LANKESTERIANA 7(3): 515-537. 2007.
GENERIC REALIGNMENTS IN MAXILLARIINAE (ORCHIDACEAE)
MARIO A. BLANCO1,2, GERMAN CARNEVALI3, W. MARK WHITTEN4, RODRIGO B. SINGER5,
SAMANTHA KOEHLER6, NORRIS H. WILLIAMS4, ISIDRO OJEDA7, KURT M. NEUBIG1
& LORENA ENDARA1
1
Department of Botany, University of Florida, 220 Bartram Hall, Gainesville, FL 32611-8526, USA.
(mablanco@ufl.edu, kneubig@ufl.edu, lendara@flmnh.ufl.edu)
2
Jardín Botánico Lankester, Universidad de Costa Rica, Apdo. 1031-7050, Cartago, Costa Rica
3
Herbario CICY, Centro de Investigación Científica de Yucatán (CICY), calle 43 No. 130,
Col. Chuburná de Hidalgo, 97200 Mérida, Yucatán, México. (carneval@cicy.mx)
4
Florida Museum of Natural History, University of Florida, P.O. Box 117800, Gainesville, FL 32611-7800, USA
(whitten@flmnh.ufl.edu, orchid@flmnh.ufl.edu)
5
Departamento de Botânica, Instituto de Biociências, Universidade Federal de Rio Grande do Sul,
CEP 91501-970, Porto Alegre, RS, Brazil. (rbsinger1@yahoo.com)
6
Departamento de Genética, Escola Superior de Agricultura “Luiz de Queiróz”, Universidade de São Paulo,
C.P. 83, Piracicaba, SP, Brazil 13400-970. (samantha.koehler@gmail.com)
7
Center for Plant Research, University of British Columbia, #302-Macmillan Building, 2357 Main Mall,
Vancouver, B.C., Canada V6T 1Z4. (isidro@interchange.ubc.ca)
ABSTRACT. A recent phylogenetic analysis of four DNA regions for ca. 354 species of core Maxillariinae
strongly indicate that the genus Maxillaria, as traditionally circumscribed, is grossly polyphyletic. We present a new phylogenetic classification for core Maxillariinae that recognizes 17 genera. Necessary realignments include: 1) resurrection of the genera Camaridium, Heterotaxis, and Ornithidium; 2) recognition of
the recent segregates Brasiliorchis (=Maxillaria sect. Repentes), Christensonella (=Maxillaria sect.
Urceolatae), Nitidobulbon (in press), and a recircumscribed Sauvetrea (=Maxillaria sect. Trigonae); 3)
adoption of the new genera Inti (=Maxillaria sect. Polyphyllae), Mapinguari, Maxillariella (=Maxillaria
sections Ebulbes and Erectae), and Rhetinantha; 4) transfers from Maxillaria sect. Reflexae to Ornithidium,
and Maxillaria sect. Rufescens to Mormolyca; and 5) synonymizing of the genera Adamanthus,
Pseudomaxillaria, Psittacoglossum, and Sepalosaccus (under Camaridium), Anthosiphon (under
Cryptocentrum), Chrysocycnis (under Mormolyca), Dicrypta, Marsupiaria, and Pentulops (under
Heterotaxis), and Laricorchis, Neo-urbania, and Siagonanthus (under Ornithidium). Some new synonyms at
the specific level are also presented.
RESUMEN. Un reciente análisis filogenético de cuatro regiones de ADN para ca. 354 especies de la subtribu
Maxillariinae indican fuertemente que el género Maxillaria, en su circunscripción tradicional, es altamente
polifilético. Presentamos una nueva clasificación filogenética para Maxillariinae que reconoce 17 géneros.
Los cambios necesarios incluyen: 1) la resurrección de los géneros Camaridium, Heterotaxis, y
Ornithidium; 2) el reconocimiento de los recientes segregados genéricos Brasiliorchis (=Maxillaria sección
Repentes), Christensonella (=Maxillaria sección Urceolatae), Nitidobulbon (en prensa), y una Sauvetrea
recircunscrita (=Maxillaria sección Trigonae); 3) la adopción de los nuevos géneros Inti (=Maxillaria sección Polyphyllae), Mapinguari, Maxillariella (=Maxillaria secciones Ebulbes y Erectae), y Rhetinantha; 4)
transferencias de Maxillaria sección Reflexae a Ornithidium, y Maxillaria sección Rufescens a Mormolyca;
y 5) puesta en sinonimia de los géneros Adamanthus, Pseudomaxillaria, Psittacoglossum y Sepalosaccus
(bajo Camaridium), Anthosiphon (bajo Cryptocentrum), Chrysocycnis (bajo Mormolyca), Dicrypta,
Marsupiaria y Pentulops (bajo Heterotaxis), y Laricorchis, Neo-Urbania, y Siagonanthus (bajo
Ornithidium). Algunos sinónimos nuevos al nivel de especie también son presentados.
KEY WORDS: Cymbidieae, generic recircumscription, Inti, Mapinguari, Maxillariella, Maxillariinae,
Orchidaceae, Rhetinantha.
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LANKESTERIANA
Subtribe Maxillariinae (sensu Chase et al. 2003)
includes a number of genera endemic to the
Neotropics. The circumscription of genera in the
core Maxillariinae (sensu Whitten et al. 2007, equal
to Maxillariinae sensu Dressler, 1993) has been
problematic since the creation of the genus
Maxillaria by Ruiz and Pavón (1794, 1798). This
large genus (ca. 580 species as traditionally defined;
e.g., Govaerts et al. 2005) has for a long time been
considered an assemblage of morphologically disparate taxa (Christenson 2002a, 2002b; Whitten et
al. 2007 and references therein), and thus probably
non-monophyletic. The complicated species-level
taxonomy of Maxillaria has hampered attempts of
providing a subgeneric classification (Christenson
2002a, 2002b1), a situation mirrored in other large
plant genera (Pfeil & Crisp 2005, Monro 2006,
Parnell et al. 2007).
The recent molecular phylogenetic analysis of
Dathe & Dietrich (2006) provided the first concrete
evidence for the grossly polyphyletic nature of
Maxillaria. However, the limited taxon sampling (30
species, one individual each) and low number of
DNA regions used in their study (nrITS only) precluded any sound taxonomic decisions based on their
results. More recently, Whitten et al. (2007) presented a vastly more detailed phylogenetic analysis of
core Maxillariinae (619 individuals representing ca.
354 species; four DNA regions used) that corroborated Dathe and Dietrich’s preliminary results and
revealed a worst-case scenario: all the currently
accepted minor genera of core Maxillariinae
1
Christenson (2002a) presented a partial subgeneric classification of Maxillaria with 19 sections, 13 of them newly
proposed. A French translation of the same article
(Christenson 2002b) was meant to appear much later, but
the publication of the Proceedings of the 16 th World
Orchid Conference (here abbreviated “Proc. 16th World
Orchid Conf.”) was delayed substantially. The two publications appeared in 2006, and the exact date of printing is
not indicated in either one. The Proceedings were published ca. six weeks before the translation in Richardiana,
and thus constitute the place of valid publication of most
of Christenson’s sections. The only exception is section
Ornithidium, which missed the basionym citation in the
Proceedings, and was validated in Richardiana (E. A.
Christenson, personal comunication 2007).
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
(Anthosiphon, Cryptocentrum, Chrysocycnis,
Cyrtidiorchis, Mormolyca, Pityphyllum, and
Trigonidium, Govaerts et al. 2005), are firmly nested
in Maxillaria sensu lato. Thus, the current generic
classification in core Maxillariinae is untenable on
phylogenetic grounds.
Backlund & Bremer (1998) presented guidelines on
how to modify existing classifications in order to
comply with phylogenetic requirements. Their primary principle is to designate only monophyletic groups
as ranked taxa. Secondary principles (in no particular
order of importance) are: a) to designate only highly
supported clades as ranked taxa; b) to minimize
nomenclatural disruption; c) to designate easily recognizable clades as ranked taxa; and d) to minimize
taxonomic redundancy by lumping monotypic taxa
with their sister group, if feasible. Entwisle & Weston
(2005) independently advanced similar guidelines,
although they also recommended to minimize taxonomic change in “charismatic” or economically
and/or horticulturally important groups, and to avoid
the use of epithets already in existence in potential
congeners (when creating new names or describing
new species). Entwisle & Weston (2005) concluded
that ease of recognition of genera is not always
achievable, and that certain “biological criteria” (e.g.,
hybridization, special morphological characters)
should not be emphasized when designating genera.
In the new classification presented here, we attempt
to follow these guidelines.
We favor the recognition of 17 genera in core
Maxillariinae (clades A through Q in Whitten et al.
2007) that correspond to well supported clades (fig.
1). This inevitably requires the resurrection of some
generic names previously placed in synonymy with
Maxillaria, and the creation of several new genera.
These new generic realignments are presented here.
The alternative, to lump the minor genera in
Maxillaria, would be less nomenclaturally disruptive,
but would substantially add to the already baffling
morphological diversity of that genus and make it
even less morphologically diagnosable. Further arguments in support of our new classification and morphological characters for each genus are presented in
Whitten et al. (2007). Keys and detailed descriptions
to all the genera in subtribe Maxillariinae will be published elsewhere.
BLANCO et al. - Generic realignments in Maxillariinae
FIGURE 1. Summary of generic relationships within the
core Maxillariinae, simplified from figure 8 of Whitten et
al. (2007). Based on a maximum parsimony analysis of
combined nrITS, matK+trnK, atpB-rbcL spacer, and
rpoC1 DNA sequence data. Values above branches are
bootstrap percentages. Numbers next to generic names
indicate their known or estimated number of species.
The realization that many large and/or charismatic
genera are polyphyletic or need to be lumped with
others based on phylogenetic principles has recently
spurred support from part of the botanical community for the acceptance of paraphyletic taxa (most of
the debate has taken place in recent issues of
Taxon). However, we are convinced that the designation of monophyletic genera is a far better, less
subjective option. Discussion of our arguments for
this lies outside of the realm of the present contribution; we refer readers to Pfeil & Crisp (2005), who
provide a lucid argument in favor of phylogenetic
classifications and against the use of paraphyletic
supraspecific taxa.
We also effect the nomenclatural transfers
required by our new classification. Following Cribb
517
et al.’s (1985) recommendation, we only transfer
species for which we are confident of their systematic position either because we have sampled them for
our molecular phylogeny (Whitten et al. 2007) or
because of their clear morphological affinity with
species included in our analyses.
We refrain from transferring names that might be
synonyms (even if not yet currently regarded as
such), and treat taxonomically complicated groups
with caution. We do not transfer a few names that
appear in our phylogenies (Whitten et al. 2007)
because now we consider them as synonyms. Many
species of Camaridium and Ornithidium already
have combinations in those genera, although in
some cases their epithets are different than those in
Maxillaria. For each species transferred, homotypic
synonyms with a different epithet are listed after the
basionym when these have been in recent use. Some
heterotypic synonyms are given after the abbreviation “syn.”.
To further clarify the circumscription of
Camaridium, Maxillaria sensu stricto and
Ornithidium, we list the names of their constituent
species. These lists mostly follow Govaerts et al.
(2005) (synonyms not listed) except for cases in
which we disagree on synonymization. We have not
been able to locate extant type material for many
names of Schlechter and other authors published
without illustrations; these are not included here,
awaiting further research. It is possible that many of
these will prove to be synonyms with other better
known species.
Most species for which adequate descriptions or
type material is available are easy to assign to each
genus. However, we prefer to await molecular data
to confirm the systematic position of a few species
with unusual morphology (e.g., Maxillaria grobyoides Garay & Dunst., M. muscoides J. T. Atwood,
M. poifolia Schltr.).
BRASILIORCHIS
Brasiliorchis R. Singer, S. Koehler & Carnevali,
Novon 17: 94. 2007.
Type species: Maxillaria picta Hook., = Brasiliorchis
picta (Hook.) R. Singer, S. Koehler & Carnevali.
Maxillaria sect. Aggregatae Pfitz. in Engl. &
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LANKESTERIANA
Prantl, Nat. Pflanzenfam. 2(6): 187. 1889, pro parte
(excl. type).
Maxillaria sect. Repentes Pfitz. in Engl. & Prantl,
Nat. Pflanzenfam. 2(6): 187. 1889.
Plants of Brasiliorchis have aggregated or distant,
sulcate, bifoliate pseudobulbs subtended by non-foliaceous sheaths. Several inflorescences are produced
simultaneously from the base of the most recent
pseudobulb, and the floral bract is almost always
shorter than the pedicel and ovary. The campanulate
flowers are food deceptive and do not produce any
rewards. The column foot can be short or long. The
sepals lack fibers, and in most species they have dark
spots, usually more intense on the external surface.
The labellum is always markedly three-lobed. The
capsules have apical dehiscence2.
This well supported clade has been informally
known as the “Maxillaria picta alliance”; most species
are restricted to Brazil. It was recently described in
detail and given generic status by Singer et al. (2007).
Three names need to be commented upon:
Brasiliorchis picta (Hook.) R. Singer, S. Koehler &
Carnevali, Novon 17: 97. 2007.
Maxillaria acutipetala Hook., Bot. Mag. 69: t.
3966. 1842, syn. nov.
Maxillaria acutipetala was described from a plant
cultivated at Kew, supposedly collected in Central
America by Barclay during the HMS Sulphur expedition. The type specimen and the published painting
clearly correspond to the variable Brasiliorchis picta,
a taxon reliably documented only from Brazil and
northeastern Argentina. The information associated
with the plant cultivated at Kew must have been erroneous or confused, and it was most likely not collected by Barclay (the Sulphur explored only the Pacific
side of the New World); it was probably collected in
Brazil by someone else.
Brasiliorchis barbosae (Loefgr.) R. Singer, S.
Koehler & Carnevali, Novon 17: 96. 2007.
2
Fruits of Maxillariinae always dehisce through six longitudinal seams, but the resulting valves can either remain
united both at the base and the tip, or they can separate
apically. These two modes of dehiscence are termed “lateral” vs. “apical” (Blanco et al. 2006, Whitten et al. 2007).
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Basionym: Maxillaria barbosae Loefgr., Arch.
Jard. Bot. Rio de Janeiro 2: 60. 1918; ‘barbozae’.
Löfgren dedicated the species to João Barbosa
Rodrigues but spelled the epithet as “barbozae”, which
is clearly a typographical error and not an intentional
latinization of the Brazilian orchidologist’s last name.
Both spellings have been used over time, and Singer et
al. (2007) used “barbozae” during the generic transfer
to Brasiliorchis. However, the epithet should be corrected to “barbosae” under article 60 of the Code
(McNeill et al. 2006; also see article 45.2).
Brasiliorchis schunkiana (Campacci & Kautsky) R.
Singer, S. Koehler & Carnevali, Novon 17: 97. 2007.
Basionym: Maxillaria schunkiana Campacci &
Kautsky, Orquidário 7: 136. 1993; ‘schunkeana’.
The adjectival epithet honors Vital Schunk; thus, it
must be spelled “schunkiana”, not “schunkeana”
(Rec. 60C.1, McNeill et al. 2006). Singer et al.
(2007) and other authors have used the latter spelling,
which is an incorrect orthographical variant.
Species of Brasiliorchis include B. barbosae
(Loefgr.) R. Singer, S. Koehler & Carnevali, B.
chrysantha (Barb. Rodr.) R. Singer, S. Koehler &
Carnevali, B. consanguinea (Klotzsch) R. Singer, S.
Koehler & Carnevali, B. gracilis (G. Lodd.) R.
Singer, S. Koehler & Carnevali, B. heismanniana
(Barb. Rodr.) R. Singer, S. Koehler & Carnevali, B.
kautskyi (Pabst) R. Singer, S. Koehler & Carnevali,
B. marginata (Lindl.) R. Singer, S. Koehler &
Carnevali, B. phoenicanthera (Barb. Rodr.) R. Singer,
S. Koehler & Carnevali, B. picta (Hook.) R. Singer,
S. Koehler & Carnevali, B. polyantha (Barb. Rodr.)
R. Singer, S. Koehler & Carnevali, B. porphyrostele
(Rchb.f.) R. Singer, S. Koehler & Carnevali, B.
schunkiana (Campacci & Kautsky) R. Singer, S.
Koehler & Carnevali, and B. ubatubana (Hoehne) R.
Singer, S. Koehler & Carnevali (Singer et al. 2007).
CAMARIDIUM
Camaridium Lindl., Bot. Reg. 10: sub t. 844. 1824.
Type species: Camaridium ochroleucum Lindl.
Adamanthus Szlach., Richardiana 7: 30. 2007, pro
parte (incl. type).
Maxillaria sect. Camaridium (Lindl.) Christenson,
Proc. 16th World Orchid Conf. 282. 2002.
BLANCO et al. - Generic realignments in Maxillariinae
Maxillaria sect. Cucullatae Christenson, Proc. 16th
World Orchid Conf. 283. 2002.
Maxillaria sect. Pseudomaxillaria (Hoehne)
Christenson, Proc. 16th World Orchid Conf. 285.
2002.
Pseudomaxillaria Hoehne, Arq. Bot. Estado Sao
Paulo n.s., f.m. 2: 71. 1947.
Psittacoglossum LaLlave & Lex., Nov. Veg. Desc.
fasc. 2. (Orch. Opusc.): 29. 1825.
Sauvetrea Szlach., Richardiana 7: 28. 2007, pro
parte (excl. type).
Sepalosaccus Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19: 244. 1923.
Species of Camaridium are variable in growth
habit; most have pseudobulbs separated by rhizome
segments of variable length. Some species are cespitose, and others lack pseudobulbs completely and
have monopodial shoots. A few species have dimorphic growth (juvenile sympodial shoots with tightly
spaced pseudobulbs, and monopodial mature shoots
without pseudobulbs). In almost all species, the floral
bract is longer than the pedicel and ovary, and overlaps with the base of the dorsal sepal (this feature is
useful in separating Camaridium from Maxillariella
and Ornithidium). The column foot can be short or
long. The sepals and petals lack fiber bundles and
have a sparkling appearance. Most species appear to
have deceptive flowers, but some produce nectar. The
pendent fruits have apical dehiscence.
As circumscribed here, Camaridium has ca. 80
species distributed throughout the Neotropics, with
the highest diversity in Central America. The
“Maxillaria neglecta complex” (= Pseudomaxillaria)
is nested within Camaridium; the Mesoamerican
species were revised by Atwood (1993). The
“Maxillaria cucullata complex”, which is sister to the
rest of Camaridium, is a taxonomically difficult
group in dire need of revision.
519
Camaridium amabile (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria amabilis J. T. Atwood,
Lindleyana 9: 239-241. 1994.
Camaridium ampliflorum (C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria ampliflora C. Schweinf.,
Bot. Mus. Leafl. 8: 188. 1940. Camaridium grandiflorum Ames, Proc. Biol. Soc. Wash. 34: 149-150.
1921, nom. illeg. [non (Lindl.) Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 9: 165. 1921].
Camaridium anceps (Rchb.f.) M. A. Blanco, comb.
nov.
Basionym: Ornithidium anceps Rchb.f., Beitr. Orch.K. Centr. Amer. 75-76. 1866. Maxillaria pseudoneglecta J. T. Atwood, Lindleyana 8: 30-31. 1993.
Camaridium atratum (Lex.) M. A. Blanco, comb.
nov.
Basionym: Psittacoglossum atratum Lex., Nov.
Veg. Descr. 2: 30. 1825. Maxillaria lexarzana Soto
Arenas & F. Chiang, Orquídea (Méx.) n.s., 12: 238239. 1992.
Camaridium aurantiacum (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium aurantiacum Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 19: 241. 1923.
Syn.: Maxillaria lankesteri Ames, Sched. Orch. 7:
11-12. 1924. Maxillaria jugata Garay,
Orquideología 4: 159. 1969, syn. nov.
Camaridium bomboizense (Dodson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria bomboizensis Dodson,
Orquideología 19: 59. 1994.
Camaridium alfaroi (Ames & C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria alfaroi Ames & C.
Schweinf., Sched. Orch. 10: 83-84. 1930.
Camaridium brevilabium (Ames & Correll) M. A.
Blanco, comb. nov.
Basionym: Maxillaria brevilabia Ames & Correll,
Bot. Mus. Leafl. 11: 15, fig. 1. 1943. Ornithidium
alfaroi Ames & C. Schweinf., Sched. Orch. 10: 98.
1930. Non Camaridium alfaroi (Ames & C.
Schweinf.) M. A. Blanco.
Camaridium allenii (L. O. Williams) M. A. Blanco,
comb. nov.
Basionym: Maxillaria allenii L. O. Williams, Ann.
Missouri Bot. Gard. 27: 282-283, t. 35. 1940.
Camaridium burgeri (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria burgeri J. T. Atwood,
Lindleyana 9: 233-236. 1994.
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LANKESTERIANA
Camaridium campanulatum (C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria campanulata C. Schweinf.,
Bot. Mus. Leafl. 5: 94-95. 1938.
Camaridium cedralense (J. T. Atwood & Mora-Ret.)
M. A. Blanco, comb. nov.
Basionym: Maxillaria cedralensis J. T. Atwood &
Mora-Ret., Selbyana 18: 31-32. 1997.
Camaridium cucullatum (Lindl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria cucullata Lindl., Edwards’s
Bot. Reg. 26: t. 12. 1840.
Camaridium densum (Lindl.) M. A. Blanco, comb.
nov.
Basionym: Maxillaria densa Lindl., Edwards’s Bot.
Reg. 21: t. 1804. 1835.
Camaridium falcatum (Ames & Correll) M. A.
Blanco, comb. nov.
Basionym: Maxillaria falcata Ames & Correll, Bot.
Mus. Leafl. 11: 15. 1943. Ornithidium costaricense
Schltr., Repert. Spec. Nov. Regni Veg. 8: 456.
1910. Non Camaridium costaricense Schltr.
(Repert. Spec. Nov. Regni Veg. 3: 249. 1907).
Orquídea (Mexico) n.s., 12: 252. 1992.
Camaridium horichii (Senghas) M. A. Blanco, comb.
nov.
Basionym: Maxillaria horichii Senghas, Orchidee
(Hamburg) 28: 13. 1977.
Camaridium inauditum (Rchb.f.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria inaudita Rchb.f., Beitr.
Orch.-K. Centr. Am. 76. 1866.
Camaridium insolitum (Dressler) M. A. Blanco,
comb. nov.
Basionym: Maxillaria insolita Dressler,
Orquideología 14: 204. 1981.
Camaridium lankesteri (Ames) M. A. Blanco, comb. nov.
Basionym: Ornithidium lankesteri Ames, Sched.
Orch. 4: 52-53. 1923. Maxillaria quadrata Ames &
Correll, Bot. Mus. Leafl. 6:16. 1943.
Camaridium longicolumna (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria longicolumna J. T. Atwood,
Selbyana 22: 132. 2001.
The epithet longicolumna is a noun in apposition,
and therefore retains its feminine gender.
Camaridium fragrans (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria fragrans J. T. Atwood,
Selbyana 22: 131. 2001.
Camaridium lutheri (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria lutheri J. T. Atwood,
Selbyana 19: 257. 1998.
Camaridium gomezianum (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria gomeziana J. T. Atwood,
Lindleyana 11: 202-204. 1996.
Camaridium meleagris (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria meleagris Lindl., Edwards’s
Bot. Reg. 30: misc. 3. 1844.
Camaridium grisebachianum (Nir & Dod) M. A.
Blanco, comb. nov.
Basionym: Maxillaria grisebachiana Nir & Dod,
Orchid. Antill.: 243. 2000.
Camaridium haberi (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria haberi J. T. Atwood,
Selbyana 16: 245. 1995.
Camaridium hagsaterianum (Soto Arenas) M. A.
Blanco, comb. nov.
Basionym: Maxillaria hagsateriana Soto Arenas,
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Camaridium micranthum M. A. Blanco, nom. nov.
Scaphyglottis parviflora Poepp. & Endl., Nov. Gen.
ac Sp. 1: 58, t. 97. 1835. Maxillaria parviflora
(Poepp. & Endl.) Garay, Bot. Mus. Leafl. 21: 258.
1967. Non Camaridium parviflorum Fawc. (Symb.
Antill. 1: 472. 1910).
Camaridium microphyton (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria microphyton Schltr., Repert.
Spec. Nov. Regni Veg. 8: 457. 1910.
Camaridium mombachoense (A. H. Heller ex J. T.
Atwood) M. A. Blanco, comb. nov.
BLANCO et al. - Generic realignments in Maxillariinae
Basionym: Maxillaria mombachoensis A. H. Heller
ex J. T. Atwood, Selbyana 5: 302. 1981.
Camaridium monteverdense (J. T. Atwood & G.
Barboza) M. A. Blanco, comb. nov.
Basionym: Maxillaria monteverdensis J. T. Atwood
& G. Barboza, Lindleyana 9: 241-242. 1994.
Camaridium neglectum (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium neglectum Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 242. 1923.
Camaridium obscurum (Linden & Rchb.f.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria obscura Linden & Rchb.f.,
Beitr. Orch.-K. Centr. Amer. 31-32, t. 6. 1866.
Camaridium oestlundianum (L. O. Williams) M. A.
Blanco, comb. nov.
Basionym: Maxillaria oestlundiana L. O. Williams,
Amer. Orchid Soc. Bull. 11: 133. 1942.
Camaridium paleatum (Rchb.f.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium paleatum Rchb.f., Linnaea
41: 36. 1877.
Camaridium praestans (Rchb.f.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria praestans Rchb.f., Gard.
Chron., n.s. 23: 566. 1885.
Camaridium pygmaeum M. A. Blanco, nom. nov.
Ornithidium wercklei Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19: 60. 1923. Maxillaria wercklei
(Schltr.) L. O. Williams, Ann. Missouri Bot. Gard.
27: 284. 1923. Non Camaridium wercklei Schltr.
(Repert. Spec. Nov. Regni Veg. Beih. 19: 58-59.
1923).
521
Camaridium scalariforme (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria scalariformis J. T. Atwood,
Selbyana 19: 257. 1998.
Camaridium sigmoideum (C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Ornithidium sigmoideum C. Schweinf.,
Bot. Mus. Leafl. 4: 121-122. 1937.
Camaridium soconuscanum (Breedlove & D. Mally)
M. A. Blanco, comb. nov.
Basionym: Maxillaria soconuscana Breedlove &
D. Mally, Amer. Orchid Soc. Bull. 58: 1234.
1989.
Camaridium standleyi M. A. Blanco, nom. nov.
Maxillaria parvilabia Ames & C. Schweinf.,
Sched. Orch. 8: 62-64. 1925, nom. illeg. (non
Rolfe, Orchid Rev. 26: 232. 1918).
Camaridium stenophyllum (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium stenophyllum Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 19: 59.
1923. Maxillaria concavilabia Ames &
Correll, Bot. Mus. Leafl. 11: 15, fig. 2. 1953.
Non Maxillaria stenophylla Rchb.f. (Bonplandia
2: 17. 1854), nec Maxillaria stenophylla F.
Lehm. & Kraenzl. (Bot. Jahrb. Syst. 26: 481.
1899).
Camaridium strumatum (Endres & Rchb.f.) M. A.
Blanco, comb. nov.
Basionym: Ornithidium strumatum Endres &
Rchb.f., Gard. Chron. 2: 772. 1874.
Camaridium suaveolens (Barringer) M. A. Blanco,
comb. nov.
Basionym: Maxillaria suaveolens Barringer,
Brittonia 37: 44-46. 1985.
Camaridium ramonense (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium ramonense Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 243-244. 1923.
Maxillaria flava Ames, Hubbard & C. Schweinf.,
Bot. Mus. Leafl. 3: 41. 1934.
Camaridium synsepalum (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria synsepala J. T. Atwood,
Selbyana 19: 260. 1998.
Camaridium rhombeum (Lindl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria rhombea Lindl., Edwards’s
Bot. Reg. 26: t. 12. 1840.
Camaridium tigrinum (C. Schweinf.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria tigrina C. Schweinf., Amer.
Orchid Soc. Bull. 37: 409-410. 1968.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
LANKESTERIANA
522
Camaridium tricarinatum (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria tricarinata J. T. Atwood,
Selbyana 19: 260. 1998.
Camaridium tuberculare (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria tubercularis J. T. Atwood,
Lindleyana 9: 229-231. 1994.
Camaridium tutae (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria tutae J. T. Atwood, Selbyana
19: 262. 1998.
Camaridium vaginale (Rchb.f.) M. A. Blanco, comb.
nov.
Basionym: Maxillaria vaginalis Rchb.f., Beitr.
Orch.-K. Centr. Amer. 77. 1866.
Camaridium valerioi (Ames & C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria valerioi Ames & C.
Schweinf., Sched. Orch. 10: 96-97. 1930.
Camaridium vittariifolium (L. O. Williams) M. A.
Blanco, comb. nov.
Basionym: Maxillaria vittariifolia L. O. Williams,
Ceiba 4: 38. 1953.
Other members of Camaridium include C. adolphi
Schltr., C. amparoanum Schltr. (Maxillaria serrulata Ames & Correll), C. biolleyi (Schltr.) Schltr., C.
bracteatum (Schltr.) Schltr., C. bradeorum Schltr.,
C. brenesii Schltr. (M. trilobata Ames & C.
Schweinf.), C. costaricense Schltr. (M. tonduzii
(Schltr.) Ames & Correll), C. ctenostachys (Rchb.f.)
Schltr., C. dendrobioides Schltr., C. dichotomum
Schltr., C. hoehnei Pabst (M. imbricata Barb.
Rodr.), C. imbricatum Schltr. (M. schlechteriana J.
T. Atwood), C. latifolium Schltr. (M. planicola C.
Schweinf.), C. minus Schltr. [M. minor (Schltr.) L.
O. Williams], C. nutantiflorum Schltr. (M.
umbratilis L. O. Williams), C. ochroleucum Lindl.
(M. camaridii Rchb.f.), and C. pulchrum Schltr.
CHRISTENSONELLA
´
Christensonella Szlach., Mytnik, Górniak & Smiszek,
Polish Bot. J. 51: 57. 2006.
Type species: Maxillaria paulistana Hoehne, =
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Christensonella subulata (Lindl.) Szlach., Mytnik,
´
Górniak & Smiszek
[= Maxillaria subulata Lindl.],
syn. nov.
Maxillaria sect. Urceolatae Christenson, Proc. 16th
World Orchid Conf. 286-287. 2002.
Species of Christensonella are either epiphytes or
lithophytes. Plants of this genus are often very
small, compared to other members of Maxillariinae.
In most species, the roots show characteristic constrictions. Epiphytic species tend to grow pendent
and have distant pseudobulbs, while the lithophytes
are erect and more cespitose. The pseudobulbs are
often ridged. Each pseudobulb bears from one to
four apical leaves, and has several non-foliaceous
subtending sheaths. The leaves can be flat and
conduplicate to subulate or hemiterete. The flowers
are usually yellow to dark red and have a shiny, dry
callus; perianth fibers are present. Christensonella
uncata and C. squamata have prominent stelidia
projecting downward from the apex of the clinandrium, which support a long, tegular stipe. The
fusiform fruits have apical dehiscence. In C. nardoides, the endocarpic trichomes are extruded as a
sausage-like mass which carries the seeds outside of
the fruit (Blanco et al. 2006).
This distinctive group of mostly South American
species has been informally known as the “Maxillaria
madida alliance” (Pabst & Dungs 1977). A detailed
revision of Christensonella is in progress (S. Koehler,
unpublished manuscript). Szlachetko et al. (2006)
transferred most members of Maxillaria section
Urceolatae to Christensonella, and here we transfer a
few more. Whitten et al. (2007) wrongfully cited M.
nardoides Kraenzl. as the type of Christensonella, but
that species is the type of Maxillaria section
Urceolatae.
Christensonella cepula (Rchb.f.) S. Koehler, comb. nov.
Basionym: Maxillaria cepula Rchb.f., Bonplandia
3: 216. 1855.
Christensonella neowiedii (Rchb.f.) S. Koehler,
comb. nov.
Basionym: Maxillaria neowiedii Rchb. f., Linnaea
41: 29. 1877.
Christensonella pacholskii (Christenson) S. Koehler,
comb. nov.
BLANCO et al. - Generic realignments in Maxillariinae
Basionym: Maxillaria pacholskii Christenson,
Orchid Rev. 111: 288. 2003.
Christensonella squamata (Barb. Rodr.) Carnevali,
comb. nov.
Basionym: Maxillaria squamata Barb. Rodr., Gen.
Sp. Orchid. 1: 118. 1877.
Other members of Christensonella which should be
recognized as discrete species (S. Koehler, unpublished manuscript) include: C. acicularis (Herb. ex
´
Lindl.) Szlach., Mytnik, Górniak & Smiszek,
C.
echinophyta (Barb. Rodr.) Szlach., Mytnik, Górniak
´
& Smiszek,
C. ferdinandiana (Barb. Rodr.) Szlach.,
´
Mytnik, Górniak & Smiszek,
C. nardoides (Kraenzl.)
´
Szlach., Mytnik, Górniak & Smiszek,
C. pumila
´
(Hook.) Szlach., Mytnik, Górniak & Smiszek, C. sub´
ulata (Lindl.) Szlach., Mytnik, Górniak & Smiszek,
C. uncata (Lindl.) Szlach., Mytnik, Górniak &
´
Smiszek,
and C. vernicosa (Barb. Rodr.) Szlach.,
´
Mytnik, Górniak & Smiszek.
Other names transferred
by Szlachetko et al. (2006) are regarded as synonyms
(S. Koehler, unpublished manuscript).
523
and the labellum. Species in subgenus Caulescentes
are even more aberrant, and have dwarf shoots with
polystichous phyllotaxis. The capsules have apical
dehiscence.
Because of its unusual morphology,
Cryptocentrum has been consistently recognized as
a separate, well defined genus. Carnevali (2001)
provided a synopsis. It is significant that the three
subgenera (Caulescentes, Cryptocentrum and
Pseudobulbosa; Carnevali, 2001) form strongly
supported monophyletic groups (Whitten et al.
2007). Because we now consider Anthosiphon as
part of Cryptocentrum, the erection of a new subgenus is necessary:
Cryptocentrum Benth. subgenus Anthosiphon
(Schltr.) Carnevali, comb. et stat. nov.
Anthosiphon Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 7: 182. 1920.
Cryptocentrum sect. Anthosiphon (Schltr.) Hawkes,
Orchid J. 2: 379. 1953.
TYPE: Cryptocentrum roseans (Schltr.) A. D.
Hawkes (= Anthosiphon roseans Schltr.)
CRYPTOCENTRUM
CYRTIDIORCHIS
Cryptocentrum Benth., J. Linn. Soc. Bot. 18: 325.
1880.
Type species: Cryptocentrum jamesonii Benth., = C.
lehmannii (Rchb.f.) Garay [= Aeranthes lehmanii
Rchb.f.].
Anthosiphon Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 7: 182. 1920.
Cryptocentrum sect. Anthosiphon (Schltr.) Hawkes,
Orchid J. 2: 379. 1953.
Cryptocentrum subgen. Caulescentes Senghas, in
Schlechter Orchideen, ed. 3, I/B(29): 1798. 1994.
Cryptocentrum subgen. Pseudobulbosa Carnevali,
Harvard Pap. Bot. 5: 470. 2001.
Cryptocentrum subgen. Cryptocentrum Benth.
sensu Carnevali, Harvard Pap. Bot. 5: 468. 2001.
Pittierella Schltr., Repert. Spec. Nov. Regni Veg.
3(31-32): 80. 1906.
Cryptocentrum is unusual among core
Maxillariinae because most species have monopodial (often congested) shoots, long, wiry inflorescences, and greenish, star shaped flowers with a
nectariferous spur formed by the bases of the sepals
Cyrtidiorchis Rauschert, Taxon 31: 560. 1982.
Type species (designated by Ortiz, Orquídeas de
Colombia ed. 2: 70, 1995): Chrysocycnis rhomboglossa F. Lehm. & Kraenzl., = Cyrtidiorchis
rhomboglossa (F. Lehm. & Kraenzl.) Rauschert.
Cyrtidium Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 27: 178. 1924, nom. illeg. (non Vainio, Acta
Soc. Fauna Flora Fenn. 49: 227, 262 1921).
Plants of Cyrtidiorchis have dimorphic growth
with sympodial, pseudobulb bearing juvenile shoots,
and monopodial, branched adult shoots.
Inflorescences are supra-axillary and are only produced by the adult shoots. The flowers have spreading perianth segments that lack fibers, a tomentose,
insectiform labellum, and a strongly arched column;
they are probably sexually deceptive. The capsules
have lateral dehiscence.
Garay (1969) presented a taxonomic revision of
Cyrtidiorchis (as Cyrtidium). This small Andean
genus (five species) is well characterized and no new
combinations are necessary.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
LANKESTERIANA
524
HETEROTAXIS
Heterotaxis Lindl., Bot. Reg. 12: t. 1028. 1826.
Type species: Heterotaxis crassifolia Lindl., = H.
sessilis (Sw.) F. Barros Barros [=Epidendrum sessile Sw.].
Dicrypta Lindl., Gen. Sp. Orchid. Pl. 44. 1830.
Marsupiaria Hoehne, Arq. Bot. Estado Sao Paulo
n.s., f.m. 2: 69. 1947.
Maxillaria subgen. Heterotaxis (Lindl.) Brieger,
An. Soc. Bot. Brasil 1972: 94. 1972.
Maxillaria sect. Heterotaxis (Lindl.) Brieger, Trab.
Congr. Nac. Bot. (Rio de Janeiro) 26: 242-244,
1977.
Maxillaria sect. Iridifolieae Pfitz., Nat.
Pflanzenfam. 2(6): 187. 1889.
Pentulops Raf., Fl. Tellur. 4: 42. 1836.
Most species of Heterotaxis have sympodial growth
with laterally compressed, oblong, unifoliate
pseudobulbs subtended by several foliaceous sheaths.
Two species (H. equitans and H. valenzuelana) have
ensiform leaves and pseudo-monopodial shoots that
lack pseudobulbs. In all cases, the shoots are aggregate. The fleshy, yellowish, campanulate flowers
have perianth fibers and a very short column foot.
The labellum produces a pad of very short, glandular
trichomes that likely constitute a reward for pollinators. The capsules have lateral dehiscence.
Ojeda et al. (2005) provided the most recent
detailed account of Heterotaxis. They described H.
fritzii Ojeda & Carnevali but accidentally provided
erroneous information for the type, which is amended
here (corrections underlined):
Heterotaxis fritzii Ojeda & Carnevali, Novon 15:
574-577. 2005.
TYPE: Purchased from Orquídeas del Valle, Cali
[Colombia]. Flowered in cultivation in Gainesville,
Florida, U.S.A., 25 Jan 2004, M. W. Whitten 2672
(holotype: FLAS).
Species that belong in Heterotaxis are: H. brasiliensis
(Brieger & Illg) F. Barros, H. discolor (Lodd. ex Lindl.)
Ojeda & Carnevali, H. equitans (Schltr.) Ojeda &
Carnevali, H. fritzii Ojeda & Carnevali, H. maleolens
(Schltr.) Ojeda & Carnevali, H. microiridifolia (D. E.
Benn. & Christenson) Ojeda & Carnevali, H. santanae
(Carnevali & I. Ramírez) Ojeda & Carnevali, H. schulLANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
tesii Ojeda & G. A. Romero, H. sessilis (Sw.) F. Barros
(Maxillaria crassifolia Lindl.), H. superflua (Rchb.f.) F.
Barros, H. valenzuelana (A. Rich.) Ojeda & Carnevali,
H. villosa (Barb. Rodr.) F. Barros, and H. violaceopunctata (Rchb.f.) F. Barros (Ojeda et al. 2005).
INTI
Inti M. A. Blanco, gen. nov.
Type species: Maxillaria chartacifolia Ames & C.
Schweinf., = Inti chartacifolia (Ames & C.
Schweinf.) M. A. Blanco.
Maxillaria sect. Polyphyllae Christenson, Proc. 16th
World Orchid Conf. 284-285. 2002.
Plantae cespitosae, epiphyticae, epseudobulbosae,
surculis congestis flabellatis. Folia disticha plurima
longa angusta. Inflorescentiae axillares in foliis
infimis. Flores foetidi, perianthio fibrarum destituto.
Fructus capsulares longi dehiscentiis lateralibus.
Plants of Inti are easily recognized by their aggregate, congested shoots devoid of pseudobulbs, with
many (>10) distichous, long leaves arranged like a fan.
The yellow or maroon flowers lack perianth fibers and
have a fetid odor. The column foot is virtually non
existent, and the labellum has a pad of glandular trichomes similar to those of Heterotaxis. The capsules
are long and narrow, and have lateral dehiscence.
ETYMOLOGY: Named after Inti, the sun god of the Inca
culture. The long and narrow leaves radiating from a congested shoot are reminiscent of the rays of a rising sun.
Having no botanical tradition, we give this generic name a
feminine gender (article 62.3, McNeill et al. 2006).
This small group was treated as the “Maxillaria
bicallosa clade” in Whitten et al. (2007).
Inti bicallosa (Rchb.f.) M. A. Blanco, comb. nov.
Basionym: Zygopetalum bicallosum Rchb.f., Otia
Bot. Hamb. 1: 9. 1878. Syn.: Ornithidium
dolichophyllum Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 9: 106. 1921 (= Maxillaria caespitosa C.
Schweinf.), syn. nov.
Inti chartacifolia (Ames & C. Schweinf.) M. A.
Blanco, comb. nov.
Basionym: Maxillaria chartacifolia Ames & C.
Schweinf., Sched. Orch. 10: 92. 1930. Syn.:
BLANCO et al. - Generic realignments in Maxillariinae
Trigonidium equitans Garay, Svensk Bot. Tidskr.
47: 288, fig. 15. 1953, syn. nov.
525
MAPINGUARI
Mapinguari desvauxianus (Rchb.f.) Carnevali & R.
Singer, comb. nov.
Basionym: Maxillaria desvauxiana Rchb.f.,
Bonplandia (Hannover) 3: 67. 1854.
Mapinguari Carnevali & R. Singer, gen. nov.
Type species: Maxillaria longipetiolata Ames & C.
Schweinf., = Mapinguari longipetiolatus (Ames &
C. Schweinf.) Carnevali & R. Singer.
Mapinguari foldatsianus (Carnevali & I. Ramírez)
Carnevali & R. Singer, comb. nov.
Basionym: Maxillaria foldatsiana Carnevali & I.
Ramírez, Ann. Missouri Bot. Gard. 76: 376. 1989.
Plantae plerumque sylvicolae, epiphyticae vel
rupicolae, pseudobulbis laevis, semper unifoliatis,
aggregatis. Radices laeves, ferrugineae, brunneae vel
albae. Flores erecti, fragrantes, rigidi et sine secretiones; pedicello brevissimo rigidoque. Pollinarium
cum tegula et viscidio semilunare.
Mapinguari longipetiolatus (Ames & C. Schweinf.)
Carnevali & R. Singer, comb. nov.
Basionym: Maxillaria longipetiolata Ames & C.
Schweinf., Sched. Orch. 8: 61-62. 1925.
Mapinguari is a small (four species), primarily
Guayanan and Amazonian genus. Mapinguari
desvauxianus reaches the Brazilian states of São Paulo
and Rio de Janeiro. Two of the species are primarily
terrestrial in sandy soils or lithophytic on sandstone.
The pseudobulbs are aggregate, unifoliate and smooth
to slightly sulcate. The conduplicate leaves normally
display a well developed petiole. The very short inflorescences are produced from the base of the most
recent pseudobulb. The flowers are erect (i.e., the
labellum is held in a vertical or near-vertical position),
brown or maroon in coloration, have a very short column foot, lack any secretions or trichomes, and have
tough perianth fibers. We have not been able to determine the mode of dehiscence of the capsules.
Maxillaria Ruiz & Pav., Fl. Peruv. Prodr. 116, t. 25.
1794.
Type species (designated by Brieger & Hunt, Taxon
18: 601-603. 1969; and by Garay, Harvard Pap.
Bot. 11: 51-52. 1997): Maxillaria platypetala Ruiz
& Pav.
Adamanthus Szlach., Richardiana 7: 30. 2007, pro
parte (excl. type).
Dendrobium sect. Maxillaria (Ruiz & Pav.) Pers.,
Syn. Pl. (Persoon) 2: 523. 1807.
Maxillaria sect. Aggregatae Pfitz., Nat.
Pflanzenfam. 2(6): 187. 1889, pro parte (incl. type).
Maxillaria sect. Amazonicae Christenson, Proc. 16th
World Orchid Conf. 282. 2002.
Maxillaria sect. Arachnites Christenson, Proc. 16th
World Orchid Conf. 282. 2002.
Maxillaria sect. Axilliflorae Lindl., Gen. Sp.
Orchid. Pl. 142. 1833, pro parte.
Maxillaria sect. Maxillaria Ruiz & Pav. sensu
Christenson, Proc. 16th World Orchid Conf. 284.
2002.
Maxillaria sect. Multiflorae Christenson, Proc. 16th
World Orchid Conf. 284. 2002.
Maxillaria subgen. Aggregatae (Pfitz.) Brieger,
Trab. Congr. Nac. Bot. (Rio de Janeiro) 26: 244.
1977.
Menadena Raf., Fl. Tellur. 2: 98. 1836.
Sauvetrea Szlach., Richardiana 7: 28. 2007, pro
parte (excl. type).
ETYMOLOGY: Named after the Mapinguarí, a legendary
and elusive creature of Brazilian-Amazonian mythology, in allusion to the brownish, cryptic flowers. Having
no botanical tradition, we treat this generic name as
masculine (article 62.3, McNeill et al. 2007).
This small but distinctive group was first recognized
by Carnevali & Ramírez (1989; as the “Maxillaria
auyantepuiensis complex”) and was treated as the
“Maxillaria desvauxiana clade” in Whitten et al.
(2007). Full synonymy for each species can be found
in Carnevali & Ramírez-Morillo (2003).
Mapinguari auyantepuiensis (Foldats) Carnevali &
R. Singer, comb. nov.
Basionym: Maxillaria auyantepuiensis Foldats,
Bol. Soc. Venez. Ci. Nat. 22: 269. 1961.
MAXILLARIA
Plants of this group almost always have pseudobulbs (with the exception of some species in the M.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
526
LANKESTERIANA
exaltata alliance) and are often cespitose. The
pseudobulbs almost invariably are unifoliate, smooth
and laterally compressed, with subtending sheaths
that are either foliaceous or not. The abscission layer
of the apical leaf is often projected above the
pseudobulb in a persistent stalk (phyllopodium). The
inflorescences always emerge from the base of the
youngest pseudobulbs (from the leaf axils near the
top of the stem in the M. exaltata alliance). The floral
bract can be shorter or longer than the pedicel and
ovary. The flowers have a prominent column foot,
and abundant perianth fibers. None of the species
produce nectar, but many produce pseudopollen in
the form of moniliform, pluricellular trichomes on the
labellum surface (a character not seen in any other
genera). The capsules have lateral dehiscence.
Almost half of the species traditionally treated in
Maxillaria sensu lato will remain in Maxillaria sensu
stricto, as here circumscribed. Maxillaria sections
Amazonicae, Maxillaria, and Multiflorae form well
supported clades (Whitten et al. 2007), but the other
sections are polyphyletic. Increased sampling of taxa
and gene regions within Maxillaria sensu stricto is
needed to support a revised infrageneric classification. Two names included in the analyses of Whitten
et al. (2007) need to be commented upon:
Maxillaria candida Lodd. ex Lindl., Edwards’s Bot.
Reg. 27 (Misc.): 28. 1841.
Syn.: Maxillaria modesta Brade, Orquídea (Rio de
Janeiro) 6: 18. 1943, nom. illeg. (non Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 28: 93. 1924),
Maxillaria modestiflora Pabst, Bradea 2: 319. 1979.
syn. nov.
Whitten et al. (2007) reported a specimen of
Maxillaria candida Lodd. ex Lindl. (Koehler 0335,
ESA) nested in the Mormolyca clade. Upon examination of the type of M. candida at Kew, we realized that
Koehler 0335 is Mormolyca cf. acutifolia (Lindl.) M.
A. Blanco. Whitten et al. (2007) also reported a specimen of Maxillaria modesta Schltr. nested in Maxillaria
sensu stricto (Koehler 0351, UEC). This specimen is,
however, the true Maxillaria candida, which was originally identified simply as “M. modesta”. The illegitimate name Maxillaria modesta Brade (non Schltr.) and
its later legitimate name M. modestiflora Pabst are previously undetected synonyms of M. candida.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Even in this exclusive circumscription, Maxillaria
remains by far the largest genus in subtribe
Maxillariinae. Species that belong in Maxillaria sensu
stricto include: M. acostae Schltr., M. aequiloba
Schltr., M. albata Lindl., M. albiflora Ames & C.
Schweinf., M. amazonica Schltr., M. anatomorum
Rchb.f., M. angustisegmenta Ames & C. Schweinf., M.
angustissima Ames, T. Hubb. & C. Schweinf., M.
arachnites Rchb.f., M. arachnitiflora Ames & C.
Schweinf., M. argyrophylla Poepp. & Endl., M. attenuata Ames & C. Schweinf., M. atwoodiana Pupulin, M.
augustae-victoriae F. Lehm. & Kraenzl., M. aurorae
D. E. Benn. & Christenson, M. azulensis D. E. Benn.
& Christenson, M. batemanii Poepp. & Endl., M. bennettii Christenson, M. bocazensis D. E. Benn. &
Christenson, M. bolivarensis C. Schweinf., M. brachybulbon Schltr., M. bradei Schltr. ex Hoehne, M. breviscapa Poepp. & Endl., M. buchtienii Schltr., M. burtonii D. E. Benn. & Christenson, M. calantha Schltr.,
M. candida Lodd. ex Lindl., M. carolii Christenson, M.
chionantha J. T. Atwood, M. chlorantha Lindl., M.
christensonii D. E. Benn., M. colemanii Carnevali &
Fritz, M. colorata Rchb.f., M. confusa Ames & C.
Schweinf., M. connellii Rolfe, M. crocea Poepp. &
Endl., M. cryptobulbon Carnevali & J. T. Atwood, M.
curvicolumna M. A. Blanco & Neubig, M. cuzcoensis
C. Schweinf., M. dalessandroi Dodson, M. dichroma
Rolfe, M. dillonii D. E. Benn. & Christenson, M. x
dunstervillei Carnevali & I. Ramírez, M. eburnea
Lindl., M. ecuadorensis Schltr., M. edwardsii D. E.
Benn. & Christenson, M. elegantula Rolfe, M. embreei
Dodson, M. endresii Rchb.f., M. exaltata (Kraenzl.) C.
Schweinf., M. fletcheriana Rolfe, M. floribunda Lindl.,
M. formosa Carnevali & G. A. Romero, M. fractiflexa
Rchb.f., M. frechettei D. E. Benn. & Christenson, M.
fucata Rchb.f., M. fuerstenbergiana Schltr., M. galantha J. T. Atwood & Carnevali, M. gentryi Dodson, M.
gorbatschowii R. Vásquez, Dodson & Ibisch, M. grandiflora (Kunth) Lindl., M. grandimentum C. Schweinf.,
M. grandis Rchb.f., M. granditenuis D. E. Benn. &
Christenson, M. grayi Dodson, M. guadalupensis
Cogn., M. hastulata Lindl., M. hennisiana Schltr., M.
hillsii Dodson, M. hirsutilabia D. E. Benn. &
Christenson, M. huanucoensis D. E. Benn. &
Christenson, M. huebschii Rchb.f., M. irrorata Rchb.f.,
M. jostii Dodson, M. jucunda F. Lehm. & Kraenzl., M.
kegelii Rchb.f., M. klugii C. Schweinf., M. langlassei
BLANCO et al. - Generic realignments in Maxillariinae
Schltr., M. leforii D. E. Benn. & Christenson, M.
lehmannii Rchb.f., M. lepidota Lindl., M. leucaimata
Barb. Rodr., M. lilliputiana D. E. Benn. & Christenson,
M. lindleyana Schltr. (M. crocea Lindl.), M. linearis
Ames & C. Schweinf., M. litensis Dodson, M. longiloba (Ames & C. Schweinf.) J. T. Atwood, M. longipes
Lindl., M. longissima Lindl., M. loretoensis C.
Schweinf., M. lueri Dodson, M. luteoalba Lindl., M.
macrura Rchb.f., M. margretiae R. Vásquez, M. marmoliana Dodson, M. mejiae Carnevali & G. A.
Romero, M. melina Lindl., M. merana Dodson, M.
meridensis Lindl., M. microtricha Schltr., M. milenae
V. P. Castro & Chiron, M. molitor Rchb.f., M. monantha Barb. Rodr., M. multiflora Barb. Rodr., M.
mungoschraderi R. Vásquez & Ibisch, M. nanegalensis Rchb.f., M. neophylla Rchb.f., M. niesseniae
Christenson, M. nigrescens Lindl., M. nuriensis
Carnevali & I. Ramírez, M. nutans Lindl., M.
ochroleuca Lodd. ex Lindl., M. pachyneura F. Lehm.
& Kraenzl., M. pannieri Foldats, M. parkeri Hook., M.
parvibulbosa C. Schweinf., M. patens Schltr., M. pauciflora Barb. Rodr., M. pentura Lindl., M. perryae
Dodson, M. platyloba Schltr., M. platypetala Ruiz &
Pav., M. plicata Schltr., M. porrecta Lindl., M. portillae Christenson, M. powellii Schltr., M. pseudoreichenheimiana Dodson, M. pterocarpa Barb. Rodr., M.
pulla Linden & Rchb.f., M. pyhalae D. E. Benn. &
Christenson, M. quelchii Rolfe, M. ramonensis Schltr.,
M. reichenheimiana Endres & Rchb.f., M. ringens
Rchb.f., M. rodriguesii Cogn., M. rodrigueziana J. T.
Atwood & Mora-Ret., M. rotundilabia C. Schweinf.,
M. rubioi Dodson, M. sanderiana Rchb.f. ex Sander,
M. setigera Lindl., M. silvana Campacci, M. simacoana Schltr., M. simplicilabia C. Schweinf., M. speciosa Rchb.f., M. spiritu-sanctensis Pabst, M. splendens Poepp. & Endl., M. striata Rolfe, M. tenuis C.
Schweinf., M. thurstoniorum Dodson, M. tiaraensis
Carnevali & G. A. Romero, M. tonsbergii Christenson,
M. trilobulata D. E. Benn. & Christenson, M. triloris
E. Morren, M. tristis Schltr., M. tuerosii D. E. Benn. &
Christenson, M. turkeliae Christenson, M. valleculata
D. E. Benn. & Christenson, M. venusta Linden &
Rchb.f., M. whittenii Dodson, M. williamsii Dodson,
M. winaywaynaensis D. E. Benn. & Dodson, M. wojii
Christenson, M. woytkowskii C. Schweinf., M. xylobiiflora Schltr., M. yanganensis Dodson, and M.
yauaperyensis Barb. Rodr.
527
MAXILLARIELLA
Maxillariella M. A. Blanco & Carnevali, gen. nov.
Type species: Maxillaria diuturna Ames & C.
Schweinf., = Maxillariella diuturna (Ames & C.
Schweinf.) M. A. Blanco & Carnevali.
Adamanthus Szlach., Richardiana 7: 30. 2007, pro
parte (excl. type).
Maxillaria sect. Ebulbes Pfitz., Nat. Pflanzenfam.
2(6): 187. 1889.
Maxillaria sect. Erectae Pfitz., Nat. Pflanzenfam.
2(6): 187. 1889.
Plantae epiphyticae, pro parte maxima rhizomatibus longis, pseudobulbosae vel epseudoblbosae;
pseudobulbi 1-2 foliis apicalibus. Flores solitarii in
axillis foliorum vel bractearum. Flores plerumque
parvi campanulati. Labella parviloba vel elobulata,
callis nitidis. Fructus capsulares dehiscentiis lateralibus.
Species of Maxillariella are variable in terms of
growth habit; a few species are subcespitose, but
most have pseudobulbs separated by medium to long
rhizome segments. The ovoid pseudobulbs are either
uni- or bifoliate. Several species with long rhizomes
have foliaceous bracts covering the segments between
pseudobulbs. Others have pseudobulbs reduced or
even absent, and these species show a clear gradient
from sympodial to monopodial growth. The most
derived members of this genus are the species formerly treated as Maxillaria section Ebulbes (the
“Maxillaria graminifolia suballiance”, Atwood
2003), which have thin, wiry, monopodial stems completely devoid of pseudobulbs, and narrow, acute
leaves. Invariably, only one flower is produced from
each leaf or bract axil (e.g., the inflorescences are not
fasciculate nor produced sequentially), and the floral
bract is shorter than the pedicel and ovary. The column foot is very short, and the labellum is simple or
obscurely three-lobed and has a glossy callus. The
flowers seem to be food deceptive in most cases. The
capsules have lateral dehiscence.
This group was treated as the “Maxillaria variabilis
clade” by Whitten et al. (2007). Maxillaria sect.
Ebulbes (the “Maxillaria graminifolia suballiance”)
was revised by Atwood (2003). Camaridium dendrobioides, the type of Adamanthus, is morphologically
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
528
LANKESTERIANA
convergent with members of that suballiance, but it is
firmly nested in Camaridium (Whitten et al. 2007).
The floral bract in C. dendrobioides is longer than the
pedicel and ovary.
Maxillariella cobanensis (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria cobanensis Schltr., Repert.
Spec. Nov. Regni Veg. 10: 295. 1912.
ETYMOLOGY: The name Maxillariella is a diminutive
of Maxillaria, in reference to the often much smaller
flowers of Maxillariella.
Maxillariella costaricensis (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria costaricensis Schltr., Rep.
Spec. Nov. Regni Veg. Beih. 19: 232-233. 1923.
Maxillariella acervata (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria acervata Rchb.f., Bonplandia
(Hannover) 3: 217. 1855.
Maxillariella alba (Hook.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Dendrobium album Hook.f., Exot. Fl. t.
142. 1825.
Maxillariella anceps (Ames & C. Schweinf.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Maxillaria anceps Ames & C.
Schweinf., Sched. Orch. 10: 84. 1930.
Maxillariella appendiculoides (C. Schweinf.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Maxillaria appendiculoides C.
Schweinf., Bot. Mus. Leafl. 4: 119-121. 1937.
Maxillariella arbuscula (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium arbuscula Lindl., Pl.
Hartw. 153. 1845.
Maxillariella brevifolia (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium brevifolium Lindl., Benth.
Pl. Hartw. 154. 1845.
Maxillariella caespitifica (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria caespitifica Rchb.f., Linnaea
41: 73. 1877.
Maxillariella cassapensis (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria cassapensis Rchb.f., Ann.
Bot. Syst. 6: 539. 1863.
Maxillariella caucana (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria caucana Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 7: 167. 1920.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Maxillariella curtipes (Hook.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria curtipes Hook., Icon. Pl. 4: t.
384. 1841
Maxillariella densifolia (Poepp. & Endl.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Dicrypta densifolia Poepp. & Endl.,
Nov. Gen. Sp. Pl. 1: 39. 1836.
Maxillariella diuturna (Ames & C. Schweinf.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Maxillaria diuturna Ames & C.
Schweinf., Sched. Orch. 8: 58. 1925.
Maxillariella elatior (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Dicrypta elatior Rchb.f., Linnaea 18:
403. 1844.
Maxillariella estradae (Dodson) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria estradae Dodson, Icon. Pl.
Trop. 1: t. 152. 1980.
Maxillariella funicaulis (C. Schweinf.) M. A. Blanco
& Carnevali, comb. nov.
Basionym: Maxillaria funicaulis C. Schweinf., Bot.
Mus. Leafl. 11: 273. 1945.
Maxillariella graminifolia (Kunth) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Isochilus graminifolius Kunth, Nov.
Gen. Sp. 1: 340, t. 78. 1816.
Maxillariella guareimensis (Rchb.f) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria guareimensis Rchb.f.,
Bonplandia (Hannover) 2: 16. 1854.
Maxillariella houtteana (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
BLANCO et al. - Generic realignments in Maxillariinae
Basionym: Maxillaria houtteana Rchb.f., Hamb.
Gartenz. 14: 212. 1858.
Maxillariella infausta (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria infausta Rchb.f., Bonplandia
(Hannover) 3: 216. 1855.
Maxillariella lawrenceana (Rolfe) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium lawrenceanum Rolfe,
Bull. Misc. Inform. Kew 1894: 185. 1894.
Maxillariella linearifolia (Ames & C. Schweinf.) M.
A. Blanco & Carnevali, comb. nov.
Basionym: Maxillaria linearifolia Ames & C.
Schweinf., Sched. Orch. 10: 95-96. 1930.
Maxillariella longibracteata (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium longibracteatum Lindl.,
Benth. Pl. Hartw. 154. 1845.
Maxillariella luteorubra (F. Lehm. & Kraenzl.) M.
A. Blanco & Carnevali, comb. nov.
Basionym: Ornithidium luteorubrum F. Lehm. &
Kraenzl., Bot. Jahrb. Syst. 26: 486. 1899.
Maxillaria cuencana Garay, Cand. J. Bot. 34: 257.
1956. Non Camaridium luteo-rubrum Lindl.
(Orchid. Linden. 22. 1846).
Maxillariella mexicana (J. T. Atwood) M. A. Blanco
& Carnevali, comb. nov.
Basionym: Maxillaria mexicana J. T. Atwood,
Selbyana 24: 35-36. 2003.
Maxillariella microdendron (Schltr.) M. A. Blanco
& Carnevali, comb. nov.
Basionym: Maxillaria microdendron Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 8: 94. 1921.
Maxillariella nitidula (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria nitidula Rchb.f., Linnaea 41:
29. 1876.
In Whitten et al. (2007) we stated that this species
probably belongs in Camaridium. Upon re-examination of herbarium material, we are now convinced it
belongs in Maxillariella.
Maxillariella oreocharis (Schltr.) M. A. Blanco &
529
Carnevali, comb. nov.
Basionym: Maxillaria oreocharis Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 17: 69. 1922.
Maxillariella pardalina (Garay) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria pardalina Garay, Bot. Mus.
Leafl. 26: 28. 1978. Maxillaria pantherina Rchb.f.
1855, Bonplandia (Hannover) 3: 239. 1855, nom.
illeg. (non Hoffmanns., Verz. Orchid. ed. 3: 71. 1844).
Maxillariella pastensis (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria pastensis Rchb.f.,
Bonplandia (Hannover) 3: 239. 1855.
Maxillariella ponerantha (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria ponerantha Rchb.f.,
Bonplandia (Hannover) 2: 17. 1854.
Maxillariella procurrens (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria procurrens Lindl., Ann.
Mag. Nat. Hist. 15: 383. 1845.
Maxillariella prolifera (Sw.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Epidendrum proliferum Sw., Prodr.:
124. 1788. Maxillaria swartziana C. D. Adams,
Amer. Orchid Soc. Bull. 35: 998. 1966.
Maxillariella purpurata (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium purpuratum Lindl.,
Orchid. Linden.: 22. 1846.
Maxillariella robusta (Barb. Rodr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium robustum Barb. Rodr.,
Gen. Spec. Orchid. 2: 210. 1881. Maxillaria johannis Pabst, Sellowia 10: 165. 1959.
Maxillariella sanguinea (Rolfe) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria sanguinea Rolfe, Bull. Misc.
Inform. Kew 1895: 8.
Maxillariella spilotantha (Rchb.f.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria spilotantha Rchb.f.,
Bonplandia (Hannover) 2: 17. 1854.
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530
LANKESTERIANA
Maxillariella stenophylla (Rchb.f. ) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria stenophylla Rchb.f.,
Bonplandia (Hannover) 2: 17. 1854. Non F. C.
Lehm. ex Kraenzl. (Bot. Jahrb. Syst. 26: 481. 1899).
Maxillariella stictantha (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria stictantha Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 8: 97. 1921.
Maxillariella tenuifolia (Lindl.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria tenuifolia Lindl., Edwards’s
Bot. Reg. 23: sub. t. 1986. 1837.
Maxillariella tuerckheimii (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium tuerckheimii Schltr.,
Repert. Spec. Nov. Regni Veg. 10: 296. 1912.
Maxillaria nagelii L. O. Williams ex Correll,
Lloydia 10: 212. 1947. Non Maxillaria tuerckheimii Schltr. (Repert. Spec. Nov. Regni Veg. 10:
295. 1912).
Maxillariella variabilis (Bateman ex Lindl.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Maxillaria variabilis Bateman ex Lindl.,
Edwards’s Bot. Reg. 23: sub. t. 1986. 1837.
Maxillariella vinosa (Rolfe) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Camaridium vinosum Rolfe, Bull. Misc.
Inform. Kew 1922: 25-26. 1922. Non Camaridium
vinosum Schltr. (Repert. Spec. Nov. Regni Veg.
Beih. 19: 240. 1923), nec Maxillaria vinosa
Senghas (Schlechter Orchideen, ed. 3, I/B(28):
1751. 1993, nom. illeg.).
Maxillariella vulcanica (F. Lehm. & Kraenzl.) M. A.
Blanco & Carnevali, comb. nov.
Basionym: Maxillaria vulcanica F. Lehm. &
Kraenzl., Bot. Jahrb. Syst. 26: 484. 1899.
Maxillariella xanthorhoda (Schltr.) M. A. Blanco &
Carnevali, comb. nov.
Basionym: Maxillaria xanthorhoda Schltr., Notizbl.
Bot. Gart. Berlin-Dahlem 7: 279. 1918.
Maxillariella x yucatanensis (Carnevali & R.
Jiménez) M. A. Blanco & Carnevali, comb. nov.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Basionym: Maxillaria x yucatanensis Carnevali &
R. Jiménez, Harvard Pap. Bot. 5: 428. 2001.
MORMOLYCA
Mormolyca Fenzl, Denkschr. Kaiserl. Akad. Wiss.,
Wien. Math.-Naturwiss. Kl. 1: 253. 1850.
Type species: Mormolyca lineolata Fenzl, = M. ringens (Lindl.) Gentil [=Trigonidium ringens Lindl.].
Chrysocycnis Linden & Rchb.f., Bonplandia
(Hannover) 2: 280. 1854.
Cyrtoglottis Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 7: 181. 1920.
Maxillaria sect. Rufescens Christenson, Proc. 16th
World Orchid Conf. 285-286. 2002.
In this expanded circumscription, Mormolyca can
be distinguished by its unifoliate pseudobulbs having a minutely verrucose texture and subtended by
non-foliaceous sheaths, the inflorescences arising
from the axils of rhizome bracts well behind the
most recent pseudobulb, perianth segments that lack
fibers and open widely, and a conspicuously clavate,
arcuate column. Mormolyca polyphylla (which is
sister to the rest of the genus) is atypical because its
long, narrow pseudobulbs have up to three apical
leaves and two subtending foliaceous sheaths, and
the inflorescences are produced from the base of the
most recent pseudobulb. The rhizome segments and
the floral peduncles can be short or long, depending
on the species. The column foot is always very
short. The labellum of most species (those formerly
treated in Maxillaria section Rufescens) have a pad
of short, glandular trichomes on the callus, but the
labellum of the other species is tomentose and insectiform. The capsules have apical dehiscence.
Chrysocycnis and Maxillaria section Rufescens
are firmly nested within Mormolyca, and thus their
constituent species need to be transferred.
Mormolyca sensu stricto was revised by Garay &
Wirth (1959), and Chrysocycnis was revised by
Sweet (1971). Species of Maxillaria section
Rufescens, informally known as the “Maxillaria
rufescens complex”, constitute a taxonomically difficult group; the Mesoamerican species were
revised by Carnevali et al. (2001), but many more
are found in South America. There probably are a
number of cryptic species, difficult to tell apart as
BLANCO et al. - Generic realignments in Maxillariinae
herbarium specimens, but distinguishable in life
especially by their floral fragrances (Christenson
2002a, 2002b, and personal observation of the
authors).
531
Mormolyca rufescens (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria rufescens Lindl., Edwards’s
Bot. Reg. 22: t. 1848. 1836.
Mormolyca acutifolia (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria acutifolia Lindl., Edwards’s
Bot. Reg. 25: misc. 92. 1839.
Mormolyca sanantonioensis (Christenson) M. A.
Blanco, comb. nov.
Basionym:
Maxillaria
sanantonioensis
Christenson, Orchids 71: 128. 2002.
Mormolyca aureoglobula (Christenson) M. A.
Blanco, comb. nov.
Basionym: Maxillaria aureoglobula Christenson,
Orchids 71: 125-126. 2002.
Mormolyca schlimii (Linden & Rchb.f.) M. A.
Blanco, comb. nov.
Basionym: Chrysocycnis schlimii Linden &
Rchb.f., Bonplandia (Hannover) 2: 280. 1854.
Mormolyca chacoensis (Dodson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria chacoensis Dodson, Icon. Pl.
Trop., II, 6: t. 531. 1989.
Mormolyca sotoana (Carnevali & Gómez-Juárez) M.
A. Blanco, comb. nov.
Basionym: Maxillaria sotoana Carnevali &
Gómez-Juárez, Brittonia 53: 461-463.
Mormolyca cleistogama (Brieger & Illg) M. A.
Blanco, comb. nov.
Basionym: Maxillaria cleistogama Brieger & Illg,
Trab. Congr. Nac. Bot. 26: 247. 1977.
Mormolyca dressleriana (Carnevali & J. T. Atwood)
M. A. Blanco, comb. nov.
Basionym: Maxillaria dressleriana Carnevali & J.
T. Atwood, Lindleyana 11: 29-31. 1996.
Mormolyca hedwigiae (Hamer & Dodson) M. A.
Blanco, comb. nov.
Basionym: Maxillaria hedwigiae Hamer &
Dodson, Icon. Pl. Trop. 8: t. 800. 1982.
Mormolyca lehmanii (Rolfe) M. A. Blanco, comb. nov.
Basionym: Chrysocycnis lehmanii Rolfe, Bull.
Misc. Inform. Kew 1918: 235.
Mormolyca moralesii (Carnevali & J. T. Atwood) M.
A. Blanco, comb. nov.
Basionym: Maxillaria moralesii Carnevali & J. T.
Atwood, Lindleyana 11: 31-32. 1996.
Mormolyca pudica (Carnevali & Tapia-Muñoz) M.
A. Blanco, comb. nov.
Basionym: Maxillaria pudica Carnevali & TapiaMuñoz, Brittonia 53: 463-465. 2001. Maxillaria
rufescens var. minor Fawcett & Rendle, J. Bot. 48:
108. 1910.
Mormolyca richii (Dodson) M. A. Blanco, comb. nov.
Basionym:
Maxillaria
richii
Dodson,
Orquideología 19: 81. 1994.
Mormolyca suarezorum (Dodson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria suarezorum Dodson, Icon.
Pl. Trop., II, 6: t. 547. 1989.
Mormolyca tenuibulba (Christenson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria tenuibulba Christenson,
Orchid Rev. 109: 41. 2001.
The other members of the genus constitute the paraphyletic Mormolyca sensu stricto: M. aurorae D. E.
Benn. & Christenson, M. gracilipes (Schltr.) Garay &
M. Wirth, M. peruviana C. Schweinf., M. polyphylla
Garay & M. Wirth, M. ringens (Lindl.) Gentil, and M.
schweinfurthiana Garay & M. Wirth. The obscure
name M. galeata (Scheidw.) Garay & M. Wirth is not
a true Mormolyca, but probably represents a species
in the Camaridium cucullatum alliance.
NITIDOBULBON
Nitidobulbon Ojeda, Carnevali & G. A. Romero,
ined.
Species of this group have oblong, shiny, smooth
pseudobulbs with several (4-6) subtending foliaceous
sheaths, and one or two apical leaves. The flowers are
campanulate and the rigid perianth segments have
abundant fibers. The column foot is very short, and
the labellum is frequently reflexed at the tip. The
labellar callus is ligulate and secretes an abundant,
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LANKESTERIANA
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resinous substance, but lacks glandular trichomes.
The capsules have lateral dehiscence.
This small group was referred to as the “Maxillaria
nasuta clade” in Whitten et al. (2007), and will be
elevated to generic rank by Ojeda et al. (in press). It
comprises the following three species currently in
Maxillaria: M. cymbidioides Dodson, J. T. Atwood &
Carnevali, M. nasuta Rchb.f., and M. proboscidea
Rchb.f.
ORNITHIDIUM
Ornithidium Salisb. ex R. Br., Hort. Kew. ed. 2, 5:
210. 1813.
Type species: Epidendrum coccineum Jacq., =
Ornithidium coccineum (Jacq.) Salisb. ex R. Br.
Adamanthus Szlach., Richardiana 7: 30. 2007, pro
parte (excl. type).
Laricorchis Szlach., Richardiana 7: 27. 2007.
Maxillaria sect. Ornithidium (Salisb. ex R. Br.)
Christenson, Richardiana 2: 52. 2002.
Maxillaria sect. Reflexae Christenson, Proc. 16th
World Orchid Conf. 285. 2002.
Maxillaria sect. Siagonanthus (Poepp. & Endl.)
Christenson, Proc. 16th World Orchid Conf. 286.
2002.
Neo-urbania Fawc. & Rendle, J. Bot. 47: 125.
1909.
Siagonanthus Poepp. & Endl., Nov. Gen. Sp. Pl.
(Poeppig & Endlicher) 1: 40. 1836.
Species of Ornithidium can be either sympodial
(cespitose to long-rhizomatous) or monopodial; a few
species have dimorphic growth (sympodial juvenile
shoots and monopodial adult shoots). The stems and
leaves of most species have an olive green coloration,
which is persistent upon drying. When present, the
ovoid pseudobulbs have a shiny, minutely cracked
texture reminiscent of old varnish. The thick roots
have a characteristic orangish coloration.
Inflorescences are usually fascicled, and the pedicel
and ovary invariably are much longer than the floral
bract. The flowers are usually small, fleshy, campanulate or more often subglobose, and often produce
nectar, and the perianth lacks fibers. Many species
have yellow, orange, or red flowers. Capsules have
apical dehiscence.
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Ornithidium adendrobium (Rchb.f.) M. A. Blanco &
Ojeda, comb. nov.
Basionym: Ponera adendrobium Rchb.f., Flora 48:
278. 1964.
Ornithidium affine (Poepp. & Endl.) M. A. Blanco &
Ojeda, comb. nov.
Basionym: Scaphyglottis affinis Poepp. & Endl.,
Nov. Gen. Sp. Pl. 1: 59, t. 99A. 1836.
Ornithidium cachacoense (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria cachacoensis J. T. Atwood,
Selbyana 24: 30-31. 2003.
Ornithidium canarense (J. T. Atwood) M. A. Blanco
& Ojeda, comb. nov.
Basionym: Maxillaria canarensis J. T. Atwood,
Selbyana 24: 31-32. 2003.
Ornithidium condorense (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria condorensis J. T. Atwood,
Selbyana 24: 32-33. 2003.
Ornithidium fimbriatilobum (Carnevali & G. A.
Romero) M. A. Blanco & Ojeda, comb. nov.
Basionym: Maxillaria fimbriatiloba Carnevali & G.
A. Romero, Orchids Venezuela, ed. 2, 3: 11381139. 2000.
Ornithidium gualaquizense (Dodson) M. A. Blanco
& Ojeda, comb. nov.
Basionym: Maxillaria gualaquizensis Dodson,
Orquideología 19: 69. 1994.
Ornithidium haemathodes (Ruiz & Pav.) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Fernandezia haemathodes Ruiz & Pav.,
Syst. Veg. Fl. Peruv. Chil. 1: 240. 1798.
Ornithidium lasallei (Foldats) M. A. Blanco &
Ojeda, comb. nov.
Basionym: Maxillaria lasallei Foldats, Contr. Ocas.
Mus. Hist. Nat. Colegio de la Salle 3: 2. 1961.
Ornithidium machinazense (D. E. Benn. &
Christenson) M. A. Blanco, comb. nov.
Basionym: Maxillaria machinazensis D. E. Benn.
& Christenson, Lindleyana 13: 71. 1998.
Ornithidium maldonadoense (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
BLANCO et al. - Generic realignments in Maxillariinae
Basionym: Maxillaria maldonadoensis J. T.
Atwood, Selbyana 24: 33-35. 2003.
Ornithidium minutiflorum (D. E. Benn. &
Christenson) M. A. Blanco & Ojeda, comb. nov.
Basionym: Maxillaria minutiflora D. E. Benn. &
Christenson, Icon. Orchid. Peruv.: t. 700. 2001.
Ornithidium nicaraguense (Hamer & Garay) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Neo-urbania nicaraguensis Hamer &
Garay, Icon. Pl. Trop. 13: t. 1238. 1985.
Ornithidium oxapampense (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria oxapampensis J. T. Atwood,
Selbyana 24: 36-37. 2003.
Ornithidium patellum (J. T. Atwood) M. A. Blanco
& Ojeda, comb. nov.
Basionym: Maxillaria patella J. T. Atwood,
Selbyana 24: 37-39. 2003.
Ornithidium patulum (C. Schweinf.) M. A. Blanco
& Ojeda, comb. nov.
Basionym: Maxillaria patula C. Schweinf.,
Fieldiana, Bot. 28: 197. 1951.
Ornithidium pseudonubigenum (J. T. Atwood) M.
A. Blanco & Ojeda, comb. nov.
Basionym: Maxillaria pseudonubigena J. T.
Atwood, Selbyana 24: 39-41. 2003.
Ornithidium pustulosum (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria pustulosa J. T. Atwood,
Selbyana 24: 41-43. 2003.
Ornithidium rauhii (D. E. Benn. & Christenson) M.
A. Blanco & Ojeda, comb. nov.
Basionym: Maxillaria rauhii D. E. Benn. &
Christenson, J. Orchideenfr. 12: 34. 2005.
Ornithidium repens (L. O. Williams) M. A. Blanco
& Ojeda, comb. nov.
Basionym: Maxillaria repens L. O. Williams,
Amer. Orchid Soc. Bull. 10: 273. 1942.
Ornithidium rigidum (Barb. Rodr.) M. A. Blanco &
Ojeda, comb. nov.
Basionym: Maxillaria rigida Barb. Rodr., Gen.
Spec. Orchid. 2: 206. 1881.
533
Ornithidium scandens (D. E. Benn. & Christenson)
M. A. Blanco & Ojeda, comb. nov.
Basionym: Maxillaria scandens D. E. Benn. &
Christenson, Icon. Orchid. Peruv.: t. 707. 2001.
Ornithidium scullianum (J. T. Atwood) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria sculliana J. T. Atwood,
Selbyana 24: 1-3. 2003.
Ornithidium sillarense (Dodson & Vásquez) M. A.
Blanco & Ojeda, comb. nov.
Basionym: Maxillaria sillarensis Dodson &
Vásquez, Icon. Pl. Trop., II, 3: pl. 261. 1989.
Ornithidium simplex (J. T. Atwood) M. A. Blanco &
Ojeda, comb. nov.
Basionym: Maxillaria simplex J. T. Atwood,
Selbyana 24: 43. 2003.
Other members of Ornithidium include O. aggregatum Rchb.f., O. aureum Poepp. & Endl., O. breve
Schltr. (Maxillaria palmensis Dodson), O. chrysocycnoides Schltr., O. coccineum (Jacq.) Salisb. ex R.
Br., O. conduplicatum Ames & C. Schweinf., O.
croceorubens Rchb.f., O. distichum Lindl., O. fulgens Rchb.f., O. giganteum Lindl., O. hystrionicum
Rchb.f., O. jamesonii Rchb.f., O. mapiriense
Kraenzl., O. miniatum Lindl., O. multicaule (Poepp.
& Endl.) Rchb.f., O. niveum Lindl., O. nubigenum
Rchb.f., O. pendens (Pabst) Senghas, O. pendulum
(Poepp. & Endl.) Cogn., O. pittieri Ames, O. purpureolabium (D. E. Benn. & Christenson) Senghas,
O. quitense Rchb.f., O. ruberrimum (Lindl.) Rchb.f.,
O. sanaense (D. E. Benn. & Christenson) Senghas,
O. semiscabrum Lindl., O. serrulatum Lindl. (M.
alticola C. Schweinf.), O. sophronitis Rchb.f., O.
squarrosum Schltr., and O. tonsoniae (Soto Arenas)
Senghas.
PITYPHYLLUM
Pityphyllum Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 7: 162. 1920.
Type species (designated by Sweet, 1972: 205):
Pityphyllum antioquiense Schltr.
Plants of Pityphyllum are minute, pendulous epiphytes,
with long rhizome segments between the tiny pseudobulbs. The pseudobulbs are completely covered and fused
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LANKESTERIANA
534
to a subtending sheath (tunica), and can have from one to
20 apical leaves, depending on the species. There is a
pair of ligular projections at the apex of each leaf sheath,
next to the abscission layer of the blade. The diminutive
flowers are borne from the axils of rhizome bracts
between the pseudobulbs. The column foot is virtually
non existent, and the capsules have apical dehiscence.
This small and distinctive Andean genus was
revised by Sweet (1972) and recently expanded by
Whitten et al. (2006) to accommodate P. huancabambae (Kraenzl.) Whitten and P. saragurense (Dodson)
Whitten, previously placed in Maxillaria. The other
species are P. amesianum Schltr., P. antioquiense
Schltr., P. hirtzii Dodson, P. laricinum (Kraenzl.)
Schltr., and P. pinoides Sweet.
RHETINANTHA
Rhetinantha M. A. Blanco, gen. nov.
Type species: Maxillaria acuminata Lindl., =
Rhetinantha acuminata (Lindl.) M. A. Blanco.
Sauvetrea Szlach., Richardiana 7: 28. 2007, pro
parte (excl. type).
Plantae epiphyticae, rhizomatibus longis et
pseudobulbis 1-4 foliis apicalibus vel plantae pendulae foliis equitantibus. Flores plerumque virides, nunquam late expansi, partibus perianthii rigidis acuminatis. Labella elobulata, secretionibus resinosis vel
ceraceis. Fructus capsulares dehiscentiis lateralibus.
Plants of Rhetinantha are sub-cespitose to long rhizomatous; the pseudobulbs are often ridged, and have
two to four apical leaves, and usually one or two subtending foliaceous sheaths. The inflorescences often
arise from rhizome bracts a few shoots behind the
most recent pseudobulb. The flowers are campanulate, with rigid, acuminate perianth parts with strong
fibers, and a very short column foot. The labellum
secretes a sticky, resinous substance in most species.
The margins of the clinandrium are conspicuously
ciliate, and the pollinarium stipe is long with involute
margins. A few species have a pair of stelidia projected downward from the apex of the clinandrium. The
capsules have lateral dehiscence. Rhetinantha witsenioides is highly modified and has pendent, monopodial shoots with ensiform, glaucous leaves.
E TYMOLOGY : From the Greek words “rhetinos”
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(resin) and “anthos” (flower), in reference to the
flowers of most species, which secrete resin on the
labellum and occasionally on the petals.
This well defined group was treated as the
“Maxillaria acuminata clade” by Whitten et al.
(2007).
Rhetinantha aciantha (Rchb.f.) M. A. Blanco, comb. nov.
Basionym: Maxillaria aciantha Rchb.f., Bot.
Zeitung (Berlin) 10: 858. 1852.
Rhetinantha acuminata (Lindl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria acuminata Lindl., Pl. Hartw.
155. 1845. Sauvetrea acuminata (Lindl.) Szlach.,
Richardiana 7: 29. 2006.
Rhetinantha cerifera (Barb. Rodr.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria cerifera Barb. Rodr. Gen.
Spec. Orchid. 1: 118. 1877.
Rhetinantha divaricata (Barb. Rodr.) M. A. Blanco,
comb. nov.
Basionym: Ornithidium divaricatum Barb. Rodr.,
Gen. Spec. Orchid. 2: 209. 1881.
Rhetinantha encyclioides (J. T. Atwood & Dodson)
M. A. Blanco, comb. nov.
Basionym: Maxillaria encyclioides J. T. Atwood &
Dodson, Orquideología 20: 268. 1997.
Rhetinantha friedrichsthalii (Rchb.f) M. A. Blanco,
comb. nov.
Basionym: Maxillaria friedrichsthalii Rchb.f., Bot.
Zeitung (Berlin) 10: 858. 1852.
Rhetinantha mariaisabeliae (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria mariaisabeliae J. T.
Atwood, Selbyana 7: 250. 1984.
Rhetinantha monacensis (Kraenzl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria monacensis Kraenzl.,
Repert. Spec. Nov. Regni Veg. 24: 49. 1927.
Rhetinantha neilii (Dodson) M. A. Blanco, comb.
nov.
Basionym: Maxillaria neilii Dodson, Orquideología
19: 79. 1994.
BLANCO et al. - Generic realignments in Maxillariinae
Rhetinantha notylioglossa (Rchb.f.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria notylioglossa Rchb.f.,
Bonplandia (Hannover) 2: 16. 1854.
Rhetinantha ophiodens (J. T. Atwood) M. A.
Blanco, comb. nov.
Basionym: Maxillaria ophiodens J. T. Atwood,
Selbyana 7: 248. 1984.
535
callus, and the midlobe is much longer than the lateral
lobes. There are no secretions produced by the flowers.
The capsules have lateral dehiscence.
Sauvetrea was polyphyletic as originally circum´
scribed by Szlachetko & Smiszek
(2007). We already
indicated the species that do not belong in this clade
(Whitten et al. 2007), and here we transfer some that do.
Sauvetrea bomboizensis (Dodson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria bomboizensis Dodson,
Orquideología 19(3): 59-61. 1994.
Rhetinantha pastorellii (D. E. Benn. & Christenson)
M. A. Blanco, comb. nov.
Basionym: Maxillaria pastorellii D. E. Benn. &
Christenson, Lindleyana 13: 74. 1998. Maxillaria
unguiculata D. E. Benn. & Christenson, Brittonia
47: 197. 1995, nom. illeg. (non Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 7: 173. 1920).
Sauvetrea chicana (Dodson) M. A. Blanco, comb.
nov.
Basionym: Maxillaria chicana Dodson,
Orquideología 19(3): 61. 1994.
Rhetinantha schistostele (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria schistostele Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 303-304. 1923.
Sauvetrea cornuta (C. Schweinf.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria cornuta C. Schweinf., Bot.
Mus. Leafl. 11: 265. 1945.
Rhetinantha scorpioidea (Kraenzl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria scorpioidea Kraenzl.,
Svensk. Vet. Akad. Handl. 46: 71. 1911.
Sauvetrea laevilabris (Lindl.) M. A. Blanco, comb.
nov.
Basionym: Maxillaria laevilabris Lindl., Pl.
Hartw.: 155. 1845. Syn.: Maxillaria piestopus
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
302. 1923, syn. nov.; Maxillaria koehleri Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 9: 103. 1921,
syn. nov.
Rhetinantha witsenioides (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria witsenioides Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 7: 175. 1920.
SAUVETREA
Sauvetrea Szlach., Richardiana 7: 28. 2007.
Type species: Maxillaria alpestris Lindl., = Sauvetrea
alpestris (Lindl.) Szlach.
Maxillaria sect. Trigonae Christenson, Proc. 16th
World Orchid Conf. 286. 2002.
Plants of Sauvetrea are cespitose to moderately longrhizomatous. The unifoliate pseudobulbs are frequently
ancipitous and are subtendend by a pair of non-foliaceous bracts. The inflorescences arise from the base of
the newly emerging pseudobulbs and have strongly
ancipitous bracts. The column foot is short. The flowers have spreading perianth segments without strong
fibers. Both the ovaries and the mature fruits are
strongly trigonous. The labellum has a ligulate, sulcate
Sauvetrea machupicchuensis (Christenson) M. A.
Blanco, comb. nov.
Basionym: Maxillaria machupicchuensis
Christenson, Orchids 71: 718-719. 2002.
Sauvetrea napoensis (Dodson) M. A. Blanco, comb. nov.
Basionym: Maxillaria napoensis Dodson, Icon. Pl.
Trop., II, 6: t. 540. 1989.
Sauvetrea sessilis (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria sessilis Lindl., Pl. Hartw.
155. 1843.
Sauvetrea trigona subsp. amaroensis (D. E. Benn. &
Christenson) M. A. Blanco, comb. nov.
Basionym: Maxillaria trigona subsp. amaroensis
D. E. Benn. & Christenson, Icon. Orchid. Peruv.: t.
708. 2001.
Other species in the genus include S. alpestris
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LANKESTERIANA
536
(Lindl.) Szlach., S. peruviana (C. Schweinf.) Szlach.,
S. trigona (C. Schweinf.) Szlach., S. unicarinata (C.
Schweinf.) Szlach., and S. xantholeuca (Schltr.)
Szlach. However, Sauvetrea needs a thorough taxonomic revision. The strange Maxillaria grobyoides
Garay & Dunst. may belong in Sauvetrea, but we prefer to wait until we have molecular data to confirm or
reject this hypothesis.
TRIGONIDIUM
Trigonidium Lindl., Bot. Reg. 23: t. 1923. 1837.
Type species: Trigonidium obtusum Lindl.
Plants of Trigonidium are either cespitose or longrhizomatous. The ridged pseudobulbs have one to
four apical leaves and are subtended by non-foliaceous bracts. The inflorescences are erect, often long
and wiry, and emerge from the most recent pseudobulbs. The flowers are erect and campanulate and lack
a column foot. The sepals form a cup around the
smaller petals and lip and are reflexed at the middle.
The petals have characteristic shiny thickenings at the
apex. The capsules have lateral dehiscence.
Trigonidium has been consistently recognized as a
genus since its creation. No nomenclatural transfers
are needed. A synoptical treatment of the genus is in
progress (M. A. Blanco, unpublished manuscript).
ACKNOWLEDGEMENTS. Eric A. Christenson (BRIT) provided general discussions on taxonomy and data about the
publication of his sections of Maxillaria. Gustavo Romero
(AMES), Cassio van den Berg (HUEFS) and two anonmous reviewers provided suggestions for improving the
manuscript. Funding was provided by the U.S. National
Science Foundation (grant No. DEB-0234064) to N.H.W.
and W.M.W., a FAPESP postdoctoral grant (01/08958-1)
to R.B.S., a M.Sc. grant from CONACYT to I.O., KLARF
fellowships from the Royal Botanic Garden, Kew, to
M.A.B. (to study type material in European herbaria) and
S.K. (a Prance Fellowship to study systematics of
Christensonella), and a Furniss Foundation Ph.D.
Fellowship from the American Orchid Society to M.A.B.
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LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
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