the Orchid Herbarium, Royal Botanic Gardens, Kew); H. Chondrorhyncha sp., Colombia,
watercolor from Iconografía Mutisiana, pl. 464 (courtesy of the Real Jardín Botánico, Madrid).
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Figure 6. Lateral view of the column and lip of Aetheorhyncha andreettae, from Pupulin 6560 (the lip sectioned). Note
the longitudinal twisting of the column. Scale bar = 1 cm.
Figure 8. Pollinarium of Euryblema anatonum (A, B. Pupulin 6282).
Scale bar = 5 mm.
Figure 7. Pollinarium of Stenotyla species. A, B. Stenotyla picta (JBL-01970;
JBL-01905); C. Stenotyla panamensis (Pupulin 5867); D. Stenotyla lendyana
(JBL-02053). Scale bar = 5 mm.

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lip is short and narrow, placed almost in the center of the blade, and it has only 2-4 distinct teeth. e
pollinarium of Stenotyla is characteristic, with a narrow, triangular stipe that curls after removal and a
small, semitubular and almost indistinct viscidium (Pupulin, in press; Fig. 7). ese features were not
consistently described in the past. e genus Euryblema is easily characterized by the leaf sheaths that
are boldly marked with purple (indigenous people from the region of Bocas del Toro in Panama refer
Figure 12. Pollinarium of a still undescribed species of Zy-
gopetalinae (Pupulin 6605). From left to right: abaxial view,
two adaxial views, and lateral view. Scale bar = 5 mm.
Figure 9. Column of Echinorhyncha antonii, in ventral (left)
and lateral views (right), showing the ventral appendages
(arrows), from Pupulin 6556. Scale bar = 1 cm.
Figure 11. Pollinarium of Chondrorhyncha thienii (adaxial,
oblique, and ventral views), from Pupulin 6924. Scale bar =
3 mm.
Figure 10. Labella of “Chondrorhyncha” velastiguii (A) and
Ixyophora viridisepala (B). A. Pupulin 6622; B. Pupulin 5941.
Scale bar = 3 cm.
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to E. anatonum as “patirroja,” or red-footed); the broad and laminar callus completely covers the basal
half of the lip, and the pollinarium is provided with a large, broad stipe, and a narrow, thick, ligular
viscidium, unlike any other species in the Chondrorhyncha complex (Pupulin, 2007; Fig. 8). In all
known species currently assigned to the genus Echinorhyncha, the ventral surface of the column bears
bristly, urchin-like appendages (Fig. 9) and the pollinarium has a pandurate stipe, two morphological
features that easily characterize the genus if consistently described. ese features may be obscured or
overlooked in poorly prepared herbarium specimens, and access to cultivated material has facilitated
comparison of overlooked characters.
Second, the new systematic arrangement of genera previously included under a broad Chondro-
rhyncha improves the predictability of generic placement in the case of dubious and not yet sampled
species. Chondrorhyncha velastiguii Dodson was originally described and illustrated by Dodson in
1989 on the base of a plant collected along the Río Negro in eastern Ecuador (Dodson and Dodson,
1989; Fig. 5, F). To my knowledge, the species has been collected only ve times after the original col-
lection, always from the eastern Ecuadorian provinces of Morona-Santiago, Pastaza, and Tungurahua.
In realigning Chondrorhyncha and the allied genera, Whitten et al. (2005) had no material to sample
C. velastiguii and include it in the data matrix, but they suggested it can likely considered a true species
of the genus Chondrorhyncha s.s. I recently had the opportunity to study living material of C. velastiguii
from Ecuador, and the vegetative morphology, as well as the shape of the lip callus (Fig. 10) and the
pollinarium agree with the set of characters now considered diagnostic for the genus Ixyophora, so
requiring a new nomenclatural combination for C. velastiguii.
ird, a good taxonomic system improves the recognition of anomalous patterns. Whitten et
al. (2005) were not able to obtain material of Cochleanthes thienii Dodson for their study, and they
considered this species to be possibly congeneric with the two known species of Euryblema. However,
the recent examination of new living specimens of C. thienii (Fig. 5, E) revealed several morphologi-
cal traits that are anomalous with respect to the diagnostic characters of the genus Euryblema. e leaf
sheaths of Euryblema are heavily spotted and blotched with purple, but this feature is absent in plants
of C. thienii. e lip of C. thienii is deeply cymbiform and provided with a central, narrow callus, con-
trasting with the open lip of Euryblema, provided with a broad, laminar callus. Also, the pollinarium of
Euryblema has a large, sagittate stipe and a ventrally distinct, ligular, thick viscidium (Pupulin, 2007),
whereas in C. thienii the stipe is small, shield-shaped, and the viscidium is a large but scarcely distinct
adhesive pad (Fig. 11).
Finally, molecular data have the benecial eect of reminding botanists that new species are
nothing more than scientic hypotheses, and such data provide a revolutionary method to test them.
is is true, of course, not only for the complex of genera and species close to Chondrorhyncha and not
only because of molecular systematics. Although frequently neglected, the need to compare morpho-
logical information with data sets from other disciplines (i.e., biochemistry, cytology, anatomy, ecol-
ogy, etc.) has been generally recognized among the requisites of standard taxonomic practice. While
examining Zygopetalinae specimens during a recent visit to Ecuador, I found a species that apparently

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does not t any of the known genera related to Chondrorhyncha (Fig. 5, D). e large, yellow ower
has a distinctly three-lobed lip with a laminar callus restricted to the center of the blade, and the pol-
linarium includes a large viscidium provided with lateral teeth and a cuspidate, sigmoid stipe (Fig.
12), two features not previously recorded among the Zygopetalinae. On the basis of morphological
comparison, I strongly suspect that this not only represents a new species but an undescribed genus in
Zygopetalinae. Names in the “Chondrorhyncha complex” by far exceed the actual number of species,
and it is perhaps advisable to refrain from the urgency to describe new taxa in this group without the
necessary evidence. Now, we fortunately have more tools to test our hypotheses, and as soon as permits
for molecular analyses are issued by the Ecuadorian authorities, we will sequence this hypothetical new
genus to place it into the phylogenetic framework of Zygopetalinae.
What about Chondrorhyncha? It is unfortunate that, more than two centuries after the rst
western botanists discovered a plant of Chondrorhyncha, our knowledge of this genus is still fragmen-
tary owing to the lack of available material for many of the taxa. We have now a better picture of
what is not a true Chondrorhyncha, and we can, by default, infer some information about this genus.
Chondrorhyncha s.s. is strictly South American in distribution, ranging from Venezuela to Bolivia along
the western slopes of the Andes, with a probable center of diversity in northwestern South America,
where most of the species have been recorded from Colombia and Ecuador (Fig. 5, G—I). e actual
number of Chondrorhyncha species is still debatable. We demonstrated that C. velastiguii, still included
in the genus, must be segregated on the basis of a set of morphological characters. I also suspect that C.
fosterae Dodson is probably misplaced in Chondrorhyncha. Although Dressler and Dalström (2004), in
the rst synopsis of Chondrorhyncha s.s. reduced the genus to only ve species, I would suggest that a
gure of 7-9 species is probably closer to reality. Chondrorhyncha species may be mainly characterized
morphologically by the presence of a narrow, not eshy callus near the middle of the lip, wide basally
and tapering at apex, and by the pollinarium provided with a narrowly triangular-peltate stipe with a
viscidium that becomes scarcely distinct toward the apical portion of the stipe. e owers of Chon-
drorhyncha are always resupinate, with the lateral sepals sometimes subfalcate and subuncinate at the
apex, spreading-recurved to abruptly reexed, concave, strongly inrolled-folded toward the base. e
petals are obtuse and gently recurved at apex. e lip, sometimes three-lobed, is rounded-subsaccate at
the base, with the apex obtuse to emarginate or minutely bilobed and the basal margins erect to ank
the column. Species of Chondrorhyncha occur as shaded epiphytes, mostly in evergreen premontane
wet forests at 500-1800 m along the eastern slopes of the Andes. eir owering has been recorded
throughout the year, with a owering peak from January to July.
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