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LANKESTERIANA 7(3): 515-537. 2007.
GENERIC REALIGNMENTS IN MAXILLARIINAE (ORCHIDACEAE)
MARIO A. BLANCO
1,2
, GERMAN CARNEVALI
3
, W. MARK WHITTEN
4
, RODRIGO B. SINGER
5
,
S
AMANTHA KOEHLER
6
, NORRIS H. WILLIAMS
4
, ISIDRO OJEDA
7
, KURT M. NEUBIG
1
& LORENA ENDARA
1
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)
A
BSTRACT. 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 pre-
sent a new phylogenetic classification for core Maxillariinae that recognizes 17 genera. Necessary realign-
ments 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 sec-
ción
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.
09 Blanco [24] 5/12/07 00:23 Página 1
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 dis-
parate 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, 2002b
1
), 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) pre-
cluded any sound taxonomic decisions based on their
results. More recently, Whitten
et al. (2007) present-
ed a vastly more detailed phylogenetic analysis of
core Maxillariinae (619 individuals representing ca.
354 species; four DNA regions used) that corroborat-
ed Dathe and Dietrich’s preliminary results and
revealed a worst-case scenario: all the currently
accepted minor genera of core Maxillariinae
(
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 prima-
ry 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 rec-
ognizable 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 taxo-
nomic 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 argu-
ments in support of our new classification and mor-
phological characters for each genus are presented in
Whitten
et al. (2007). Keys and detailed descriptions
to all the genera in subtribe Maxillariinae will be pub-
lished elsewhere.
LANKESTERIANA
516
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
1
Christenson (2002a) presented a partial subgeneric classi-
fication 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. 16
th
World
Orchid Conf.”) was delayed substantially. The two publi-
cations appeared in 2006, and the exact date of printing is
not indicated in either one. The Proceedings were pub-
lished 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).
09 Blanco [24] 5/12/07 00:23 Página 2
BLANCO et al. - Generic realignments in Maxillariinae
517
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
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 communi-
ty for the acceptance of paraphyletic taxa (most of
the debate has taken place in recent issues of
Taxon). However, we are convinced that the desig-
nation of monophyletic genera is a far better, less
subjective option. Discussion of our arguments for
this lies outside of the realm of the present contribu-
tion; 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
et al.’s (1985) recommendation, we only transfer
species for which we are confident of their systemat-
ic 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 abbrevia-
tion “
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 groby-
oides
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. &
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.
09 Blanco [24] 5/12/07 00:23 Página 3
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
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-folia-
ceous 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 dehiscence
2
.
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 expedi-
tion. 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 erro-
neous or confused, and it was most likely not collect-
ed 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.
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 cor-
rected 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. 16
th
World Orchid Conf. 282. 2002.
LANKESTERIANA
518
2
Fruits of Maxillariinae always dehisce through six longi-
tudinal 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 “lat-
eral” vs. “apical” (Blanco
et al. 2006, Whitten et al. 2007).
09 Blanco [24] 5/12/07 00:23 Página 4
BLANCO et al. - Generic realignments in Maxillariinae
519
LANKESTERIANA 7(3), diciembre 2007. © Universidad de Costa Rica, 2007.
Maxillaria sect. Cucullatae Christenson, Proc. 16
th
World Orchid Conf. 283. 2002.
Maxillaria sect. Pseudomaxillaria (Hoehne)
Christenson, Proc. 16
th
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 cespi-
tose, and others lack pseudobulbs completely and
have monopodial shoots. A few species have dimor-
phic 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 over-
laps 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.
Camaridium alfaroi (Ames & C. Schweinf.) M. A.
Blanco,
comb. nov.
Basionym: Maxillaria alfaroi Ames & C.
Schweinf., Sched. Orch. 10: 83-84. 1930.
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 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 grandi-
florum
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 pseudone-
glecta
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: 238-
239. 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 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 burgeri (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria burgeri J. T. Atwood,
Lindleyana 9: 233-236. 1994.
09 Blanco [24] 5/12/07 00:23 Página 5
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).
Camaridium fragrans (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria fragrans J. T. Atwood,
Selbyana 22: 131. 2001.
Camaridium gomezianum (J. T. Atwood) M. A.
Blanco,
comb. nov.
Basionym: Maxillaria gomeziana J. T. Atwood,
Lindleyana 11: 202-204. 1996.
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,
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 lutheri (J. T. Atwood) M. A. Blanco,
comb. nov.
Basionym: Maxillaria lutheri J. T. Atwood,
Selbyana 19: 257. 1998.
Camaridium meleagris (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria meleagris Lindl., Edwards’s
Bot. Reg. 30: misc. 3. 1844.
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.
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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).
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 rhombeum (Lindl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria rhombea Lindl., Edwards’s
Bot. Reg. 26: t. 12. 1840.
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 synsepalum (J. T. Atwood) M. A.
Blanco,
comb. nov.
Basionym: Maxillaria synsepala J. T. Atwood,
Selbyana 19: 260. 1998.
Camaridium tigrinum (C. Schweinf.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria tigrina C. Schweinf., Amer.
Orchid Soc. Bull. 37: 409-410. 1968.
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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 serrula-
ta
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, =
Christensonella subulata (Lindl.) Szlach., Mytnik,
Górniak & Smiszek [=
Maxillaria subulata Lindl.],
syn. nov.
Maxillaria sect. Urceolatae Christenson, Proc. 16
th
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 con-
strictions. 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 clinandri-
um, which support a long, tegular stipe. The
fusiform fruits have apical dehiscence. In
C. nar-
doides
, 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.
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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, unpub-
lished 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).
CRYPTOCENTRUM
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 monopodi-
al (often congested) shoots, long, wiry inflores-
cences, and greenish, star shaped flowers with a
nectariferous spur formed by the bases of the sepals
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 sub-
genus 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.)
CYRTIDIORCHIS
Cyrtidiorchis Rauschert, Taxon 31: 560. 1982.
Type species (designated by Ortiz, Orquídeas de
Colombia ed. 2: 70, 1995):
Chrysocycnis rhom-
boglossa
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 pro-
duced by the adult shoots. The flowers have spread-
ing 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.
´
´
´
´
´
´
´
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HETEROTAXIS
Heterotaxis Lindl., Bot. Reg. 12: t. 1028. 1826.
Type species:
Heterotaxis crassifolia Lindl., = H.
sessilis
(Sw.) F. Barros Barros [=Epidendrum ses-
sile
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 aggre-
gate. 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 pollina-
tors. 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. schul-
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. violaceop-
unctata
(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. 16
th
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 aggre-
gate, 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 tri-
chomes similar to those of
Heterotaxis. The capsules
are long and narrow, and have lateral dehiscence.
E
TYMOLOGY: Named after Inti, the sun god of the Inca
culture. The long and narrow leaves radiating from a con-
gested 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.:
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Trigonidium equitans Garay, Svensk Bot. Tidskr.
47: 288, fig. 15. 1953,
syn. nov.
MAPINGUARI
Mapinguari Carnevali & R. Singer, gen. nov.
Type species: Maxillaria longipetiolata Ames & C.
Schweinf., =
Mapinguari longipetiolatus (Ames &
C. Schweinf.) Carnevali & R. Singer.
Plantae plerumque sylvicolae, epiphyticae vel
rupicolae, pseudobulbis laevis, semper unifoliatis,
aggregatis. Radices laeves, ferrugineae, brunneae vel
albae. Flores erecti, fragrantes, rigidi et sine secre-
tiones; pedicello brevissimo rigidoque. Pollinarium
cum tegula et viscidio semilunare.
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 inflo-
rescences 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 col-
umn foot, lack any secretions or trichomes, and have
tough perianth fibers. We have not been able to deter-
mine the mode of dehiscence of the capsules.
E
TYMOLOGY: Named after the Mapinguarí, a legendary
and elusive creature of Brazilian-Amazonian mytholo-
gy, 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.
Mapinguari desvauxianus (Rchb.f.) Carnevali & R.
Singer,
comb. nov.
Basionym: Maxillaria desvauxiana Rchb.f.,
Bonplandia (Hannover) 3: 67. 1854.
Mapinguari foldatsianus (Carnevali & I. Ramírez)
Carnevali & R. Singer,
comb. nov.
Basionym: Maxillaria foldatsiana Carnevali & I.
Ramírez, Ann. Missouri Bot. Gard. 76: 376. 1989.
Mapinguari longipetiolatus (Ames & C. Schweinf.)
Carnevali & R. Singer,
comb. nov.
Basionym: Maxillaria longipetiolata Ames & C.
Schweinf., Sched. Orch. 8: 61-62. 1925.
MAXILLARIA
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. 16
th
World Orchid Conf. 282. 2002.
Maxillaria sect. Arachnites Christenson, Proc. 16
th
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. 16
th
World Orchid Conf. 284.
2002.
Maxillaria sect. Multiflorae Christenson, Proc. 16
th
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).
Plants of this group almost always have pseudob-
ulbs (with the exception of some species in the
M.
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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 classifica-
tion. 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 examina-
tion 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 speci-
men of
Maxillaria modesta Schltr. nested in Maxillaria
sensu stricto (Koehler 0351, UEC). This specimen is,
however, the true
Maxillaria candida, which was origi-
nally identified simply as “
M. modesta”. The illegiti-
mate name
Maxillaria modesta Brade (non Schltr.) and
its later legitimate name
M. modestiflora Pabst are pre-
viously undetected synonyms of
M. candida.
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. attenu-
ata
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. ben-
nettii
Christenson, M. bocazensis D. E. Benn. &
Christenson,
M. bolivarensis C. Schweinf., M. brachy-
bulbon
Schltr., M. bradei Schltr. ex Hoehne, M. bre-
viscapa
Poepp. & Endl., M. buchtienii Schltr., M. bur-
tonii
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. galan-
tha
J. T. Atwood & Carnevali, M. gentryi Dodson, M.
gorbatschowii
R. Vásquez, Dodson & Ibisch, M. gran-
diflora
(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
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527
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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. longilo-
ba
(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. mar-
moliana
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. monan-
tha
Barb. Rodr., M. multiflora Barb. Rodr., M.
mungoschraderi
R. Vásquez & Ibisch, M. nanegalen-
sis
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. pau-
ciflora
Barb. Rodr., M. pentura Lindl., M. perryae
Dodson, M. platyloba Schltr., M. platypetala Ruiz &
Pav.,
M. plicata Schltr., M. porrecta Lindl., M. por-
tillae
Christenson, M. powellii Schltr., M. pseudore-
ichenheimiana
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. sima-
coana
Schltr., M. simplicilabia C. Schweinf., M. spe-
ciosa
Rchb.f., M. spiritu-sanctensis Pabst, M. splen-
dens
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. xylobi-
iflora
Schltr., M. yanganensis Dodson, and M.
yauaperyensis
Barb. Rodr.
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 rhizomati-
bus 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 lateral-
ibus.
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 former-
ly treated as
Maxillaria section Ebulbes (the
“
Maxillaria graminifolia suballiance”, Atwood
2003), which have thin, wiry, monopodial stems com-
pletely 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 col-
umn 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 dendro-
bioides
, the type of Adamanthus, is morphologically
09 Blanco [24] 5/12/07 00:23 Página 13
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.
E
TYMOLOGY: The name Maxillariella is a diminutive
of
Maxillaria, in reference to the often much smaller
flowers of
Maxillariella.
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.
Maxillariella cobanensis (Schltr.) M. A. Blanco &
Carnevali,
comb. nov.
Basionym: Maxillaria cobanensis Schltr., Repert.
Spec. Nov. Regni Veg. 10: 295. 1912.
Maxillariella costaricensis (Schltr.) M. A. Blanco &
Carnevali,
comb. nov.
Basionym: Maxillaria costaricensis Schltr., Rep.
Spec. Nov. Regni Veg. Beih. 19: 232-233. 1923.
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.
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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-examina-
tion of herbarium material, we are now convinced it
belongs in
Maxillariella.
Maxillariella oreocharis (Schltr.) M. A. Blanco &
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 johan-
nis
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.
09 Blanco [24] 5/12/07 00:23 Página 15
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 tuerck-
heimii
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.
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. rin-
gens
(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. 16
th
World Orchid Conf. 285-286. 2002.
In this expanded circumscription, Mormolyca can
be distinguished by its unifoliate pseudobulbs hav-
ing 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 insec-
tiform. 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 dif-
ficult 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
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herbarium specimens, but distinguishable in life
especially by their floral fragrances (Christenson
2002a, 2002b, and personal observation of the
authors).
Mormolyca acutifolia (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria acutifolia Lindl., Edwards’s
Bot. Reg. 25: misc. 92. 1839.
Mormolyca aureoglobula (Christenson) M. A.
Blanco,
comb. nov.
Basionym: Maxillaria aureoglobula Christenson,
Orchids 71: 125-126. 2002.
Mormolyca chacoensis (Dodson) M. A. Blanco,
comb. nov.
Basionym: Maxillaria chacoensis Dodson, Icon. Pl.
Trop., II, 6: t. 531. 1989.
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 & Tapia-
Muñoz, 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 rufescens (Lindl.) M. A. Blanco, comb. nov.
Basionym: Maxillaria rufescens Lindl., Edwards’s
Bot. Reg. 22: t. 1848. 1836.
Mormolyca sanantonioensis (Christenson) M. A.
Blanco,
comb. nov.
Basionym: Maxillaria sanantonioensis
Christenson, Orchids 71: 128. 2002.
Mormolyca schlimii (Linden & Rchb.f.) M. A.
Blanco,
comb. nov.
Basionym: Chrysocycnis schlimii Linden &
Rchb.f., Bonplandia (Hannover) 2: 280. 1854.
Mormolyca sotoana (Carnevali & Gómez-Juárez) M.
A. Blanco,
comb. nov.
Basionym: Maxillaria sotoana Carnevali &
Gómez-Juárez, Brittonia 53: 461-463.
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 para-
phyletic
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,
09 Blanco [24] 5/12/07 00:23 Página 17
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. 16
th
World Orchid Conf. 285. 2002.
Maxillaria sect. Siagonanthus (Poepp. & Endl.)
Christenson, Proc. 16
th
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, campan-
ulate 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.
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: 1138-
1139. 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.
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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.
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. aggre-
gatum
Rchb.f., O. aureum Poepp. & Endl., O. breve
Schltr. (Maxillaria palmensis Dodson), O. chrysocy-
cnoides
Schltr., O. coccineum (Jacq.) Salisb. ex R.
Br.,
O. conduplicatum Ames & C. Schweinf., O.
croceorubens
Rchb.f., O. distichum Lindl., O. ful-
gens
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. pur-
pureolabium
(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 pseudob-
ulbs. The pseudobulbs are completely covered and fused
09 Blanco [24] 5/12/07 00:23 Página 19
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. huancabam-
bae
(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 pendu-
lae foliis equitantibus. Flores plerumque virides, nun-
quam late expansi, partibus perianthii rigidis acumi-
natis. Labella elobulata, secretionibus resinosis vel
ceraceis. Fructus capsulares dehiscentiis lateralibus.
Plants of Rhetinantha are sub-cespitose to long rhi-
zomatous; the pseudobulbs are often ridged, and have
two to four apical leaves, and usually one or two sub-
tending foliaceous sheaths. The inflorescences often
arise from rhizome bracts a few shoots behind the
most recent pseudobulb. The flowers are campanu-
late, 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 project-
ed downward from the apex of the clinandrium. The
capsules have lateral dehiscence.
Rhetinantha witse-
nioides
is highly modified and has pendent, monopo-
dial shoots with ensiform, glaucous leaves.
E
TYMOLOGY: From the Greek words “rhetinos”
(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.
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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.
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).
Rhetinantha schistostele (Schltr.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria schistostele Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19: 303-304. 1923.
Rhetinantha scorpioidea (Kraenzl.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria scorpioidea Kraenzl.,
Svensk. Vet. Akad. Handl. 46: 71. 1911.
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. 16
th
World Orchid Conf. 286. 2002.
Plants of
Sauvetrea are cespitose to moderately long-
rhizomatous. The unifoliate pseudobulbs are frequently
ancipitous and are subtendend by a pair of non-folia-
ceous bracts. The inflorescences arise from the base of
the newly emerging pseudobulbs and have strongly
ancipitous bracts. The column foot is short. The flow-
ers have spreading perianth segments without strong
fibers. Both the ovaries and the mature fruits are
strongly trigonous. The labellum has a ligulate, sulcate
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.
Sauvetrea chicana (Dodson) M. A. Blanco, comb.
nov
.
Basionym: Maxillaria chicana Dodson,
Orquideología 19(3): 61. 1994.
Sauvetrea cornuta (C. Schweinf.) M. A. Blanco,
comb. nov.
Basionym: Maxillaria cornuta C. Schweinf., Bot.
Mus. Leafl. 11: 265. 1945.
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.
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
´
09 Blanco [24] 5/12/07 00:23 Página 21
(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 taxo-
nomic revision. The strange
Maxillaria grobyoides
Garay & Dunst. may belong in Sauvetrea, but we pre-
fer 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 long-
rhizomatous. The ridged pseudobulbs have one to
four apical leaves and are subtended by non-folia-
ceous bracts. The inflorescences are erect, often long
and wiry, and emerge from the most recent pseudob-
ulbs. 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).
A
CKNOWLEDGEMENTS. Eric A. Christenson (BRIT) pro-
vided general discussions on taxonomy and data about the
publication of his sections of
Maxillaria. Gustavo Romero
(AMES), Cassio van den Berg (HUEFS) and two anon-
mous 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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