UNIVERSITY OF
ILLINOIS LIBRARY
AT URBANA-CHAMPA1GN
BIOLOGY
Botany
SERIES, NO. 40
FLORA COSTARICENSIS
William Burger, Editor
Family #39 Orchidaceae: Tribe Maxillarieae:
Subtribes Maxillariinae and Oncidiinae
John T. Atwood
Dora Emilia Mora de Retana
April 30, 1999
Publication 1500
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
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FIELDIANA
Botany
NEW SERIES, NO. 40
FLORA COSTARICENSIS
William Burger, Editor
Family #39 Orchidaceae: Tribe Maxillarieae:
Subtribes Maxillariinae and Oncidiinae
John T. Atwood
Director, Orchid Identification Center
Marie Selby Botanical Gardens
811 South Palm Avenue
Sarasota, Florida 34236
U.S. A.
Dora Emilia Mora de Retana
Director, Lankester Gardens
Universidad de Costa Rica
Escuela de Biologia
San Jose 1000, Costa Rica
Accepted October 27, 1997
Published April 30, 1999
Publication 1500
Research Associate
Department of Botany
Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605-2496
U.S.A.
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1999 Field Museum of Natural History
ISSN 0015-0746
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
DEDICATION v
INTRODUCTION v
ACKNOWLEDGMENTS vi
SUBTRIBE MAXILLARIINAE 1
Key to the Genera of Subtribe Maxillariinae
1
Descriptions of Genera and Species 2
Cryptocentrum 2
Maxillaria 32
Mormolyca 84
Trigonidium 85
SUBTRIBE ONCIDIINAE 86
Key to Subtribe Oncidiinae 87
Descriptions of Genera and Species 89
Ada 89
Amparoa 114
Aspasia 114
Brassia 116
Cischweinfia 118
Comparettia 119
Fernandezia 120
Goniochilus 120
Hybochilus 121
lonopsis 122
Leochilus 123
Lockhartia 124
Macroclinium 127
Mesospinidium 132
Miltonlopsis 133
Notylia 134
Oncidium 135
Osmoglossum 150
Otoglossum 151
Pachyphyllum 152
Plectrophora 154
Psychopsis 154
Psygmorchis 155
Rhynchostele 156
Rodriguezia 159
Rossioglossum 159
Scelochilus 160
Sigmatostalix 161
Systeloglossum 163
Ticoglosswn 164
Trie hoc entrum 165
Trichopilia 1 69
Trizeuxis 172
Warmingia 172
ADDENDUM TO SUBTRIBE MAXILLARIINAE 173
ADDENDUM TO SUBTRIBE ONCIDIINAE 175
LITERATURE CITED 176
INDEX . .178
List of Illustrations
MAXILLARIINAE
1. Cryptocentrum standleyi, C. latifolium,
C. calcaratum, C. lehmannii 3
2. Maxillaria lueri, M. alfaroi, M. inaudi-
ta, M. valerioi 4
3. Maxillaria linearifolia, M. appendicu-
loides, M. dendrobioides, M. trilobata
5
4. Maxillaria adendrobium, M. nicara-
guensis, M. tonduzii, M. parvilabia 6
5. Maxillaria biolleyi, M. quadrata, M.
adolphii, M. conduplicata 7
6. Maxillaria fulgens, M. pittieri, M. exal-
tata, M. meridensis 8
7. Maxillaria monteverdensis, M. amabi-
lis, M. falcata, M. acervata, M. poner-
antha, M. elatior 9
8. Maxillaria tubercularis, M. ctenostach-
ya, M. friedrichsthalii, M. scorpioidea
10
9. Maxillaria aciantha, M. brevilabia, M.
neglecta, M. parviflora, M. concavila-
bia 11
10. Maxillaria horichii, M. pseudoneglecta,
M. uncata, M. tenuifolia, M. sanguinea
12
1 1 . Maxillaria planicola, M. tigrina, M.
variabilis, M. costaricensis, M. haberi
13
12. Maxillaria muscoides, M. microphyton,
M. burgeri, M. flava 14
13. Maxillaria wercklei, M. lankesteri, M.
minor, M. sigmoidea, M. paleata, M.
vaginalis 15
14. Maxillaria schlechteriana, M. dichoto-
ma, M. suaveolens, M. bradeorum, M.
ampliflora 16
15. Maxillaria umbratilis, M. campanulata,
M. oreocharis, M. alba 17
16. Maxillaria anceps, M. diuturna, M.
bracteata, M. gomeziana 18
17. Maxillaria serrulata, M. crassifolia, M.
maleolens, M. nasuta 19
111
18. Maxillaria valenzuelana, M. angustissi-
ma, M. chartacifolia, M. bicallosa 20
19. Maxillaria longipetiolata, M. strumata,
M. piestopus, M. longiloba 21
20. Maxillaria ringens, M. endresii, M. an-
gustisegmenta, M. cryptobulbon 22
21. Maxillaria brunnea, M. acutifolia, M.
hedwigiae, M. dressleriana 23
22. Maxillaria moralesii, M. obscura, M.
punctostriata, M. cedralensis 24
23. Maxillaria rodrigueziana, M. attenu-
ata, M. confusa, M. ramonensis 25
24. Maxillaria reichenheimiana, M. pachy-
acron, M. arachnitiflora, M. chionan-
tha 26
25. Maxillaria brachybulbon, M. vittariifol-
ia, M. acostae, M. caespitifica 27
26. Mormolyca ringens, Trigonidium eger-
tonianum, T. riopalenquense, T. lankes-
teri
ONCIDIINAE
27. Ada chlorops, Amparoa costaricensis,
Aspasia epidendroides, Aspasia prin-
cipissa 90
28. Brassia arcuigera, B. caudata, B. gi-
reoudiana, B. verrucosa 91
29. Cischweinfia dasyandra, Cischweinfia
pusilla, Comparettia falcata, Fernande-
zia tica 92
30. Goniochilus leochilinus, lonopsis utri-
cularioides, Leochilus labiatus, L. tri-
cuspidatus 93
31. Lockhartia amoena, L. hercodonta, L.
micrantha, L. oerstedii 94
32. Lockhartia acuta, L. pittieri, Macrocli-
nium confertum, M. generalense 95
33. Macroclinium cordesii, Macroclinium
ramonense, Mesospinidium horichii,
Mesospinidium warscewiczii 96
34. Miltoniopsis warscewiczii, Notylia tri-
sepala, N. pittieri, N. pittieri 97
35. Oncidium ascendens, O. teres, O. ce-
bolleta, O. carthagenense, O. schroe-
derianum . .98
36. Oncidium cariniferum, O. stenoglos-
sum, O. panduriforme, O. storkii 99
37. Oncidium globuliferum, O. cristagalli,
O. warscewiczii, O. bryolophotum 100
38. Oncidium bracteatum, O. luteum, O.
cheirophorum, O. ampliatum 101
39. Oncidium obryzatoides, O. klotzschia-
num, O. dichromaticum, O. parviflorum
102
40. Oncidium ansiferum, O. stenobulbon,
O. ensatum, O. stenotis 103
41. Oncidium polycladium, Oncidium isth-
mi, Osmoglossum convallarioides, Os-
moglossum egertonii, Otoglossum chi-
riquense 104
42. Pachyphyllum crystallinum, Pachyphyl-
lum hispidulum, Plectrophora alata,
Psychopsis krameriana 105
28 43. Psygmorchis pumilio, Rhynchostele
bictoniensis, R. hortensiae, R. stellata
106
44. Rodriguezia compacta, Rossioglossum
schlieperianum, Scelochilus aureus,
Sigmatostalix unguiculata 107
45. Sigmatostalix picta, S. macrobulbon, S.
brownii, S. hymenantha 108
46. Sigmatostalix adamsii, Systeloglossum
acuminatum, Systeloglossum costari-
cense, Ticoglossum krameri 109
47. Ticoglossum oerstedii, Trichocentrum
dianthum, Trichocentrum pfavii, Tri-
chocentrum estrellense 110
48. Trichocentrum caloceras, T. costari-
cense, T. brenesii, T. capistratum Ill
49. Trichocentrum cymbiglossum, Trichopi-
lia maculata, Trichopilia suavis, Tri-
chopilia turialbae 112
50. Trichopilia tortilis, Trichopilia margin-
ata, Trichopilia galeottiana, Trizeuxis
falcata 113
MAXILLARIINAE ADDENDUM
51. Maxillaria cacaoensis . .174
IV
Dedication
Costa Rica has a proud history of enlightened
botanists, both native- and foreign-born. This vol-
ume is dedicated to A. R. Endres, a little-known
but productive plant collector and illustrator who
died in 1877. Almost nothing is known about him,
and he may not have been Costa Rican. What we
do know is that he left copious specimens and was
one of the most precise illustrators of minute or-
chids. His last name is Spanish, yet he wrote his
notes in English. He was referred to as a "half-
caste" (Veitch, J. H. Hortus Veitchii, 1906), a la-
bel that may indicate parentage of a North Amer-
ican or European and a Latin American. Most of
the specimens were ignored during Reichenbach's
life, but some of these were actually new species
only recently described by C. A. Luer (Lindleyana
10:133-173. 1995). Examination of Endres'
drawings under a dissecting microscope reveals
incredibly close shade lines that could be accom-
plished only by an unusually steady hand. Still
more unbelievable was his ability to accurately
observe minute details of some of the tiniest or-
chid flowers. As Reichenbach stated, he was "one
of those collectors who cared more for science
than for sovereigns" (Card. Chron. 19:432. 1883).
The life work of Endres is preserved at the Reich-
enbach Herbarium (w) of the Naturhistorisches
Museum, Vienna, Austria.
Introduction
This is the first of a series treating the Orchi-
daceae for Flora Costaricensis. It represents the
collaborative efforts between authors of very dif-
ferent experience, backgrounds, and cultures,
which we believe have complemented one another
in the finished product. Usually, such products are
made either by a native in the tropics with little
access to original materials or by botanists from
northern countries with limited access to live ma-
terial. This product combines the assets of bota-
nists from both perspectives. German Carnevali
provided the treatment of Cryptocentrum using
his valuable doctoral dissertation, and Franco
Pupulin provided treatments of Macroclinium and
Trichocentrum with D. E. Mora.
This volume features two subtribes, Maxillari-
inae and Oncidiinae, according to the classifica-
tion of R. L. Dressier (1981, see references cited
within both subtribes). Two hundred twenty-nine
species, or about one-fifth of Costa Rican Orchi-
daceae, are treated. These are among the most
conspicuous of Costa Rica's epiphytic orchids,
and therefore this treatment should have broad
utility among scientists, horticulturists, and the lay
public. We are particularly confident in the con-
tribution of two large genera, Maxillaria (107 spe-
cies treated) and Oncidium (28 species treated),
with many previously confused, misapplied, and
superfluous names. This work is based on all
available specimens examined from AMES, CR, D,
F, INB, MO, and us, supplemented by specimens
from herbaria mentioned in the Acknowledg-
ments. Types have been examined as far as is
practical, but our access to types has been pri-
marily through microfiche collections of the Ames
Herbarium at Harvard University, the Lindley
Herbarium at Kew, and the Reichenbach Herbar-
ium in Vienna. In the text, these are indicated as
"photo seen." The senior author also has made
several trips to the Ames Herbarium in search of
types, and one trip to Kew and Vienna. Most of
the Maxillariinae have been catalogued on AMO-
DATA and will be converted to TROPICOS.
Specimens of Oncidiinae continue to be cata-
logued directly on TROPICOS.
To clarify nomenclature, several specimens are
selected as lectotypes or neotypes, and one is se-
lected as an epitype. Neotypes are selected for
Maxillaria acostae Schltr. (AMES), M. piestopus
Schltr. (AMES), M. rouseauae Schltr. (MO), and Or-
nithidium wercklei Schltr. (AMES). Lectotypes are
selected for M. acervata Rchb. f. (w), M. ringens
Rchb. f. (w), Oncidium fulgens Schltr. (AMES), On-
cidium globuliferum Kunth var. costaricense
Rchb. f. (w), Oncidium naranjense Schltr. (AMES),
Oncidium warscewiczii Rchb. f. (w), Ornithidium
costaricense Schltr. (AMES), and Ornithidium ton-
duzii Schltr. (us). An epitype is selected for M.
pachyacron Schltr. (SEL) to serve as a reference
specimen for interpreting Schlechter's original de-
scription and drawings. These names can be found
in the Index.
Many of the individual species descriptions are
based on Icones Plantarum Tropicarum (series 1),
fascicles 14, 15, and 16. Many more specimens
have been examined since the appearance of these
works, with subsequent modification of the text.
More than 92% of the species in this volume
are illustrated. The illustrations are organized ac-
cording to their occurrence in the key rather than
alphabetically, which we believe will facilitate de-
termination when consulted with the key. The in-
dividual species treatments, which are arranged
alphabetically, may be consulted as indexes.
Acknowledgments
We thank the herbarium staffs of the Oakes
Ames Herbarium of Harvard University (AMES),
Museo Nacional (CR), Field Museum of Natural
History (F), Institute de Biodiversidad (INB), Mis-
souri Botanical Garden (MO), Selby Gardens (SEL),
National Herbarium (us), and Universidad de
Costa Rica (us;) for general support in this project
and for providing access to collections. We also
thank several other herbaria and their curators for
loans, some of which have been borrowed for
more than 10 years. These include Duke Univer-
sity (DUKE), Michigan State University (MSC), the
University of Michigan (MICH), New York Botan-
ical Garden (NY), and the University of Texas
(TEX). Special thanks are extended to the Missouri
Botanical Garden, which has supported the botan-
ical efforts of Selby Botanical Gardens from the
beginning. We especially thank Peter Raven, the
director, as well as members of his staff. Michael
Grayum and Barry Hammel supported the project
from its initiation in 1988. William Haber provid-
ed much logistic support, as well as assistance in
field efforts in the Monteverde region, and served
as a knowledgeable and enthusiastic field com-
panion. William Burger of the Field Museum has
supported the project from the beginning.
There are too many modern collectors to thank
for herbarium collections, but we would like to
acknowledge some notable historical collectors:
A. Alfaro, A. Brenes, A. R. Endr6s, O. Jimenez,
C. Lankester, P. Standley, and J. Valeric. Alex-
ander Skutch, thankfully still with us, is the last
of these illustrious collectors. Much of the Selby
herbarium is based on an excellent collection of
Costa Rican Orchidaceae assembled by L. O. Wil-
liams.
We have many notable botanists to thank for
annotating the specimens before us. In particular,
we thank Louis O. Williams, Charles Schwein-
furth, and Oakes Ames. Bob Dressier reviewed
specimens of the Oncidiinae before us.
Dora E. Mora thanks Joaquin Garcia for shar-
ing locality data and vegetative material and for
valuable comments on the manuscript of the On-
cidiinae. She also thanks Maria del Carmen Calvo
for secretarial assistance.
John T. Atwood thanks W. J. Kress of the Na-
tional Museum of Natural History (us) for en-
couragement in the project and D. H. Nicolson for
sharing his expertise regarding application of
names, for helping us solve a variety of difficult
nomenclatural problems, and especially for shar-
ing his philosophy of the use of types.
We thank Calaway H. Dodson for fruitful dis-
cussions and for the use of several illustrations
from Icones Plantarum Tropicarum. Robert L.
Dressier freely shared his research on Costa Rican
Oncidiinae with us, notably concerning the genera
Notylia and Sigmatostalix. German Carnevali pro-
vided the treatment of Cryptocentrum. Franco
Pupulin provided treatments of Macroclinium and
Trichocentrum with D. E. Mora. Lastly, Joaqin
Garcia-Castro provided a treatment of Leucohyle
with D. E. Mora as an Addendum.
Special thanks are extended to curators of two
European herbaria. Phillip Cribb and Jeff Wood
granted free access to the herbarium at Kew, and
Harald Riedl was most helpful by making avail-
able the extensive Reichenbach and Kranzlin col-
lections at the Naturhistorisches Museum in Vi-
enna, Austria.
We thank the many artists who worked with us
over the several years it took to complete this
work, notably Omar Achi, Luis Alfaro, G. Bar-
boza, B. N. Culbertson, Carmen Marin, Stig Dal-
strom, and Willow Zuchowski.
Lastly, we thank our institutions, Marie Selby
Botanical Gardens and the University of Costa
Rica, for allowing us salaried time to write this
volume and to do the necessary field and library
research leading to it.
This material is based on work supported by
the National Science Foundation (NSF) and by the
Office of Forestry, Environment, and Natural Re-
sources, Bureau of Science and Technology, of
the U.S. Agency for International Development
(NSF grant DEB-9200812).
VI
FLORA COSTARICENSIS
Family #39 Orchidaceae
ORCHIDACEAE
By John T. Atwood and
Dora Emilia Mora de Retana
Subtribe Maxillariinae Benth.
REFERENCES — R. L. Dressier, The Orchids. Har-
vard University Press, Cambridge, Massachusetts,
and London, England, 332 pp. 1981. R. L. Dress-
ier, Phylogeny and Classification of the Orchid
Family. Dioscorides Press, Portland, Oregon, 314
pp. 1993. L. O. Williams, An enumeration of the
Orchidaceae of Central America, British Hondu-
ras and Panama. Ceiba 5:1-256. 1956.
Plant usually epiphytic, or opportunistically terrestri-
al, cespitose or rhizomatous, sometimes with rhizomes
forming upright canes, sympodial or less commonly
monopodial, sometimes sympodial as juveniles and
monopodial as adults; roots with a white velamen, of
varying maximum diameter depending on the species,
decurrent on upright stems. Pseudobulb present or ab-
sent, when present of a single internode, usually com-
pressed, sometimes grooved, always terminating the
shoot. Leaves articulate or rarely secondarily lacking ar-
ticulations, always conduplicate, rarely equitant, distrib-
uted along the rhizome as well as at the pseudobulb apex
or restricted to the pseudobulb apex, usually distichous
and very rarely spiraled. Inflorescence short or long,
lateral on the stem, at the base of the pseudobulb if
present, always a single-flowered scape, sometimes
formed in clusters (fascicles) within each rhizome bract
of what appear to be very condensed, aborted, vegetative
shoots. Flowers exhibiting a great deal of variability,
spreading or campanulate, of various colors including
green and brown, fragrant or not. Sepals free or vari-
ously connate, especially the lateral sepals, sometimes
forming a spur. Petals usually similar to the sepals. Lip
fixed or hinged to the receptacle or column foot, simple
to 3-lobate, variously adorned with calli and hairs. Col-
umn usually cylindric, straight or more commonly ar-
cuate, sometimes adorned with wings (Cryptocentrum);
pollinia 4, dorso-ventrally superposed in 2 pairs, sup-
ported on a well-defined viscidium and usually a stipe.
Fruit a capsule.
A Neotropical subtribe of 460-500 species con-
tained in eight genera (recognizing Maxillaria,
sensu laid). Four genera with a total of 117 spe-
cies are found in Costa Rica.
This well-defined subtribe with conduplicate
leaves is easily distinguished from closely related
Lycastinae Schltr. By the plicate leaves of the lat-
ter. One of the most interesting aspects of the
Maxillariinae is the variety of growth patterns
summarized above, especially those that are sym-
podial as juveniles, with each abbreviated shoot
terminated by a pseudobulb, but monopodial as
adults, producing extensive canes lacking pseu-
dobulbs altogether. Perhaps the pseudobulb serves
to buffer water stress in small plants, which often
live in usually wet but extremely windy habitats.
The genus Maxillaria is here accepted in the tra-
ditional broad sense, including segregates Cama-
ridium Lindl., Ornithidium R. Br., Pseudomaxilla-
ria Brieger, and Sepalosaccus Schltr. We ac-
knowledge that data from DNA studies currently
being generated and analyzed cladistically should
help define useful genera within the Maxillariinae.
Key to the Genera of Subtribe Maxillariinae
la. Plants with distinct, cylindric sepaline spur Cryptocentrum
Ib. Plants not with cylindric sepaline spur, although sepals may be joined 2
2a. Flowers with sepals adpressed into a cup at the base with reflexed apices, petals barely or not
exerted beyond the reflexed part of the sepal apices giving the flower a trigonous appearance
Trigonidium
2b. Flowers not as above 3
3a. Column lacking a distinct foot, i.e., the base of the column not extended below the ovary;
FIELDIANA: BOTANY, N.S., NO. 40, APRIL 30, 1999, PP. 1-182
scape wiry, the internodes well exposed between scape bracts (Costa Rican), pedicel longer
than the ovary Mormolyca (ringens)
3b. Column with a distinct foot extended below the ovary, scapes with internodes entirely or
nearly concealed by bracts, pedicel absent or if present much shorter than ovary
Maxillaria
Cryptocentrum Benth. & Hook.
(G. Carnevali)
REFERENCES — R. L. Dressier, The systematic
position of Cryptocentrum (Orchidaceae). Britton-
ia 13:266-270. 1961. G. Carnevali, Systematics,
phylogeny, and twig epiphytism in Cryptocentrum
(Orchidaceae). Ph.D. diss., University of Missou-
ri, St. Louis. 1996.
Herb, epiphytic, rarely subterrestrial or lithophytic,
monopodial or less commonly sympodial, if sympodial
then stems pseudobulbous-thickened. Roots originating
from the lower portions of the stems, frequently decur-
rent on and totally hiding the stems. Leaves distichous
or polystichously arranged, conduplicate, coriaceous or
succulent, flat or hemicylindric or 3-edged in cross sec-
tion, articulate, glabrous, margins ciliate or erase; leaf
sheaths usually wider than the blades, imbricate, con-
spicuously nerved. Inflorescences I-flowered, usually
originating from the axils of the lowermost leaf sheaths;
ovary with pedicel 6-edged, subterete, smooth, rarely
winged. Flowers resupinate, nocturnally fragrant, green-
ish or yellowish, often with purplish suffusion. Sepals
basally connate into a tube, free and somewhat divergent
above; tube obconic, often fleshy-thickened; lateral se-
pals often longer than dorsal, basally produced into a
backwardly oriented sepaline spur; sepaline spur cylin-
dric, apically acute or somewhat saccate. Petals free,
basally enclosed by the sepaline tube, the free portion
usually lanceolate or oblong, erect, spreading to some-
what reflexed and then called the bent portion; bent por-
tion frequently somewhat tailed, commonly triangular or
triangular-lanceolate, fleshier than the basal portion. Lip
basally produced into a backwardly oriented spur con-
tained within the sepaline spur, the walls of the labellar
spur fused with the roof of the sepaline spur; the lip
blade without callus, elliptic or ovate when flattened,
divided into a deeply concave or saccate hypochile con-
tained in the sepaline tube and an erect, spreading, or
reflexed, somewhat concave epichile. Column erect,
stout, with 2 wings abaxially; anther terminal, pollinia
4 in 2 superposed pairs, waxy, supported on a stipe and
viscidium. Fruit a capsule, oblongoid or ellipsoid, beak-
less.
A Neotropical genus of 17 species ranging
from the Cordillera de Tilaran in northern Costa
Rica to the Amazonian slopes of the Andes in
Bolivia with two disjunct species in the Guayana
region.
A genus well-defined by several apomorphies,
Cryptocentrum is closely related to Anthosiphon
(G. Carnevali, unpubl. diss.) and appears to have
evolved from that genus by elongation of the se-
paline spur and proportional reduction of the se-
paline cup.
Key to the Species of Cryptocentrum
la. Leaves spiraled (polystichous) on the stem, subterete, < 3 cm long; inflorescence longer than to
subequaling the subtending leaves; floral bract conspicuously shorter than the spur, only enclosing
the apical Vs or less of it 2
2a. Spur 12-14.5 mm long, inflorescence subequaling or little longer than subtending leaves; mono-
podia shorter than leaves on mature plants; leaves (30)40-60(70) mm long, 0.5-1.5 mm thick
when fresh; leaf sheaths not or inconspicously attenuated apically, not becoming clawed; lateral
sepals always 3-nerved C. flavum
2b. Spur (15)16-19 mm long; inflorescence always conspicuously longer than subtending leaves;
monopodia longer than leaves on mature plants; leaves (10)15-25(30) mm long, 0.3-0.5 mm
thick when fresh; leaf sheath apically conspicuously attenuated and becoming clawed; lateral
sepals usually 5 -nerved, only rarely 3-nerved C. standleyi
Ib. Leaves distichous on the stem, subterete or flat, usually > 4 cm long; inflorescence shorter or rarely
subequaling the subtending leaves; floral bract longer than the spur, enclosing at least % of its length
3
3a. Leaves terete or hemiterete, 1-2(3) mm wide and thick C. gracillimum
3b. Leaves flat, 4-10(20) mm wide 4
FIELDIANA: BOTANY
5cm
FIG. 1. A, Cryptocentrum standleyi (Icon. PI. Trop. 14: t. 1309. Illustrator: S. Dalstrom). B, C. latifolium (Icon. PI.
Trop. 14: t. 1308. Illustrator: B. N. Culbertson). C, C. calcaratum (Icon. PI. Trop. 14: t. 1307. Illustrator: B. N.
Culbertson). D, C. lehmannii (Icon. PI. Trop. 1: t. 33. Illustrator: B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
B
1 cm
FIG. 2. A, Maxillaria lueri (Icon. PI. Trop. 2: t. 155. Illustrator: L. Megahee). B, M. alfaroi (illustrator: S. Dalstrom).
C, M. inaudita (Icon. PI. Trop. 14: t. 1349. Illustrator: B. N. Culbertson). D, M. valerioi (illustrator: S. Dalstrom).
FIELDIANA: BOTANY
FIG. 3. A, Maxillaria linearifolia (Icon. PI. Trop. 14: t. 1350. Illustrator: B. N. Culbertson). B, M. appendiculoides
(illustrator: S. Dalstrom). C, M. dendrobioides (illustrator: W. Zuchowsky). D, M. trilobata (Icon. PI. Trop. 14: t.
1367. Illustrator: B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
0.5 mm
FIG. 4. A, Maxillaria adendrobium (illustrator: S. Dalstrom). B, M. nicaraguensis (Icon. PI. Trop. 14: t. 1355. Illus-
trator: S. Dalstrom). C, M. tonduzii (illustrator: S. Dalstrom). D, M. parvilabia (illustrator: S. Dalstrom).
FIELDIANA: BOTANY
FIG. 5. A, Maxillaria biolleyi (illustrator: S. Dalstrom). B, M. quadrata (illustrator: L. Alfaro). C, M. adolphii (illus-
trator: B. N. Culbertson). D, M. conduplicata (illustrator: S. Dalstrom).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
FIG. 6. A, Maxillaria fulgens (Icon. PI. Trop. 14: t. 1348. Illustrator: B. N. Culbertson). B, M. pittieri (illustrator: S.
Dalstrom). C, M. exaltata (illustrator: S. Dalstrom). D, M. meridensis (Icon. PI. Trop. 14: t. 1352. Illustrator: B. N.
Culbertson).
FIELDIANA: BOTANY
10cm
1 cm
FIG. 7. A, Maxillaria monteverdensis (Lindleyana 9:241. Illustrator: G. Barboza). B, M. amabilis (Lindleyana 9:241.
Illustrator: J. T. Atwood). Note: Plant habit is similar to that in Figure 7A. C, M. falcata (illustrator: S. Dalstrom).
D, M. acervata (Icon. PI. Trop. 14: t. 1346 [as synonym, M.foliosa]. Illustrator: B. N. Culbertson). E, M. ponerantha
(Icon. PI. Trop. 14: t. 1358. Illustrator: B. N. Culbertson). F, M. elatior (illustrators: plant habit, B. N. Culbertson;
floral diagnosis, L. Alfaro).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
FIG. 8. A, Maxillaria tubercularis (Lindleyana 9:229. Illustrator: G. Barboza). B, M. ctenostachya (illustrator: S.
Dalstrom). 'C, M. friedrichsthalii (Icon. PI. Trop. 14: t. 1347. Illustrator: B. N. Culbertson). D, M. scorpioidea (Icon.
PI. Trop. 14: t. 1363. Illustrator: B. N. Culbertson).
10
FIELDIANA: BOTANY
B
5 mm
FIG. 9. A, Maxillaria aciantha (Icon. PI. Trop. 14: t. 1333. Illustrator: B. N. Culbertson). B, M. brevilabia (illustrator:
S. Dalstrom). C, M. neglecta (illustrators: plant habit, S. Dalstrom; floral diagnosis, Lindleyana 8:27, S. Dalstrom).
D, M. parviflora (Lindleyana 8:27. Illustrator: S. Dalstrom). Note: Plant habit is similar to that in Figure 9E. E, M.
concavilabia (illustrators: plant habit, S. Dalstrom; floral diagnosis, Lindleyana 8:27, S. Dalstrom).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
11
FIG. 10. A, Maxillaria horichii (illustrators: plant habit, S. Dalstrom; floral diagnosis, Lindleyana 8:27, S. Dalstrom).
B, M. pseudoneglecta (Lindleyana 8:27. Illustrator: S. Dalstrom). Note: Plant habit is similar to that in Figure 10A.
C, M. uncata (Icon. PI. Trop. 14: t. 1369. Illustrator: S. Dalstrom). D, M. tenuifolia (Icon. PL Trap. 14: t. 1366.
Illustrator: B. N. Culbertson). Note: Lip from rehydrated flower, somewhat straightened. E, M. sanguined (illustrators:
plant habit, Icon. PL Trop. 14: t. 1362, S. Dalstrom; floral diagnosis, Icon. PL Trop. 15: t. 1362a, L. Alfaro).
12
FTELDIANA: BOTANY
FIG. 11. A, Maxillaria planicola (illustrator: S. Dalstrom). B, M. tigrina (Icon. PI. Trop. 14: t. 1306 [as synonym,
Chrysocycnis tigrinum]. Illustrator: B. N. Culbertson). C, M. variabilis (illustrator: S. Dalstrom). Note: Plant habit is
similar to that in Figure 1 ID. D, M. costaricensis (illustrator: S. Dalstrom). E, M. haberi (Selbyana 16:244. Illustrator:
B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
13
FIG. 12. A, Maxillaria muscoides (Lindleyana 9:235. Illustrator: G. Barboza). B, M. microphyton (illustrator: S.
Dalstrom). C, M. burgeri (Lindleyana 9:234. Illustrator: G. Barboza). D, M. flava (illustrator: S. Dalstrom).
14
FIELDIANA: BOTANY
FIG. 13. A, Maxillaria wercklei (illustrator: S. Dalstrom). B, M. lankesteri (illustrator: S. Dalstrom). Note: Plant habit
is similar to that in Figure 13A. C, M. minor (Icon. PI. Trop. t. 14: 1353. Illustrator: B. N. Culbertson). D, M.
sigmoidea (illustrator: S. Dalstrom). E, M. paleata (illustrator: S. Dalstrom). F, M. vaginalis (illustrator: S. Dalstrom).
Note: Plant habit is similar to that in Figure 13D.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
15
FIG. 14. A, Maxillaria schlechteriana (Lindleyana 9:230. Illustrator: S. Dalstrom). B, M. dichotoma (illustrator: S.
Dalstrom). Note: Plant habit is similar to that in Figure 14A. C, M. suaveolens (Icon. PI. Trop. 14: t. 1365. Illustrator:
B. N. Culbertson). D, M. bradeorum (illustrator: S. Dalstrom). E, M. ampliflora (illustrator: S. Dalstrom).
16
FIELDIANA: BOTANY
FIG. 15. A, Maxillaria umbratilis (Icon. PI. Trop. 14: t. 1368. Illustrator: B. N. Culbertson). B, M. campanulata
(illustrator: S. Dalstrom). C, M. oreocharis (Icon. PI. Trop. 14: t. 1356. Illustrator: B. N. Culbertson). D, M. alba
(Icon. PI. Trop. 14: t. 1334. Illustrator: B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
17
1 cm
FIG. 16. A, Maxillaria cmceps (Icon. PI. Trop. 14: t. 1335. Illustrator: B. N. Culbertson). B, M. diuturna (illustrator:
S. Dalstrom). C, M. bracteata (illustrators: plant habit, S. Dalstrom; floral diagnosis, Lindleyana 9:240, J. T. Atwood).
D, M. gomeziana (illustrator: S. Dalstrom).
18
FIELDIANA: BOTANY
FIG. 17. A, Maxillaria serrulata (illustrator: S. Dalstrom after J. T. Atwood). B, M. crassifolia (Icon. PI. Trop. 14: t.
1341. Illustrator: B. N. Culbertson). C, M. maleolens (Icon. PI. Trop. 14: t. 1351. Illustrator: B. N. Culbertson). D,
M. nasuta (Icon. PI. Trop. 14. t. 1354. Illustrator: B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
19
FIG. 18. A, Maxillaria valenzuelana (Icon. PI. Trop. 14: t. 1370. Illustrator: B. N. Culbertson). B, M. angustissima
(Icon. PI. Trop. 14: t. 1337. Illustrator: B. N. Culbertson). C, M. chartacifolia (Icon. PI. Trop. 14: t. 1340. Illustrator:
B. N. Culbertson). D, M. bicallosa (illustrator: S. Dalstrom).
20
FIELDIANA: BOTANY
1 cm
FIG. 19. A, Maxillaria longipetiolata (Icon. PI. Trop. 16: t. 1563. Illustrator: S. Dalstrom). B, M. strumata (illustrator:
L. Alfaro). C, M. piestopus (illustrator: S. Dalstrom after J. T. Atwood). D, M. longiloba (Icon. PI. Trop. 16: t. 1562.
Illustrator: S. Dalstrom).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
21
2cm
4cm
FIG. 20. A, Maxillaria ringens (illustrator: S. Dalstrom). B, M. endresii (Icon. PI. Trop. 14: t. 1345. Illustrator: S.
Dalstrom). C, M. angustisegmenta (Icon. PI. Trop. 14: t. 1336. Illustrator: B. N. Culbertson). D, M. cryptobulbon
(illustrator: S. Dalstrom).
22
FIELDIANA: BOTANY
B
FIG. 21. A, Maxillaria brunnea (illustrator: S. Dalstrom). B, M. acutifolia (illustrator: S. Dalstrom). C, M. hedwigiae
(illustrator: S. Dalstrom). D, M. dressleriana (illustrator: B. N. Culbertson).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
23
B
FIG. 22. A, Maxillaria moralesii (Lindleyana 11:31. 1996. Illustrator: B. N. Culbertson). B, M. obscura (illustrator:
S. Dalstrom). C, M. punctostriata (illustrator: S. Dalstrom). D, M. cedralensis (illustrator: J. T. Atwood).
24
FIELDIANA: BOTANY
B
FIG. 23. A, Maxillaria rodrigueziana (Icon. PI. Trop. 14: t. 1361. Illustrator: B. N. Culbertson). B, M. attenuata
(illustrator: S. Dalstrom). C, M. confusa (illustrator: S. Dalstrom). D, M. ramonensis (illustrator: S. Dalstrom).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
25
FIG. 24. A, Maxillaria reichenheimiana (Icon. PI. Trop. 14: t. 1359. Illustrator: B. N. Culbertson). B, M. pachyacron
(Icon. PI. Trop. 14: t. 1357. Illustrator: B. N. Culbertson). C, M. arachnitiflora (Icon. PI. Trop. 14: t. 1338. Illustrator:
B. N. Culbertson). D, M. chionantha (Selbyana 16:242. Illustrator: B. N. Culbertson after G. Barboza).
26
FIELDIANA: BOTANY
B
2mm
M
FIG. 25. A, Maxillaria brachybulbon (Icon. PI. Trop. 14: t. 1339. Illustrator: B. N. Culbertson). B, M. vittariifolia
(Icon. PI. Trop. 14: t. 1372. Illustrator: S. Dalstrom). C, M. acostae (illustrators: plant habit, S. Dalstrom; floral
diagnosis, J. T. Atwood). D, M. caespitifica (illustrator: S. Dalstrom, based on a Nicaraguan collection [Atwood
6978]).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
27
5 mm
FIG. 26. A, Mormolyca ringens (illustrator: S. Dalstrom). B, Trigonidium egertonianum (Icon. PL Trop. 4: t. 345.
Illustrator: L. Megahee). C, T. riopalenquense (Icon. PL Trop. 14: t. 1399. Illustrator: S. Dalstrom). D, T. lankesteri
(Icon. PL Trop. 4: t. 346. Illustrator: W. B. Zomlefer, based on an Ecuadorian plant probably misdetermined as T.
insigne Rchb. f. ex Benth. & Hook.).
28
FIELDIANA: BOTANY
4a. Leaves (8)10-16(20) mm wide, leaf apex with longer lobe exceeding the shorter by (2)3-
5(12) mm C. latifolium
4b. Leaves (3)5-6(8) mm wide, leaf apex with longer lobe exceeding the shorter by 0.5-1.5
mm 5
5a. Inflorescence with 3-4(5) internodes, the floral bract comprising V^-Vi of total inflores-
cence length; bracts of the inflorescence somewhat inflated; perianth segments clear
green or yellow-green, usually not tinged with other colors C. calcaratum
5b. Inflorescence with (5)6-11 internodes, the floral bract comprising /^-^(Vi) of total
inflorescence length; inflorescence bracts all tubulose; flowers dull green tinged with
brown or maroon . C. lehmannii
Cryptocentrum calcaratum (Schltr.) Schltr., Or-
chideen 449. 1914. Pittierella calcarata Schltr.,
Repert. Spec. Nov. Regni Veg. 3:80-81. 1907.
TYPE: Costa Rica, Province of Alajuela, La
Palma, A. Tonduz 9682 (holotype: B, destroyed;
drawing at AMES). Figure 1C.
Epiphytic herb (5)8-20(28) cm tall, monopodial, rarely
branching from the base; stems (1)2-7(10) cm long, usually
erect, distichously 4-8-leaved at the apical Vz, basal Vz cov-
ered with defibrating leaf sheaths and decurrent roots that
give it a pseudobulbous appearance. Leaves coriaceous, flat
or somewhat convex or concave, straight or arcuate, mid-
nerve shallowly sulcate on ventral surface, slightly keeled
on dorsal surface especially toward apex, (4)5-15(20) cm
long, (4)6-11(12) mm wide; sheath 1-2 cm wide; blades
oblong-elliptic to narrowly oblong-elliptic, apex obtuse to
subacute, asymmetrically 2-lobulate; margins and keel
erose. Inflorescence (2)3-6 cm long, much shorter than
the leaves, 1-3(5) appearing simultaneously from the root-
ing zone, erect, sheathed with inflated and dorsally sharply
keeled bracts; ovary with pedicel smooth or somewhat ver-
ruculose, (26)30-36(38) mm long, subterete, straight, sub-
tended by a subequal floral bract. Flowers spreading,
greenish or yellowish green, the segments rarely suffused
with purple or brownish apically. Sepals subfleshy, flat or
with margins slightly involute, obtuse to rarely acute; dorsal
narrowly triangular, narrowly oblong-elliptic to narrowly el-
liptic, 9-15 mm long, (3.5)4-5(6) mm wide; lateral sepals
narrowly lanceolate to narrowly elliptic-lanceolate, 12-16
mm long, 3.5-5 mm wide; tube 4-5 mm long, 2.5-3 mm
thick, fusiform to broadly fusiform-obconic; sepaline spur
22-32 mm long, cylindric, straight. Petals narrowly trullate
to trullate-elliptic, (9)10-15(17) mm long, 2.5-4 mm wide,
the basal section contained in the sepaline tube, concave;
bent portion 5-11 mm long starting ca. 2 mm above the
mouth of the sepaline tube, 2.5-3.5 mm wide at base, apex
obtuse to subacute, margins flat or somewhat revolute. Lip
laxly covered with long white or greenish hairs in some
clones, sigmoid in profile, 6-9 mm long, 4.5-5 mm wide
when spread; hypochile broadly elliptic to suborbicular
when expanded, 4-5 mm long, 2-3 mm deep; epichile
fleshier than hypochile, conduplicate-concave, lanceolate to
lanceolate-triangular, 3-4 mm long, 1 .5-2 mm wide, obtuse
to subacute. Column stout, 2.5-3.5 mm long, flanked by
dolabriform wings for % of its length, anterior margins of
the wings projected forward; anther ca. 2 mm long, ven-
trally with a pair of retrorse teeth; pollinia 4, ellipsoid. Cap-
sule ellipsoid or oblongoid, 2.5-3.2 cm long.
Apparently restricted to open places at the for-
est edges or high on trees within cloud forests at
1000-2400 m. Flowering mostly July to October.
Costa Rica and adjacent Panama. Frequent on
pasture trees at the Monteverde Reserve in the
Cordillera de Tilaran but rather rare in southeast-
ern Costa Rica and Panama.
Cryptocentrum flavum Schltr., Repert. Spec.
Nov. Regni Veg. 12:214. 1913. TYPE: Colom-
bia, Cauca, Cordoba, Dagua Valley, Pacific
coastal zone, 30-100 m, H. Pittier 591 (holo-
type: us).
Plant an epiphytic herb, erect or ascendent, mono-
podial, often branching with age, 4-8 cm tall. Stem ba-
sally enveloped by decurrent roots, leaf arrangement
polystichous. Leaves straight or slightly recurved, usu-
ally speckled with dull red-maroon, linear or linear-ob-
long, hemicylindric or subtriquetrous in cross section,
obscurely keeled beneath, (30)40-60(70) mm long, 1 .5-
2.5 mm wide; apex obtuse to acutely obtuse, microscop-
ically mucronulate; margins and keel microscopically
erose. Inflorescence erect, spreading, or descending,
subequaling or slightly longer than the subtending
leaves, 1-2(4) appearing simultaneously from the lower
stem, (1.5)3-6(7) cm long, bracts much shorter than the
internodes, tubulose; ovary with pedicel green tinged
with red-maroon, subterete, (8)10-14.5(16) mm long,
subtended by a floral bract 5-7 mm long and enclosing
only the apical ]/lo-V5 of the spur. Flowers greenish or
dull ocher-yellow, often tinged with dull red-maroon to-
ward the bases of the perianth segments, subcampanu-
late or with perianth segments spreading to an angle of
45-80°. Sepals somewhat fleshy, convex, often with re-
curved margins; dorsal narrowly lanceolate to lanceolate
or oblong-lanceolate, 5-7 mm long, 1.3-2.1 mm wide,
obtuse to obtusely acute; lateral sepals narrowly lanceo-
late to lanceolate, 5-6 mm long, 1.3-2 mm wide, obtuse
or acute; tube broadly obconic, broadly 3-keeled, 1-1.5
mm thick; sepaline spur cylindric, broadly and obscurely
clavate, straight, rarely somewhat curved, 12-14.5 mm
long. Petals narrowly lanceolate to lanceolate, 6-8.5
mm long, 1.5-2 mm wide; basal section shallowly con-
cave and projecting well beyond the sepaline cup for
1.5-2 mm; bent portion fleshier than the basal portion,
bending angle 35-45°, 3-5 mm long, 1-1.3 mm wide at
base, apex acute and acuminate. Lip sigmoid in profile,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
29
6-7.5 mm long, 2.5-2.7 mm wide when spread; hypo-
chile shallowly saccate, elliptic when expanded, 4 mm
long, 1-1.2 mm deep; epichile straight or slightly down-
flexed, bending angle 0-45°, narrowly triangular to tri-
angular-lanceolate, 3.2-4 mm long, 1.5-1.8 mm wide at
base, acuminate. Column dorsally flat or convex, ven-
trally flanked by dolabriform wings for V4-% its length,
2.4-2.6 mm long, 1.3-1.5 mm thick; anther ca. 1 mm
long and tall, ventrally with an obscure pair of teeth,
0.2-0.3 mm long. Capsule 1.5-2 cm long.
Epiphytic in tropical wet to cloud forests at 0-
1650 m. Flowering in Costa Rica little known;
elsewhere apparently throughout the year. Ex-
treme southeastern Costa Rica to Ecuador.
Cryptocentrwn flavum and closely related C.
standleyi are easily recognized by the small stat-
ure of the plants but are difficult to distinguish
from each other, especially in dried specimens.
Plants of C. flavum usually have longer leaves that
are also thicker, straighter, and stiffen The stems
of C. flavum are always shorter than those of C.
standleyi for an equivalent number of internodes.
The spur of C. flavum is 12-15 mm long but rare-
ly as short in C. standleyi.
Cryptocentrum gracillimum Ames & C.
Schweinf., Sched. Orch. 8:68. 1925. TYPE:
Costa Rica, Cartago, Pejivalle, 2300 ft, 16 May
1924, Lankester 866 (holotype: AMES).
Plant an epiphytic herb, erect, 7-15 cm tall, mono-
podial but older plants producing 1-many lateral offsets
and then plants densely cespitose. Stems enveloped by
decurrent roots basally, apically by leaf sheaths. Leaves
hemicylindric, linear, (4)5.5-10(15) cm long, 2-2.5 mm
thick, adaxially shallowly concave, abaxially convex
with no raised midvein; apex obtuse (acute in dried spec-
imens), unequally 2-lobate; margins erose; sheaths 5-12
mm long, ca. 5 mm wide, multinerved. Inflorescences
appearing simultaneously from the rooting zone, erect or
horizontally spreading, 4-9 cm long; peduncle terete but
the last internode obscurely compressed, sheathed with
conduplicate dorsally keeled bracts; ovary with pedicel
17-21 mm long, laxly covered with microscopic glan-
dular hairs, subtended by a floral bract 19-24 mm long,
spathaceous and inflated, open only at the apex, enclos-
ing the spur and covering the floral tube partially or
totally. Flowers with widely spreading segments, pale
greenish yellow, green, dull olivaceous, or dull tan-
brown. Sepals subfleshy, oblong-elliptic, obtuse, mar-
gins revolute; dorsal 8-10 mm long, 2.5-3.1 mm wide;
laterals 8-10 mm long, 2.3-2.8 mm wide; sepaline cup
fusiform-obconic, 2-3 mm long, 1.9-2.5 mm thick; se-
paline spur cylindric, only obscurely clavate, 13.5-19
mm long. Petals narrowly lanceolate, acute, long acu-
minate, 8.3-10 mm long, 1.9-2.1 mm wide; bent portion
4-5 mm long, starting well above the mouth of the se-
paline cup, bending angle 45-90°. Lip sigmoid in pro-
file, 7-8 mm long, 2.8-3.2 mm wide when expanded,
throat with a few white hairs; hypochile broadly elliptic
to ovate, with 3 thick nerves and 2 thinner outer ones,
margins straight; epichile narrowly lanceolate, 3-nerved,
down-flexed, bending angle 45-90°, 4-5 mm long. Col-
umn arcuate, 2.1-3 mm long, ca. 1.5 mm thick, flanked
by dolabriform wings for % its length; anther 1-1.2 mm
long, 1.2-1.5 mm tall, ventrally with a pair of retrorse
teeth; pollinia ellipsoid, dull cream-yellow.
A rare epiphyte within its range growing in ei-
ther bright or shady conditions at 500-1000 m.
Flowering May to December. Endemic to the
Central Valley of Costa Rica; replaced in Panama
by an undescribed species.
The hemiterete leaves on an abbreviated di-
stichous-leaved stem are good field characters for
recognition.
Cryptocentrum latifolium Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:247. 1923. TYPE:
Costa Rica, Alajuela, Las Angeles de San Ra-
mon, A. Brenes 56 (holotype: B, destroyed; lec-
totype: drawing at AMES). Figure IB.
Plant an epiphytic herb, occasionally growing on road
banks or rocks, monopodial, erect or horizontally
spreading, rarely producing basal offshoots, (7)12-
20(30) cm tall. Stems covered basally by decurrent
roots, apically by leaf sheaths. Leaves distichous, con-
duplicate, fleshy-coriaceous, linear-oblong to narrowly
oblong-obovate, erect to arcuate, fiat to adaxially con-
vex, 5-20(30) cm long, (8)10-16(20) mm wide; apex
obtuse to subrounded, strongly unequally 2-lobate, ob-
scurely keeled, margins erose, especially toward apex.
Inflorescences to as long as the leaves, usually shorter,
(4)6-13(20) cm long, appearing simultaneously from the
rooting zone, erect or horizontally spreading or pendent;
peduncle stout, bracts subequal to the internodes; ovary
with pedicel 28-35 mm long, subterete, laxly to sub-
densely covered with glandular hairs; subtended by a
floral bract 30-42 mm long, somewhat inflated, totally
enclosing the ovary and spur and occasionally the base
of the dorsal sepal. Flowers among the largest in the
genus with widely spreading perianth segments, greenish
brown or yellow-green, often heavily tinged with ma-
roon, or totally maroon. Sepals fleshy, obtuse to acute,
subapically mucronulate to conspicuously mucronate,
margins revolute, especially the laterals; dorsal lanceo-
late to elliptic-lanceolate, 15-20 mm long, 5-8 mm
wide; lateral sepals oblong to narrowly oblong-elliptic,
18-30 mm long, 3.5-5.5 mm wide; tube cylindric to
broadly cylindric-obconic, 2-3 mm long; sepaline spur
cylindric, not clavate or obscurely so, 26-30 mm long.
Petals elliptic to broadly elliptic, 10-16 mm long, 4-
5.5 mm wide, the basal section included in the sepaline
cup, concave; bent portion 5-7.5 mm long, 3-5.5 mm
wide at the base, conspicuously fleshier than basal por-
tion, bending angle 90-180°. Lip sigmoid in profile, 8-
12 mm long, 4.5-6 mm wide when expanded; hypochile
broadly elliptic to rounded when expanded, 4.5-5.5 mm
long, 2.5-3 mm deep, margins straight; epichile trian-
gular to broadly ovate, 2.5-3.5 mm long. Column 3-4
mm long, ca. 2 mm thick, dorsally slightly arcuate.
30
FIELDIANA: BOTANY
flanked by dolabriform wings for % its length; anther
1.8-2.2 mm long, ventrally with a pair of retrorse teeth.
Capsule 30-40 mm long, ellipsoid.
Epiphytic on larger branches in very humid to
moist forests at 0-1500 m. Flowering June to Oc-
tober. Costa Rica to Ecuador.
Cryptocentrum latifolium is the largest species
in the genus and has the largest flowers, although
small plants start flowering when they could be
confused with other species. The broad, very un-
equally 2-lobate leaf apex is unique, with the lon-
ger lobe exceeding the shorter by (2)3-5(12) mm.
Cryptocentrum lehmannii (Rchb. f.) Garay, Bot.
Mus. Leafl. 18:209. 1958. Aeranthus (Cryptoplec-
tri) lehmannii Rchb. f., Otia Bot. Hamb. 10. 1878.
TYPE: Ecuador, Tungurahua, Volcan Tungurahua,
Banos, 7400 ft (2320 m), Mar. 1877, F. Lehmann
601 (w). Campylocentrwn lehmannii (Rchb. f.)
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 8:
164. 1921. (sphalm). Cryptocentrum jamesonii
Benth., Gen. PI. 557. 1883. TYPE: Ecuador, Pi-
chincha, Jameson s.n. (holotype: K). Cryptocen-
trum gracilipes Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19:246. 1923. TYPE: Costa Rica, Al-
ajuela, Los Angeles de San Ramon, Brenes 37
(holotype: B, destroyed). Figure ID.
Plant an epiphytic monopodial herb (8)12-25(30) cm
tall, rarely branching from the base. Stems erect, basal l/i
covered with filiferous leaf sheaths and decurrent roots, dis-
tichously 5-8-leaved apically. Leaves coriaceous to some-
what fleshy, flat or somewhat convex or concave, arcuate
or stiffly erect, linear to linear-elliptic, (5)10-20(25) cm
long, (1.5)3-9(12) mm wide; apex obtuse to acute, un-
equally 2-lobate. Inflorescences rarely exceeding the
leaves, 1-4(6) appearing simultaneously from the rooting
zone, arcuate or horizontally spreading to erect, (4)5.5-
12(15) cm long; peduncle with bracts shorter to somewhat
longer than the intemodes; ovary with pedicel (15)22-30
mm long, subtended by a floral bract reaching and enclos-
ing the dorsal section of the tube. Flowers with perianth
segments opening to an angle of 70-90°, dark green with
dark reddish or purplish tinge. Sepals somewhat fleshy,
lanceolate or oblong-lanceolate, acute to subobtuse, with
margins usually revolute; dorsal 8-15(20) mm long, 3-6
mm wide; lateral sepals often somewhat oblique, (10)12-
20(22) mm long, (3)4-6 mm wide; sepaline cup broadly
obconic, obscurely keeled, 2-3(3.5) mm long and thick;
sepaline spur cylindric to strongly clavate, straight, (10)15-
27 mm long. Petals trullate-elliptic, (8)12-16(18) mm long,
the basal section contained in the sepaline tube, concave;
bent portion 5-9 mm long, margins somewhat or strongly
revolute, bending angle 45-90°. Lip sigmoid in profile, 6-
10(13) mm long, 4-6 mm wide when spread; hypochile
broadly elliptic to almost orbicular when expanded, 4.5-6
mm long, 2.6-3 mm deep; epichile conduplicate-concave
or flat, lanceolate to lanceolate-triangular, 4-7 mm long, 2-
2.5 mm wide, obtuse to acute, bending angle 10-180°. Col-
umn 2.5-4 mm long, 1.5-1.8 mm thick, flanked by dolab-
riform wings for % its length, anterior wing margins pro-
jected forward; anther 2 mm long, with a pair of retrorse
teeth.
Cloud forests mostly at 1200-2000 m. Flowering
mostly December to July. Costa Rica to Ecuador.
In Costa Rica, C. lehmannii can be easily dis-
tinguished from C. calcaratum by the inflores-
cences with (5)6-12 internodes as opposed to 3-
4(5) internodes in the latter species. It has smaller
flowers than C. latifolium.
Cryptocentrum standleyi Ames, Sched. Orch. 9:
55-56. 1925. TYPE: Costa Rica, San Jose, La
Palma, ca. 1600 m, 3 Feb. 1924, P. C. Standley
33028 (holotype: AMES). Cryptocentrum longis-
capum Brieger, Bot. Jahrb. Syst. 97:568. 1977.
TYPE: Colombia, Coteje on Rio Timbiqui, 200-
500 m, Lehmann 8961 (holotype, K). Cryptocen-
trum brenesii Schltr., nom. Figure 1A.
Plant an epiphytic herb, erect or ascendent, monopodial,
branching with age, (2)4-10(20) cm tall. Stems basally en-
veloped by decurrent roots; apical ¥2 polystichous with
many leaves. Leaves often recurved, linear or linear-obo-
vate to narrowly obovate (small leaves), hemicylindric to
triquetrous in cross section, obscurely keeled below es-
pecially apically, (10)15-25(30) mm long, apex obtuse,
mucronulate, margins erose. Inflorescences appearing si-
multaneously from the lower- or midstem intemodes, erect
but peduncle frequently curving, with 3-4 intemodes,
(2.5)3.5-6(7.5) mm long; ovary with pedicel subterete,
whitish or pinkish, or tinged with red-maroon, (14)18-22
mm long, subtended by a floral bract 5-9 mm long. Flow-
ers yellow-green or yellow-orange, often heavily tinged
with several shades of maroon, or entirely maroon to dull
wine red, subcampanulate or with perianth segments
spreading to an angle of 45-90°. Sepals subfleshy, oblong
to oblong-lanceolate, subapically mucronulate, convex, of-
ten with revolute margins; dorsal 6.5-8 mm long, 1.5-2
mm wide, obtuse to obtusely acute; lateral sepals 7-10 mm
long, 1 .8-2 mm wide, acute; tube broadly obconic, dorsally
convex, ventrally convex, broadly 3-keeled, 1-1.3 mm
long, ca. 2 mm thick; sepaline spur cylindric, broadly and
obscurely clavate, straight or curved, (15)16-19 mm long.
Petals narrowly elliptic, 7.5-8 mm long, 2.1-2.2 mm wide;
basal section concave and projecting well beyond the se-
paline tube; bent portion fleshier than the basal portion,
bending angle 35-45°, 3.2-4.2 mm long, 1.8-2 mm wide
at base. Lip sigmoid in profile, 6-6.5 mm long, 2-2.5 mm
wide when expanded; hypochile deeply saccate, when ex-
panded broadly elliptic, 1.5-2 mm long, 1.2-1.5 mm deep;
margins entire; epichile narrowly triangular-lanceolate,
straight or slightly down-flexed, bending angle 0-45°, 3-4
mm long, 1.2-1.5 mm wide at base, acuminate. Column
2-2.5 mm long, 1 .8-2 mm thick, ventrally flanked by do-
labriform wings for % its length; anther 0.8-1 mm long and
tall, ventrally with an obscure pair of teeth; pollinia irreg-
ularly ellipsoid, supported on an oblong stipe and amor-
phous viscidium.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
BKX08Y LI8RAKY
« 9DGO
31
Cryptocentrum standleyi grows both on twigs
and on larger branches over a range of light ex-
posures in rain and cloud forests from 200-2050
m. Sporadically continuously blooming, but most-
ly from June through December during the rainy
season. Costa Rica to Ecuador.
This species and C. flavum are similarly small
species but difficult to distinguish from one an-
other. Cryptocentrum standleyi usually has longer
inflorescences and longer spurs and has shorter
and thinner leaves.
Maxillaria Ruiz & Pavon
(J. T. Atwood)
REFERENCES — J. T. Atwood, Orchids of Costa
Rica 1. Icones Plantarum Tropicarum 14: plates
1301-1400. 1989. J. T. Atwood, A revision of the
Maxillaria neglecta complex (Orchidaceae) in
Mesoamerica. Lindleyana 8:25-31. 1993. J. T. At-
wood, Two new species of Maxillaria (Orchida-
ceae) from southern Central America and a new
name for a well-known taxon. Lindleyana 9:227-
231. 1994. J. T. Atwood, Two new Costa Rican
species of Maxillaria (Orchidaceae) related to
Maxillaria flava and Maxillaria microphyton.
Lindleyana 9:232-237. 1994. J. T. Atwood, Two
new species of Costa Rican Maxillaria (Orchida-
ceae), and a clarification of Maxillaria bracteata.
Lindleyana 9:239-242. 1994. J. T. Atwood, Two
overlooked species of Maxillaria from Central
America. Selbyana 16:242-245. 1995. G. Carne-
vali and J. T. Atwood, Orchidaceae Dunstervillo-
rum II. Maxillaria cryptobulbon sp. nov. and a
note on the Maxillaria brunnea complex. Novon
1:159-164. 1991. G. Carnevali and J. T. Atwood,
Two new species in the Maxillaria rufescens com-
plex from Central America. Lindleyana 11:27-33.
1996. K. Senghas, Subtribus: Maxillariinae. In R.
Schlechter. Die Orchideen 29:1727-1803. 1994.
Epiphytic herb cespitose or rhizomatous, sometimes
cespitose as juveniles but rhizomatous at maturity; rhi-
zomes often aerial, with short to elongate, often branch-
ing stems, with or without apically 1-4-leaved pseudo-
bulbs. Pseudobulbs always terminating the shoot when
present, less often absent, and forming indeterminate
shoots. Leaves conduplicate, usually ligulate and cori-
aceous, rarely fleshy. Inflorescences 1-many, apparently
reduced to 1 -several elongate 1 -flowered peduncles per
bract (rhizome scale) at the pseudobulb base; ovary usu-
ally without pedicel. Flowers campanulate to spreading,
of various colors, sometimes sweetly or offensively
odoriferous. Sepals subequal, the lateral sepals some-
what adnate to the column foot and forming a chin at
the base, rarely connate. Petals similar to the sepals but
usually smaller, the lip concave, 3-lobate or entire, the
disk usually with a fleshy or mealy, less often oily or
waxy callus. Column semiterete, rarely winged, and
usually with a conspicuous foot; pollinia 4, waxy.
A Neotropical genus of at least 450 species;
107 species in Costa Rica. The genus is best rec-
ognized by the combination of conduplicate
leaves, apparently one-flowered scapose inflores-
cences, the presence of a column foot, four pol-
linia supported on a short to long stipe, and a
distinct, usually horseshoe- shaped viscidium.
The relationships of Maxillaria are not clear.
We suggest that certain Brazilian species of Max-
illaria (e.g., M. picta Hook.) are more closely re-
lated to Trigonidium than to other species of Max-
illaria. Also, Mormolyca ringens (Lindl.) Schltr.
exhibits rugose leaves with raised veins, pedicel-
late ovaries, and a lip shape similar to that of the
Maxillaria rufescens Lindl. complex. One species
in the Costa Rican flora, Maxillaria tigrina C.
Schv/einf., has the vegetative and floral aspect of
the otherwise South American genus, Chrysocyc-
nis, but the pollinia are odd for that genus. Clear-
ly, much needs to be done to improve generic re-
alignment in the Maxillariinae.
Keys to the Species of Maxillaria
la. Plants always with elongate stems that lack pseudobulbs; pseudobulbs if present restricted to base
of plant (e.g., Figs. 2B-5D) 2
2a. Leaf blades > 30 cm, < 1 cm wide; plant pendent M. lueri
2b. Blades if 30 cm long then much wider; plant various 3
3a. Leaf axils never with > 1 scape 4
4a. Sepals > 1 .6 cm long, at least some leaves > 1 .5 cm wide 5
5a. Sepals < 2 cm long M. alfaroi
5b. Sepals > 2.5 cm long M. inaudita
4b. Sepals < 1 .5 cm long; leaves < 1 .3 cm wide 6
32
FIELDIANA: BOTANY
6a. Lip spatulate, midlobe > 2 times wider than isthmus M. valerioi
6b. Lip not as above 7
7a. Ovary > 2.5 cm long M. linearifolia
7b. Ovary < 1.5 cm long 8
8a. Leaves elliptic, thin; plants pendent M. appendiculoides
8b. Leaves lanceolate, chartaceous; plants usually stiffly erect
M. dendrobioides
3b. At least some leaf axils with > 1 scape 9
9a. Column (excluding anther) < 4 mm long 10
lOa. Ratio: foot length/column length < 0.2 11
1 la. Leaves > 3 cm wide M. trilobata
1 Ib. Leaves < 2 cm wide 12
12a. Leaves acute; column about 2 mm long M. adendrobium
12b. Leaves obtuse and unequally 2-lobate; column about 4 mm long
M. nicaraguensis
lOb. Ratio: foot length/column length > 0.2 13
13a. Sepals > 1.1 cm long 14
14a. Lip forming a sharp angle with column foot and usually readily
movable 15
15a. Leaves lanceolate, sessile, < 2 cm wide 16
16a. Sepals and petals yellow, column < 2.5 mm long ex-
cluding anther and foot M. parvilabia
16b. Sepals and petals brick red to brown or green, column
> 4 mm long M. tonduzii
15b. Leaves elliptic, long petiolate, > 3 cm wide . . . M. trilobata
14b. Lip continuous with column foot, more or less rigid, not forming
a sharp angle 17
17a. Lip with simple callus at base lacking adornments
M. biolleyi
17b. Lip with keels, papillae, or warts at base in addition to
fleshy callus 18
18a. Tepals greenish, reddish, or brownish . . M. tonduzii
18b. Tepals lavender to white 19
19a. Lip subsaccate, ovary ^ 1 1 mm long
M. adolphii
19b. Lip not saccate, ovary > 11 mm long
M. quadrata
13b. Sepals < 1 cm long 20
20a. Lip with a pair of rounded lobes at base M. conduplicata
20b. Lip lacking side lobes 21
2 la. Flowers blood red and yellow M. fulgens
21b. Flowers mauve to whitish or lavender M. pittieri
9b. The same > 4.5 mm long 22
22a. Ratio: foot length/column (less anther) length > 0.3; stem branches some-
what swollen at ends 23
23a. Ratio: midlobe (suborbicular)Tlip ^ 1A M. exaltata
23b. Ratio: midlobe (lance-ovate)/lip ^ Vz M. meridensis
22b. Ratio: foot length/column (less anther) length < 0.25, stem branches never
swollen at apex 24
24a. Column > 7 mm long 25
25a. Column (without anther) 7.5-8 mm long M. falcata
25b. Column 9-10 mm long (without anther), lip keeled on callus. . .
M. amabilis
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS 33
25c. Column 13-15 mm long (without anther), lip not keeled (very
shortly so in plants from Panama) M. monteverdensis
24b. Column < 6 mm long 26
26a. Leaves < 1.5 cm wide M. nicaraguensis
26b. Leaves > 3.5 cm wide M. trilobata
Ib. Plants various but elongate stems (if present) with pseudobulbs 27
27a. Plants with apically > 2-foliate pseudobulbs 28
28a. Plants with upright, decumbent or pendent, aerial canes 29
29a. Largest leaves 6 cm long; sepals < 12 mm long 30
30a. Flowers dark purple M. ponerantha
30b. Flowers greenish to white, lip with red stain M. acervata
29b. Largest leaves > 10 cm long; sepals > 12 mm long 31
3 la. Flowers red to orange M. elatior
31b. Flowers white to yellow 32
32a. Sepals and petals long-attenuate, ^ 4 mm wide . . . M . ctenostachya
32b. Sepals and petals oblong-elliptic, > 5 mm wide M. camaridii
28b. Plants often creeping, but never with aerial canes 33
33a. Column 9-1 1 mm long M. friedrichsthalii
33b. Column 12-14 mm long M. scorpioidea
33c. Column > 1.5 cm long M. aciantha
27b. Pseudobulbs apically 1 -foliate or pseudobulbs lacking altogether 34
34a. Plants with distinct rhizomes or canes separating pseudobulbs; usually some rhizomes
aerial Key 1
34b. Plants distinctly cespitose, lacking elongate rhizomes or canes Key 2
Key 1: Rhizomes or Canes with Pseudobulbs, Some Aerial
la. Lateral sepals partially united beyond column foot (e.g., Figs. 9B-10B) 2
2a. Leaves distinctly elliptic and obtuse; inflorescences usually forming all along most recently
matured rhizome, not just at pseudobulb base M. brevilabia
2b. Leaves lanceolate or linear, always acute; inflorescences fasciculate only at pseudobulb base
3
3a. Lateral lobes of lip suborbicular when viewed from the side M. concavilabia
3b. Lateral lobes otherwise; subquadrate, elliptic or nearly absent 4
4a. Lateral lobes of lip reduced to auricles at base of midlobe, base forming a distinct
claw 5
5a. Flowers orange to red; column foot longer than column M. horichii
5b. Flowers white or off-white, lip with yellow midlobe; column foot equal to
or shorter than column M. parviflora
4b. Lateral lobes of lip conspicuous and extending to column foot, base not forming
a distinct claw 6
6a. Lip when spread 3-4 mm across; flowers always white, lip white with yellow
midlobe; habitat below 1200 m M. neglecta
6b. Lip when spread (4.5) 5-7 mm across; flowers various colors; habitat above
900 m M. pseudoneglecta
Ib. Lateral sepals not united beyond column foot 7
7a. Elongate stem apex somewhat swollen, but rarely forming a distinct pseudobulb, which, if
present, is concealed by leaf sheaths 8
8a. Sepals and petals > 1.5 cm 9
9a. Ratio: midlobe (suborbicular)Tlip < % M. exaltata
9b. Ratio: midlobe (lance-ovate)/lip ^ Vz M. meridensis
8b. Sepals < 1.1 cm long 10
34 FIELDIANA: BOTANY
lOa. Lip with pair of rounded lobes at base M. conduplicata
lOb. Lip lacking side lobes 11
1 la. Flowers blood red and yellow M. fulgens
lib. Flowers mauve to whitish or lavender M. pittieri
7b. Stem apex clearly modified into pseudobulb (M. schistotele, known only from type photos
and original description, cannot be taken further) 12
12a. Sheaths subtending pseudobulbs lacking blades or articulations 13
13a. Pseudobulbs < 1.4 cm long; column foot about as long as column; leaves subterete
M. uncata
13b. Pseudobulbs > 1.5 cm long; foot much shorter than column; leaves never subterete
14
14a. Sepals > 1.7 cm long 15
15a. Leaves < 9 mm wide; flowers basically red, the lip simple, not at all
3-lobate 16
16a. Lip apex white spotted with reddish maroon M. tenuifolia
16b. Lip apex whitish, suffused with red or rose .... M. sanguined
15b. Leaves > 1 cm wide; flowers yellow or striped, not red, lip clearly 3-
lobate 17
17a. Flowers yellow; sepals > 3 cm long M. planicola
17b. Flowers translucent, striped with maroon; sepals < 2.9 cm long
M. tigrina
14b. Sepals < 1.5 cm long 18
18a. Sepals < 8 mm long M. caespitifica
18b. Sepals > 8 mm long 19
19a. Flowers orange-yellow with red on lip M. variabilis
19b. Flowers green stained with red, especially lip . . M. costaricen-
sis
12b. Sheath(s) at base of pseudobulb foliaceous or at least with articulation 20
20a. Column without anther > 1.7 cm long M. haberi
20b. Column without anther < 1.7 cm long 21
2 la. Petiole of foliaceous bracts abruptly narrowed above articulation and/or leaf
< 2 mm wide 22
22a. Leaf apex rounded to emarginate; neither acute nor acuminate, or if
otherwise leaf < 2 mm wide (see Figs. 12A-13B) 23
23a. Leaves terete, < 2 mm in diameter M. muscoides
23b. Leaves not terete, > 2 mm wide 24
24a. Midlobe > % length of lip 25
25a. Pseudobulbs elongate, ^ 2 times longer than wide . .
26
26a. Pseudobulbs distinctly grooved or fluted; ratio:
lip length/sepal length < % M. flava
26b. Pseudobulbs not grooved; ratio: lip length/sepal
length > 3/5 27
27a. Sepals > 1.1 cm long .... M. lankesteri
27b. Sepals < 1 cm long M. wercklei
25b. Pseudobulbs < 2 times longer than wide
M. burgeri
24b. Midlobe < % length of lip M. microphyton
22b. Leaf apex acute to acuminate, rarely obtuse and shortly acuminate . . .
28
28a. Column foot > Vi column length less anther, clearly differentiated
from base of lip M. tubercularis
28b. Column foot < Vi column length, often poorly differentiated from
base of lip 29
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS 35
29a. Column less anther > 6 mm long 30
30a. Column less anther < 1 1 mm long . . M. sigmoidea
30b. Column less anther > 13 mm long . . . M. vaginalis
29b. Column less anther < 6 mm long 31
3 la. Column less anther < 2 mm long M. minor
31b. Column less anther > 2.5 mm long 32
32a. Flowers purple to reddish M. paleata
32b. Flowers pink, lavender, white, or yellowish . . 33
33a. Lower quarter of lip subsaccate; midlobe <
Vi lip; flowers light lavender to white ....
M. schlechteriana
33b. Lower quarter of lip not at all saccate; mid-
lobe > Vz lip; flowers yellowish to pinkish
M. dichotoma
21b. Petiole not abruptly narrowed above articulation; leaves ^ 2 mm wide . . 34
34a. Blades of rhizome sheaths subtending flowers appearing half-lyre-
shaped from side (e.g., Figs. 14D-15B) 35
35a. Sepals and petals pure white 36
36a. All leaves ^1.8 cm wide M. camaridii
36b. Some, if not all, leaves ^ 1.8 cm wide . . . M. ampliflora
35b. Sepals and petals otherwise 37
37a. Column < 6 mm long M. umbratilis
37b. Column > 6 mm long 38
38a. Lip when spread wider than long 39
39a. Sepals > 3.4 cm long M. ampliflora
39b. Sepals < 3.3 cm long M. bradeorum
38b. Lip when spread narrower than long
M. campanulata
34b. Sheaths various but not appearing half-lyre-shaped from side .... 40
40a. Sepals > 1 .7 cm long 41
41a. Leaves < 1 cm wide M. oreocharis
41b. Leaves > 1.1 cm wide, or if narrower then obtuse ... 42
42a. Plant with distinctly foliaceous bracts subtending
pseudobulb 43
43a. Columns less anther < 6 mm long 44
44a. Ovary trigonous; fruit 3-alate
M. gomeziana
44b. Ovary terete, fruit not 3-alate
M. bracteata
43b. Column less anther > 6 mm long 45
45 a. Midlobe of lip suborbicular, serrulate ....
M. serrulata
45b. Midlobe of lip otherwise 46
46a. Flowers white with yellow lip . . 47
47a. Leaves < 2 cm wide
M. camaridii
47b. Leaves > 2.5 cm wide
M. suaveolens
46b. Rowers orange to red or purple . . 48
48a. Ovary with pedicel > 3 cm long
M. elatior
48b. Ovary with pedicel < 2 cm long
M. tubercularis
42b. Bract at base of pseudobulb usually with articulation
36 FIELDIANA: BOTANY
but rarely with a distinct broadened blade; lip not ser-
rulate 49
49a. Flowers clearly white with yellow on lip
M. alba
49b. Flowers green, whitish, or yellowish
M. anceps
40b. Sepals < 1.6 cm long 50
50a. Youngest rhizome concealed by several foliaceous sheaths
with fully developed blades 51
5 la. Flowers dark purple M. ponerantha
51b. Flowers light green with red stain on lip
M. acervata
50a. Youngest rhizome concealed by 0-3 blade-bearing sheaths,
rest non-blade-bearing 52
52a. Leaves never obtuse and deeply emarginate, larger >
7 cm long 53
53a. Flowers red with yellow lip M. fulgens
53b. Flowers otherwise 54
54a. Leaves > 1.2 cm wide, scapes appearing
precociously on new shoots M. diuturna
54b. Leaves < 1 . 1 cm wide, scapes appearing at
base of mature shoots .... A/, oreocharis
52b. Leaves obtuse and deeply emarginate, < 6 cm long,
elliptic 55
55a. Sepals > 1 1 mm long M. lankesteri
55b. Sepals < 9 mm long M. wercklei
Key 2: No Conspicuous Rhizomes or Canes
la. Shoots with ^ 3 leaves arranged in fan, or if pseudobulbs present, always with ^ 3 subtending,
foliaceous sheaths, leaves never broadly elliptic with petiole ^ 3 mm broad (see Figs. 17B-18D)
2
2a. Plants with conspicuous pseudobulbs 3
3a. Sepals < 2.2 cm long M. crassifolia
3b. Sepals 2.3-2.6 cm long M. maleolens
3c. Sepals > 3 cm long M. nasuta
2b. Plants lacking pseudobulbs, or these hidden by foliaceous sheaths 4
4a. Plant pendent, psygmoid, handlike; leaves equitant, iris-like M. valenzuelana
4b. Plant and leaf otherwise 5
5a. Column foot longer than column M. angustissima
5b. Column foot shorter than column 6
6a. Leaves not abruptly narrowed above the articulation, thick, obtuse
M. crassifolia
6b. Leaves abruptly narrowed above articulation, thin, distinctly acute 7
7a. Leaves ^1.3 cm wide with unspotted sheaths, flowers wine red ....
M. bicallosa
7b. Leaves > 1.7 cm wide with spotted sheaths, flowers yellow
M. chartacifolia
Ib. Shoots always with pseudobulbs, rarely with 3 foliaceous sheaths at base 8
8a. No broadened, foliaceous sheaths subtending pseudobulbs, although articulations sometimes
present (rarely in M. alba or M. oreocharis) 9
9a. Scape included within subtending rhizome bract 10
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS 37
lOa. Flowers pure white with yellow on lip M. alba
lOb. Flowers otherwise 11
1 la. Sepals > 1.8 cm long 12
12a. Leaf < 4 mm wide, linear, not petiolate M. sanguinea
12b. Leaf > 1 cm wide, elliptic, long-petiolate M. longipetiolata
1 Ib. Sepals < 1.8 cm long 13
13a. Sepals < 8 mm long M. caespitifica
13b. Sepals > 8 mm long 14
14a. Flowers orange-yellow with red on lip M. variabilis
14b. Flowers green stained with red, especially lip . . M. costaricen-
sis
9b. Scape exerted beyond subtending rhizome bract 15
15a. Leaves < 1.1 cm wide 16
16a. Sepals partially united beyond column foot M. strwnata
16b. Sepals not so united 17
17a. Leaves < 12 cm long M. piestopm
17b. Leaves > 12 cm long 18
18a. Flowers with spreading sepals, column > 1 cm long
M. sanguinea
18b. Flowers campanulate; column < 1 cm long . . . . M. oreocharis
15b. Leaves ^ 1.1 cm wide 19
19a. Petals lanceolate, distinctly broadest on the lower 1A 20
20a. Midlobe of lip fleshy, black, reflexed, and forming an angle of about
90° with base M. longiloba
20b. Midlobe yellow and white, often with purple spots, neither black, nor
so reflexed 21
2 la. Flowers > 2 per rhizome bract; lip < 1.6 cm long M. ringens
21b. Flowers l(-2?) per rhizome bract; lip > 2 cm long 22
22a. Pseudobulbs usually truncate apically; sepals > 5.5 cm long
M. endresii
22b. Pseudobulbs usually conic, not truncate; sepals < 5 cm long
M. angustisegmenta
19b. Petals various, but not broadest on lower 1A 23
23a. Midlobe of lip ^ ¥3 as long as lip 24
24a. Lip apex reflexed black, lance-ovate M. longiloba
24b. Lip apex straight, sometimes with purple spots, not black . . 25
25a. Pseudobulb suborbicular, small, and hidden by subtending
bracts M. cryptobulbon
25b. Pseudobulb ovoid, emerging from subtending bracts
M. brunnea
23b. Midlobe of lip > Yz length of lip 26
26a. Petals distinctly spatulate or oblanceolate M. piestopus
26b. Petals distinctly not spatulate or oblanceolate 27
27a. Inflorescence shorter than pseudobulb 28
28a. Petiole ^ % as long as blade .... M. longipetiolata
28b. Petiole < % as long as blade 29
29a. Sepals and petals maroon to brown
M. dressleriana
29b. Sepals and petals cream-white to yellow ... 30
30a. Lip orange M. hedwigiae
30b. Lip yellow with red stripe up center ....
M. acutifolia
30c. Lip otherwise, usually yellow with red dots
or stained dark M. moralesii
27b. Inflorescence longer than pseudobulb 31
38 FIELDIANA: BOTANY
3 la. Lip orange, midlobe broadly lanceolate
M. hedwigiae
31b. Lip red, purplish, or cream spotted with maroon, mid-
lobe ovate (cucullata group) 32
32a. Flowers with sepals and petals stained with pur-
ple; midlobe of lip with truncate to emarginate
callus M. obscura
32b. Flowers with lines of purple dots on sepals and
petals, lip usually dark red or maroon
M. punctostriata
32c. Flowers with white sepals and petals, lip cream
spotted with maroon M. cedralensis
8b. Pseudobulb subtended by > 1 foliaceous sheath, at least when young 33
33a. Sepals > 10 cm long M. rodrigueziana
33b. Sepals < 10 cm long 34
34a. Foot at least as long as column less anther 35
35a. Sepals < 2.7 cm long 36
36a. Flowers held much higher than shoots; lip twice length of foot plus
column less anther M. attenuata
36b. Flowers much shorter than shoots; lip not much longer than foot plus
column less anther 37
37a. Anther bed lacerate; lateral sepals attenuate, ^1.8 cm long . . .
M. confusa
37b. Anther bed not at all lacerate; lateral sepals not attenuate, ^1.8
cm long M. ramonensis
35b. Sepals > 2.8 cm long 38
38a. Leaves with large, white spots M. reichenheimiana
38b. Leaves lacking such spots 39
39a. Sepals ^5.5 cm long M. arachnitiflora
39b. Sepals < 5 cm long M. pachyacron
34b. Foot shorter than column 40
40a. Lip subentire to indistinctly 3-lobate M. chionantha
40b. Lip distinctly and deeply 3-lobate near the apex 41
41a. Sepals > 1.4 cm long 42
42a. Sepals > 3.5 cm long; flowers generally held higher than the
shoots M. pachyacron
42b. Sepals ^ 3 cm long; flowers much shorter than shoots .... 43
43a. Plants < 12 cm tall, ovary with pedicel < 1.7 cm long . . .
M. brachybulbon
43b. Plants > 25 cm tall, ovary with pedicel > 2.5 cm long . . .
M. cryptobulbon
41b. Sepals < 1.3 cm long 44
44a. Sepals < 6 mm long, column foot nearly absent
M. vittariifolia
44b. Sepals > 8 mm long, column foot nearly as long as column . . .
M. acostae
Maxillaria acervata Rchb. f., Bonplandia 3:217. Plant epiphytic, somewhat straggly, especially in low-
1855. TYPE: Costa Rica, "Ad Suru[n]gnes," er elevation specimens, to 30 cm tall, rooted only at the
Oersted s.n. (lectotype selected here: w; isolec- base;. shoots asce"di"g' 3r9IKcm^Pf"- ,Roots whf' to l
,..„.-., o /-i mm in diameter, rseudobulbs 2(l)-iohate, ovoid, com-
totype: K). Maxillaria fohosa Ames & C. pressed, 1-3 cm long, about 1 cm wide, concealed when
Schweinf., Sched. Orch. 8:60. 1925. TYPE: young by leaf-bearing sheaths. Leaves and leaf-bearing
Costa Rica, C. H. Lankester 478 (holotype: sheaths to more than 20 in developing shoots, all except
AMES). Figure 7D. the apical leaves of the pseudobulbs abscising at matu-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS 39
rity, oblong-lanceolate, 1.5-6 cm long, 5-10 mm wide,
obtuse to unequally 2-lobate. Inflorescence 1 per sheath
axil in the immature shoot; scape about 1 cm long, con-
cealed by few acute sheaths; ovary with pedicel 1.5-2
cm long, exceeding the subtending bract. Flowers erect,
green to white, the lip with red stain on the lower %.
Sepals ovate, elliptic, obtuse, apiculate, 1 cm long, 4
mm wide. Petals elliptic, 9 mm long, 3.5 mm wide. Lip
hinged to the column foot, simple, spatulate, 1 cm long,
4 mm wide; callus oblong, extended to just beyond the
middle, obtuse or rounded apically. Column arcuate, ex-
panded above, 7 mm long; foot 2 mm long; pollinia 4
lacking a stipe but with saddle-shaped viscidium. Cap-
sule ellipsoid, 2 cm long.
Epiphytic in primary montane and premontane
evergreen forests at 700-2000 m. Flowering Sep-
tember to March. Nicaragua to Panama; also re-
ported from Ecuador as M. foliosa.
This species is closely related to M. ponerantha
Rchb. f., but the plant is somewhat larger, and the
flowers are light green with red on the lower half
of the lip. Maxillaria ponerantha is a somewhat
smaller, more widespread, lowland species with
dark purple flowers. The pseudobulbs are nearly
always apically 2-foliate in M. acervata and 1-
foliate in Costa Rican M. ponerantha. Leaf-bear-
ing rhizome bracts of immature shoots in M. fo-
liosa may exceed 20. Lower elevation examples
are much less compact than upper elevation con-
specifics.
Authors usually treat this as the synonym M.
foliosa (e.g., Hamer, Icon. PL Trop 9: Plate 854;
Atwood, Icon. PL Trop 14: Plate 1346). Both
available types of M. acervata agree in the rhi-
zomatous habit, the foliaceous sheaths at the base
of the pseudobulbs, and the rather simple flowers.
Maxillaria aciantha Rchb. f., Bot. Zeitung (Ber-
lin) 10:858. 1852. TYPE: Costa Rica, /. de
Warszewicz s.n. (not seen). Ly caste aciantha
(Rchb. f.) Rchb. f., Bonplandia 3:216. 1855.
Figure 9 A.
Plant a loosely cespitose to straggly epiphyte, rarely
forming large clumps, some of the shoots to 3 cm apart;
often pendent with only basal shoots rooted. Roots fi-
brous, white, to 1 mm in diameter. Pseudobulbs cylin-
dric-fusiform, 3-8 cm tall and 0.7-1.2 cm wide, laterally
flattened and with several vertical ribs conspicuous es-
pecially when dried, subtended by 1-3 early deciduous,
blade-bearing bracts, apically 2-4 foliate. Leaves linear
to elliptic-oblong, 3-15 cm long and 0.8-1.6 cm wide,
unequally 2-lobed apically. Inflorescence lateral, 1-
flowered, borne singly per rhizome bract, the peduncle
about 1.5 cm long covered by 3-5 inflated bracts, with
the uppermost bract subtending the flower 2.0-2.7 cm
long; ovary with pedicel to 2 cm long at anthesis, usu-
ally much longer than the upper of 2 subtending bracts,
soon elongating after pollination. Flowers coriaceous,
persistent in fruit, cartilaginous, not opening fully and
difficult to spread, brick red to yellowish green with red
stain on the inner surface, lip dark red. Sepals similar;
dorsal lanceolate, acute, concave, apically carinate, 2.5-
3.2 cm long, 5-6 mm wide; lateral sepals narrowly
ovate, falcate, 2.5-3 cm long, 5-7 mm wide. Petals nar-
rower than the sepals, falcate, 2.2-2.8 cm long. Lip
hinged to the column foot, elliptic-oblong to narrowly
rhombic, arcuate, thickened along the margin and at the
apex, 2-2.6 cm long and 5-6 mm wide; callus linear
from the base to just below the middle, the apical fifth
appearing oily internally. Column cylindric about 1.7-
2.2 cm long; a foot 3 mm long; pollinia 4, supported on
a stipe 3 mm long; anther bed ciliate at the margin.
Capsule ellipsoid, 3-4 cm long.
A locally common epiphyte of wet forests, 50
to perhaps 1000 m. Uncommon from Mexico
(Campeche? and Chiapas) to Chiriqui, Panama;
possibly in Colombia and Peru. Flowering mostly
October to February. In Central America, M.
aciantha is found almost exclusively in the Carib-
bean drainage.
This is closely related to sympatric M. fried-
richsthalii, and the two species sometimes occur
on the same host tree. The shoots and flowers of
M. aciantha are conspicuously larger, and the
flower color is reddish to yellow-green, never
white. It also differs from closely related M. scor-
pioidea Kraenzl, by the larger flowers with dark
red, unspotted lip. That species is almost exclu-
sively restricted to the Pacific drainage.
Maxillaria acostae Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:301. 1923. TYPE: Costa
Rica, near San Ramon, Acosta s.n. (neotype se-
lected here: AMES 40543). Figure 25C.
Plant a small, cespitose epiphyte rarely to 20 cm tall.
Roots to 0.5 mm in diameter. Pseudobulbs ovoid, 1-2
cm long, subtended by nonfoliaceous, triangular sheaths
transversely spotted with purple; apex 1 -foliate. Leaves
with petiole V4-Vi the blade length; blade somewhat co-
riaceous, narrowly to broadly elliptic, 22-100 mm long,
4-17 mm wide, acute to obtuse or acuminate. Inflores-
cences 1-2 per rhizome bract axil of the mature shoot;
scape 2-3 cm long, concealed by spotted, acuminate
sheaths; ovary with pedicel 7-1 1 mm long, somewhat
exceeded by the floral bract. Flowers white turning or-
ange-brown, the lip with crimson callus and midlobe.
Sepals dissimilar; dorsal lance-oblong, obtuse to acute,
1-1.2 cm long, 3-4 mm wide; lateral sepals oblique,
oblong, 1.2-1.3 cm long, 3.5-4.5 mm wide, acute to
obtuse. Petals lanceolate-elliptic to oblanceolate, 1 cm
long, 2.5-3 mm wide, acute to obtuse. Lip hinged to the
column foot, cuneate to obovate, 9-1 1 mm long, 3-5.5
mm wide when spread, the apical '/5 deeply 3-lobate;
lateral lobes embracing the column rounded; midlobe
suborbicular to ovate, somewhat recurved; callus an ob-
long, fleshy ligule ending about 3/5 from the base. Col-
umn arcuate, 5-6 mm long excluding the anther; foot
40
FIELDIANA: BOTANY
2.5-3.5 mm long; anther about 2 mm long. Capsule
ellipsoid, about 1.5 cm long.
Epiphytic at 650-1500 m. Flowering March to
December. Rarely collected, perhaps because of
its small size. Costa Rica and Panama.
There appear to be two kinds of M. acostae
with identical flowers but different leaves, those
with a maximum leaf width-to-length ratio of ^
0.14 and those with a ratio of < 0.11. Both veg-
etative types are known from Costa Rica and Pan-
ama, but the vertical range of the broad leaf form
is 650-1000 m, while that of the narrow leaf form
is 1050-1500. There may be two distinct species
represented here, but the apparently identical
flowers do not support this. Similar leaf width-to-
length ratios taken from the type description are
0.09 and 0.10, corresponding with higher eleva-
tion forms. None of this form has been recently
collected. The designated neotype is all that has
been found of the original materials.
Maxillaria acutifolia Lindl., Edwards's Bot. Reg.
25: misc. 92. 1839. TYPE: Demerara, Loddiges
s.n. (holotype: K). Not Maxillaria acutifolia
Schltr. Figure 21B.
Plant a cespitose epiphyte rarely forming large
clumps; shoots to about 1 cm between pseudobulbs.
Roots fibrous, to about 1 mm in diameter. Pseudobulbs
1 -foliate, cylindric, to about 6 cm long and 1.5 cm in
diameter. Leaf shortly petiolate, narrowly elliptic, to 21
cm long and 1.5 cm wide, acute. Inflorescence usually
shorter than the pseudobulbs; ovary with pedicel 1.3-
1.8 cm long. Flower dull yellow, the lip yellow with red
spots and a red central line. Sepals similar, 1.2-1.5 cm
long, about 4 mm wide; dorsal oblong-lanceolate, lateral
sepals somewhat oblique, falcate, acute to obtuse. Petals
elliptic, 1-1.4 cm long, 2-3 mm wide, acute. Lip artic-
ulated with the short column foot, 3-lobate below the
middle, 1-1.2 cm long; lateral lobes acute; midlobe
more or less oblong, truncate, and retuse. Column ar-
cuate, clavate, 7 mm long, with a short foot about 2 mm
long.
Tropical forests at 0-1350 m. Flowering De-
cember to February. Local from Honduras to Ec-
uador and at least northern South America.
This is one of the two smaller flowered species
of the M. rufescens complex in Costa Rica, the other
being M. dressleri, with wine red to maroon flowers.
Maxillaria acutifolia differs most strikingly in the
more or less cylindric, not ovoid pseudobulb and
relatively narrow leaf. In live material the red line
up the center of the lip is the most distinctive floral
feature of the species. Maxillaria hedwigiae is a
larger plant with proportionately broader pseudo-
bulbs. Plants accepted here as M. acutifolia have a
somewhat more cuneate midlobe than the type and
may prove to be the same as M. moralesii, in which
case the latter name should be used.
Maxillaria adendrobium (Rchb. f.) Dressier,
Taxon 13:248. 1964. Ponera adendrobium
Rchb. f., Flora 48:278. 1865. TYPE: Cuba,
Wright 1697 (isotypes: AMES 6081, 6082) Pleu-
ranthium adendrobium (Rchb. f.) Benth. &
Hook, f., J. Linn. Soc. 18:312. 1881. Neour-
bania adendrobium (Rchb. f.) Fawc. & Rendle,
J. Bot. 47:125. 1909. Figure 4A.
Plant sometimes more than 1 m tall lacking pseudo-
bulbs; stems somewhat fractiflex, pendent, apparently
monopodial, mostly concealed by leaf sheaths. Roots ap-
pearing along the entire stem to 2.5 mm in diameter.
Leaves coriaceous, narrowly elliptic, articulate to the
sheaths, usually deciduous below, 5.5-11 cm long, 7-
15 mm wide, acute. Inflorescences 1-few per leaf axil,
not much exerted beyond the subtending sheath; ovary
with pedicel about 10 mm long, subtended by a shorter,
triangular bract. Flowers pale green to yellow, purplish
toward the base; column dark. Sepals subequal, narrow-
ly ovate, 5.5-7 mm long, about 2 mm wide, acute. Pet-
als narrowly ovate, 5 mm long, 2 mm wide, acute. Lip
attached to the column foot, 3-lobate, about 5 mm long;
lateral lobes short, ovate; midlobe large, ovate-elliptic,
retuse, conspicuously mucronate abaxially near the
apex; margins somewhat crenate; callus globose at the
base of the midlobe. Column stout, about 2 mm long;
foot at approximately a right angle with the column,
about 2 mm long. Capsule ellipsoid, about 1.3 cm long.
Epiphytic mostly in rain forests at 350-850 m.
Flowering September to November. Costa Rica,
Panama, Jamaica, and Cuba; doubtfully reported
from Ecuador on the basis of a similar plant with
a different lip.
Maxillaria adolphii Ames & Correll, Bot. Mus.
Leafl. 11:18. 1943. Camaridium adolphii Schltr.
(invalidly published), Repert. Spec. Nov. Regni
Veg. Beih. 19:58. 1923. Ornithidium tonduzii
Schltr., Repert. Spec. Nov. Regni Veg. 3:250-
251. 1907. TYPE: Costa Rica, on the banks of
the Rio Angel (Poas), Tonduz 10770 (lectotype
selected here: us 938677; isolectotype: us
577310; drawing of type: AMES). Not Maxillaria
tonduzii (Schltr.) Ames & Correll. 1943. Not
Camaridium tonduzii Schltr. 1910. Figure 5C.
Plant a large, cane-forming, apparently monopodial
epiphyte or terrestrial to 2 m tall, often becoming pen-
dent with size; juveniles cespitose, sympodial with each
shoot terminating in a pseudobulb. Mature stems
sheathed with closely spaced, conduplicate leaves, most-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
41
ly deciduous near the base with sheaths fragmenting into
fibers. Roots slender, to about 1 mm in diameter. Pseu-
dobulbs of juveniles to about 10 cm long, subtended by
1 or more foliaceous sheaths, the apex l(-2?)-foliate.
Leaves articulate, coriaceous, keeled beneath, shortly
petiolate above the articulation; blades 5-20 cm long,
1.2-2.8 cm wide; apex unequally 2-lobate. Inflores-
cences several to many per leaf axil; scapes to nearly
10 cm long but not much exerted beyond the subtending
leaf blade, clothed in acute sheaths; ovary with pedicel
8.5-1 1 mm long, usually exceeded by the subtending
floral bract. Flowers fragrant, with somewhat recurved
segments, pinkish lavender to white, the lip purple. Se-
pals similar, oblong-oblanceolate, 1.4-1.9 cm long, 4-5
mm wide, the apex acute and externally keeled; petals
elliptic-oblanceolate, 1.2-1.4 cm long, 4-5 mm wide,
the apex acute. Lip rigid with the column foot, subsac-
cate on the lower half, 3-lobate above the middle, 4-7
mm long, 2.5-4 mm wide in natural position; lateral
lobes little extended beyond the sinuses, rounded; mid-
lobe ovate, rounded to emarginate, recurved; callus a
truncate ligule ending just below the lobe sinuses with
an additional 3 or more papillae at the base. Column
stout, 3-4 mm long excluding the anther, the anther
about 1.5 mm in diameter. Capsules ellipsoid, about 1
cm long.
Epiphytic or terrestrial fully exposed in windy,
cold cloud forests at 2200-3300 m. Flowering
November to August. Endemic to Costa Rica and
western Panama.
Maxillaria adolphii is distinguished from close-
ly related M. biolleyi (Schltr.) L. O. Williams by
the more saccate lip with relatively complex cal-
lus that is papillose at the base and by a pair of
lateral lobes distally. Maxillaria biolleyi has or-
ange in the lip, a much more simple callus lacking
papillae, and a pair of lateral lobes at the base.
Available herbarium specimens with flowers in-
termediate with M. biolleyi suggest that natural
hybrids may occur. Similar Maxillaria quadrata
with nonsaccate lip may prove to be a variant of
M. adolphii.
The nomenclature of M. adolphii was confused
when Schlechter indicated that Ornithidium ton-
duzii Schltr. "might" (moge) be called C. adolphii
Schltr. in the future but did not accept the new
name at the time of publication, thus rendering it
invalidly published. Although Ames and Correll
published the combination as M. adolphii, it must
be accepted as a new name. The lectotype is the
more complete of two isotypes that surfaced at us
and is provided with floral analysis.
Maxillaria alba (Hook, f.) Lindl., Gen. & Sp.
Orch. PI. 143. 1832. Dendrobium album Hook,
f., Exot. Fl. t. 142. 1825. TYPE: Jamaica (ho-
lotype: K). Broughtonia alba (Hook, f.) Spreng.,
Syst. Veg. 3:735. 1826. Camaridium album
(Hook, f.) Hoehne, Arq. Bot. Est. Sao Paulo
2(4):72. 1947. ^Maxillaria hedyosma Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 42:131.
1925. Figure 15D.
Plant epiphytic with ascending rhizomes, each ter-
minated in a pseudobulb and rooted only at the base;
successive shoots 1.5-6 cm apart. Roots white, to 1.5
mm in diameter. Pseudobulbs ancipitous, ovate, smooth,
2.5-6 cm long, 1-2.5 cm broad, 1 -foliate; articulation
usually oblique, subtended by sheaths often with soon
deciduous blades. Leaves shortly petiolate, subcoria-
ceous, linear to narrowly elliptic, 20-40 cm long, 1.4-
2.5 cm wide, tapering to an obtuse apex. Inflorescences
1 in each of several rhizome bract axils of the devel-
oping shoot; scapes 2-3 cm long, hidden by the sub-
tending rhizome bract; ovary with pedicel 2.5-3.5 long,
much exceeding the attenuate floral bract. Flowers fra-
grant, white, the lip creamy white with yellow apex.
Sepals lanceolate, acuminate, 2.2-2.5 cm long, 5-7 mm
wide. Petals obliquely lanceolate, 1.8-2.2 cm long, 5-6
mm wide, acute. Lip articulated to the column foot,
ovate, puberulent, lightly 3-lobate on the apical l/3, 1.3-
1.6 cm long, 4-6 mm wide; callus extending to just
beyond the middle, parabolic in front. Column arcuate,
about 1 cm long; foot 3 mm long. Capsule ellipsoid.
Large crown epiphyte of wet forests at 250-
1200 m. Flowering throughout the year. Guate-
mala to Panama, Jamaica, Cuba, and much of
tropical South America.
This is most similar to M. anceps Ames & C.
Schweinf. and M. nagelii L. O. Williams (not in
Costa Rica), both of higher elevations, with de-
cidedly greenish flowers and more conspicuous
callus. It also resembles M. oreocharis Schltr., a
species with smaller, reddish-stained flowers that
form at the base of mature shoots.
Maxillaria alfaroi Ames & C. Schweinf., Sched.
Orch. 10:83-84. 1930. TYPE: Costa Rica, Car-
retera Fuentes, Alfaro 284 (holotype: AMES).
Figure 2B.
Plant stout, with erect stems perhaps 50 or more cm
tall, apparently lacking pseudobulbs; stems erect, con-
cealed by the distichous, imbricate, conduplicate leaf
sheaths. Roots large, 2.5 mm in diameter. Leaves linear-
oblong to narrowly elliptic, to 23 cm long, 2.5 cm wide,
becoming smaller toward the stem apex, the midvein
sulcate above and carinate below; blade articulate to the
sheath base; apex rather abruptly narrowed, obtuse or
rounded. Inflorescence single per leaf axil, the scape to
about 8 cm long and concealed by several acute sheaths.
Flowers orange-yellow. Sepals dissimilar; dorsal con-
cave, oblong-ovate, about 1.9 cm long, 8-9 mm wide,
acute and mucronate; lateral sepals somewhat oblique,
broadly lanceolate, 1 .9 cm long, about 7 mm wide, acute
and mucronate. Petals ovate-oblong, 1.6 cm long, 6.5-
42
FIELDIANA: BOTANY
7 mm wide, acute, somewhat oblique. Lip simple, sub-
quadrate-oblong, slightly constricted in the middle,
about 1.4 cm long, 6.5 mm wide; apex broadly rounded
and somewhat retuse. Column stout, strongly arcuate,
7-8.5 mm long. Capsule apparently narrowly ellipsoid.
Maxillaria alfaroi is represented here only by
the type collection and by a photograph of a sin-
gle plant in a live collection in British Columbia.
The above description is condensed from the orig-
inal and augmented from a photograph of the sin-
gle live plant. Little is known of its ecology ex-
cept that the type was collected at 1400 m and in
flower in August. It appears to be related to, if
not identical with, M. densifolia (Poeppig &
Endl.) Rchb. f. from South America. It should be
easily recognized by the relatively large canes
lacking pseudobulbs and the rather simple flower
with simple lip.
Maxillaria amabilis J. T. Atwood, Lindleyana 9:
239-241. 1994. TYPE: Costa Rica: Province of
Alajuela: Zona Protectora, Chayote, Llano Bo-
nito, Naranjo, 1200 m, L. D. Gomez, G. Her-
rera, & W. Murillo 23979 (holotype: MO; iso-
types: F, SEL). Figure 7B.
Plant a large, erect, cane-forming epiphyte to at least
90 cm tall; canes of mature shoots lacking pseudobulbs,
concealed by rugose leaf sheaths, mostly leafless on the
lower canes. Roots mostly restricted to the base, to about
1.5 mm in diameter. Pseudobulbs present only at base
(as cespitose juveniles), about 5 cm tall and 2.5 cm wide,
compressed, sessile, subtended by 1-2 foliaceous
sheaths, apically 1 -foliate with shortly petiolate leaf.
Leaves thin, articulate; blade keeled below, 4-7 cm
long, 1.3-2.8 cm wide, the apex obtuse to rounded and
unequally 2-lobate. Inflorescences several per leaf axil,
long for the genus and sometimes exceeding the sub-
tending leaf; scapes to about 9 cm long; ovary with ped-
icel 9-11 mm long, subtended by a subequal to longer
subulate, acute floral bract. Flowers large for the genus,
campanulate, to about 2.5 cm long, white to lavender,
or the outer surfaces of sepals and petals with pink
stripe, lip yellow or orange. Sepals similar, oblong-ellip-
tic, 2.2-2.5 cm long, 4.5-6 mm wide, the apex acute
and carinate externally. Petals oblanceolate, about 2.2
cm long, 5.5-6.5 mm wide, acute to acuminate. Lip
loosely rigid with the column foot, canaliculate on the
lower half, 3-lobate on the upper half, about 9 mm long
and 3 mm wide in natural position, 1.2 cm long when
spread; lateral lobes embracing the column; midlobe
ovate, obtuse; callus a truncate lamina terminating at
about the lobe sinuses with a keel on the lower Vi ele-
vated and acute in front. Column nearly straight, cylin-
dric, 9 mm long excluding foot and anther; foot short,
about 2 mm long; anther about 1.5 mm in diameter.
Maxillaria amabilis is a poorly known epiphyte
apparently of premontane and montane rain for-
ests at 1200-1900 m. Flowering at least in Oc-
tober and January. Endemic to Costa Rica.
Maxillaria ampliflora C. Schweinf., Bot. Mus.
Leafl. 8:188. 1940. Camaridium grandiflorum
Ames, Proc. Biol. Soc. Wash. 34:149-150.
1921. TYPE: Panama, Chiriqui, east of the Rio
Caldera, Killip 3565 (holotype: AMES; isotype:
us). Not Camaridium grandiflorum Schltr. Not
Maxillaria grandiflora Lindl. Figure 14E.
Plant a straggly terrestrial or epiphyte forming erect
to decumbent canes to about 2 m long; stems comprised
of successively borne shoots, each terminated by a pseu-
dobulb. Roots forming only on the lowest shoots, to 2.5
mm in diameter. Pseudobulbs ovoid, compressed, to
about 5 cm long; base subtended by several leaves; apex
1 -foliate. Leaves distinctly and shortly petiolate, con-
duplicate; blade narrowly elliptic, increasing in size to-
ward apex within each shoot, to about 30 cm long, 4 cm
wide, acute. Inflorescence 1(2) per bract axil, subtended
by a foliaceous bract with distinct articulation; scapes
sheathed in acute bracts, to about 7 cm long; ovary with
pedicel 1.3-1.6 cm long, exceeded by the subtending
floral bract. Flowers somewhat campanulate, white or
pinkish to light yellow, the recurved segments lightly
barred with lavender; lip white and yellow with reddish
transverse bars. Sepals oblong to lanceolate, the apical
margins often involute; dorsal 3.5-5 cm long, about 1
cm wide; lateral sepals somewhat oblique 3.5-5 cm
long, about 1 cm wide, acute. Petals elliptic-lanceolate,
3.2-4.5 cm long, about 1 cm wide; apical margins some-
what involute; apex acute to attenuate. Lip hinged to the
column foot, 3-lobate, about 1 cm long and wide when
spread; lateral lobes somewhat clasping the column,
rounded to bluntly acute, the midlobe rounded to acu-
minate, often with involute margin; callus a raised ligule
ending at about the sinuses of the lobes, 3-dentate in
front, papillose on the lower %. Column arcuate, 8-10
mm long excluding the anther; anther about 2.5 mm in
diameter.
Epiphytic and terrestrial in cloud forests at
1000-2100 m. Flowering January to March and
August to October. Apparently more common in
Panama than in Costa Rica, if available herbarium
specimens are an indication. Also reported from
Colombia and Ecuador as M. bradeorum (Schltr.)
L. O. Williams (sensu laid).
Maxillaria ampliflora is most similar to M. bra-
deorum (Schltr.) L. O. Williams, a species with
smaller flowers, more purple in the floral seg-
ments, and a lip with a proportionately smaller
midlobe. These species are sympatric.
Maxillaria anceps Ames & C. Schweinf., Sched.
Orch. 10:84. 1930. TYPE: Costa Rica, La Pal-
ma A. M. Brenes (133) 463 (holotype: AMES,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
43
photo; isotype: CR). Not Ornithidium anceps
Rchb. f. Figure 16A.
Plant large, epiphytic, with ascending shoots each ter-
minated by a pseudobulb and borne 3.5-7 cm apart;
rooted to the substrate only on lower shoots. Roots
white, to 2 mm in diameter. Pseudobulbs ovoid, 1 -fo-
liate, strongly compressed, shiny, 2-5.5 cm long, 1-2.5
cm wide, subtended by rhizome bracts, which are soon
deciduous along the oblique articulation, the apex 1 -fo-
liate. Leaves subcoriaceous, narrowly elliptic to oblong,
shortly petiolate, the blade 6-30 cm long and 1-3.5 cm
wide, apically obtuse to unequally bilobed. Inflores-
cences 1 per each of several rhizome bracts of the newly
forming shoot; scapes 1-5 cm long, covered by 3 or
more acute sheaths and usually concealed by the rhi-
zome bract; ovary with pedicel 2.5-4 cm long, much
longer than the subtending bract. Flowers fragrant,
greenish white to yellow. Sepals lanceolate-ovate, 1.8-
2.1 cm long, 6-7 mm wide. Petals elliptic-oblong, 1.6-
1.8 cm long, 4-5 mm wide. Lip hinged to the column
foot, ovate, strongly 3-lobate on the apical third, 1 .5-1 .7
cm long, 7-9 mm wide, thickened at the base, glabrous
inside; callus orbicular just below the middle (difficult
to rehydrate in pressed specimens). Column arcuate,
about 1 cm long; foot about 2 mm long. Capsule ellip-
soid, 3 cm long.
Large clump-forming epiphyte of tree trunks
and larger branches in wet forests at 1000-1600
m. Flowering November to February. Guatemala
to Costa Rica.
Maxillaria nagelii L. O. Williams, ranging
from Mexico to Guatemala and El Salvador, is a
similar middle elevation species with similar or-
bicular callus but with distinctly acute perianth
segments and less distinctly 3-lobate lip. Maxil-
laria anceps also resembles M. alba (Hook, f.)
Lindl., a species with white flowers and longer,
more acute floral segments, a more pronounced 3-
lobate lip, and ligulate callus extending beyond
the middle. Maxillaria anceps also is more re-
stricted in flowering season, geography, and ele-
vation than the widespread M. alba.
Maxillaria angustisegmenta Ames, Hubbard, &
C. Schweinf., Sched. Orch. 10:86-87. 1930.
TYPE: Costa Rica, La Palma, A. M. Brenes
(132) 462 (holotype: AMES). Maxillaria endresii
Rchb. f. var. angustisegmenta (Ames, Hubbard
& C. Schweinf.) C. Schweinf., nom. Maxillaria
luteoalba of authors, not Lindl. Figure 20C.
Plant a densely cespitose epiphyte with approximate
shoots. Roots white, to 1.5 mm in diameter. Pseudo-
bulbs 1 -foliate, conic to ovoid, rarely approaching or-
bicular, somewhat compressed, 2-7 cm long and 1 .5-2.5
cm wide, partially concealed at the base by several
smooth, chartaceous, minutely spotted, triangular, acute
bracts 3-10 cm long, shredding with age. Leaves long
(rarely short) petiolate; petiole sometimes more than 1A
the length of the blade; blade oblong-elliptic, 15-24 cm
long and 2.5-6 cm wide, acute. Inflorescence 1 -flow-
ered, the scapes 4-15 cm long, concealed by 4-6 in-
flated and minutely spotted bracts; ovary with pedicel
2-3 cm long, enclosed and concealed by the floral bract
before pollination. Flowers fragrant, mostly white, the
inner surface of the sepals yellow, the lip yellow and
centrally lined with purple on the side lobes, column
white to purple. Sepals similar; dorsal elliptic-oblong,
somewhat concave, 3-5 cm long, 7-10 mm wide, acute
to obtuse; lateral sepals oblong, tapering from about the
middle to a broadly acute apex, 3-5 cm long, 6-10 mm
wide at the base. Petals narrowly elliptic, falcate, strong-
ly incurved, 4 cm long and 4-7 mm wide, acute. Lip
obovate, sharply 3-lobate on the apical quarter, 2.1-2.6
cm long, 12-18 mm wide; midlobe about 6 mm long,
6.5 mm wide; callus on the lower l/i puberulent, raised,
thickened, and V-shaped in front. Column 1.8-2 cm
long, including the 5-8-mm foot. Capsule 4 cm long.
Epiphyte of larger branches or less often ter-
restrial, often forming large clumps; mostly in
cloud forests from (450)700-1800(2000) m.
Flowering from October to March. Costa Rica and
Panama.
This species is most similar to the earlier de-
scribed and probably sister species, M. luteoalba
Lindl. (TYPE: Venezuela, Linden 678, photo
seen), which doubtfully occurs in Panama as re-
ported. Maxillaria luteoalba has larger flowers
and a proportionately larger midlobe and shorter
callus in the lip. In Costa Rica M. angustisegmen-
ta is easily distinguished from the lower elevation
M. endresii Rchb. f. by the shorter and usually
proportionately broader perianth parts (despite the
name) and the proportionately smaller midlobe.
The anther bed of M. endresii is coarsely and ir-
regularly lacerate but low and minutely dentate in
M. angustisegmenta. Vegetatively, the pseudobulb
of M. endresii is nearly always suborbicular and
truncate or emarginate, while that of M. angusti-
segmenta is ovoid to conic.
Maxillaria angustissima Ames, Hubbard, & C.
Schweinf., Bot. Mus. Leafl. 3:41. 1934. Max-
illaria acutifolia Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:229-230. 1923 (not Lindl.,
1839). TYPE: Costa Rica, San Ramon, A. M.
Brenes 235 (holotype: B, destroyed; drawing of
type: AMES), Brenes 173 (504) (neotype: AMES).
Figure 18B.
Plant a cespitose epiphyte; shoots approximate, often
pendent, to 30 cm tall. Roots white, to 0.5 mm in di-
ameter. Pseudobulbs hidden from view by 3-4 subtend-
ing, leaf-bearing rhizome bracts, apically 1 -foliate.
44
FIELDIANA: BOTANY
Leaves linear, attenuate both toward the base and the
apex, 5-25 cm long, 4-11 mm wide, the leaves sub-
tending the pseudobulb with broadened sheaths below
the articulation. Inflorescences lateral in the rhizome
bract axils, the single-flowered scapes 1-3 cm long; ova-
ry with pedicel about 1.5 cm long, subtended by a sub-
equal, acute, floral bract. Flowers white to pink, the side
lobes of the lip often reddish. Sepals dissimilar; dorsal
ovate, concave, 9 mm long, 3.5-4 mm wide, apex
strongly concave; lateral sepals triangular, adnate to the
long column foot, 1.6 cm long, apex acute and strongly
concave. Petals oblong, about 1 cm long, 3 mm wide,
constricted just below the erose apex. Lip hinged to the
column foot, cuneate, strongly 3-lobate on the apical
fifth, 13-14 mm long, 4 mm wide, midlobe broadly
acute; callus ligulate, pubescent, rather deep and para-
bolic in front, extending to just below the base of the
midlobe. Column 4 mm long; foot 10 mm long. Cap-
sule somewhat curved, narrowly ellipsoid, 3.5-4 cm
long.
A rare epiphyte of wet forests at 900-1500 m.
Flowering August and September. Costa Rica,
Panama, and southern Nicaragua (Volcan Mom-
bacho).
The very narrow leaves and small, hidden pseu-
dobulbs along with the small white flowers pro-
duced in August and September are good features
for field recognition.
Maxillaria appendiculoides C. Schweinf., Bot.
Mus. Leafl. 4:119-121. 1937. TYPE: hills of
San Pedro de San Ramon, Brenes (239) 1427
(holotype: AMES; isotype: NY). Figure 3B.
Plant a straggly, often pendent epiphyte; stems elon-
gate, lacking pseudobulbs, concealed by sheaths. Roots
white, to about 1 mm in diameter. Leaves distichous,
deciduous on the lower stems, thin; blade elliptic to
broadly lanceolate, 1-4 cm long, 5-13 mm wide, round-
ed and unequally 2-lobate apically. Inflorescence 1 per
leaf axil, shorter than the subtending leaf; peduncle 1-2
cm long; ovary with pedicel about 1 cm long, subtended
by a somewhat shorter, acute bract. Flowers yellow or
greenish; lip with purple. Sepals ovate-lanceolate, some-
what concave, ovate-lanceolate, 8-9 mm long, 3-3.5
mm wide, lateral sepals somewhat oblique, acute. Petals
narrowly elliptic-linear, somewhat falcate, acute, 6.5-7.5
mm long, 1.5-1.75 mm wide. Lip hinged to the foot,
rhombic-ovate in outline when spread, somewhat 3-lo-
bate above the middle, 6 mm long, 4.5-5 mm wide when
spread; lateral lobes rounded and somewhat clasping the
column; midlobe subquadrate, rounded to truncate; cal-
lus simple, from near the base to about the middle, lig-
ulate, rounded in front. Column about 4 mm long ex-
cluding the anther; foot 2-2.5 mm long. Capsule ellip-
soid about 1.5 cm long, separating into 6 valves, with
persistent column.
Epiphytic and apparently rare in premontane
rain forests around 800-1600 m. Flowering Au-
gust to November. Endemic to Costa Rica, al-
though it may not have caught the eye of collec-
tors because of its understated appearance.
Maxillaria appendiculoides is closely related to
M. luteorubra (Lindl.) Rchb. f. from South Amer-
ica, a species with somewhat larger flowers. It is
easily distinguished from sympatric M. linearifol-
ia Ames & C. Schweinf. by the shorter and more
elliptic leaves and lower elevation range.
Maxillaria arachnitiflora Ames & C. Schweinf.,
Sched. Orch. 10:87-89. 1930. TYPE: Costa
Rica, Navarro, C. H. Lankester 1212 (holotype:
AMES). Figure 24C.
Plant a cespitose epiphyte, not forming large clumps.
Roots white, to 1 mm in diameter. Pseudobulbs slender,
to 4.5 cm long, concealed and subtended by 1-2 leaf-
bearing rhizome bracts. Leaves petiolate, the petiole to
V2 the blade length; blade narrowly to broadly elliptic,
8-20 cm long and 2-3.5 cm long, acute. Inflorescence
usually not longer than the leaves, 5-8 cm long, nearly
concealed by 5-6 obtuse, apiculate bracts 1-3 cm long;
ovary with pedicel 2-3 cm long, surpassed by the sub-
tending bract. Flowers with sepals creamy yellow, the
petals white, lip yellow, column with maroon at base.
Sepals 5.5-7 cm long, acuminate, apically spiraled or
pendent; dorsal 8 mm wide, the lateral sepals 1.4 cm
along the column foot. Petals acuminate, abruptly nar-
rowed below the middle, about 4 cm long and 6 mm
wide. Lip obovate, 1.8 cm long and 1 cm wide, 3-lobate
on the apical quarter, the apex thickened; callus sparsely
pilose, ligulate, thickened and raised in front, to just
above the lower Vz of the lip. Column arcuate, 9 mm
long from the receptacle and passing into a laterally and
shallowly sigmoid foot 12-14 mm long; pollinia 4, sup-
ported on a saddle-shaped viscidium.
Very little known; cloud forests at 400-1400
m. Flowering August to October and March. Cos-
ta Rica and Panama.
This species is most similar to M. chionantha,
M. pachyacron, and M. reichenheimiana, but the
leaves are unspotted, the callus reaches to just
above middle of the lip, the sepals and petals are
more acuminate, the tips are often spiraled or pen-
dent, and the column foot is extremely pro-
nounced and somewhat sigmoid.
Maxillaria attenuata Ames & C. Schweinf.,
Sched. Orch. 10:89-90. 1930. TYPE: Costa
Rica, La Palma, Brenes 512 (holotype: AMES;
isotype: NY). Figure 23B.
Plant a small, cespitose epiphyte usually forming
small clumps. Roots white, to about 0.75 mm in diam-
eter. Pseudobulbs ovoid, to about 1 cm long, partially
concealed at the base by 1-2 foliate sheaths, 1 -foliate
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
45
apically. Leaves petiolate; petiole to about 1A the length
of the blade and narrowed above the articulation of fo-
liate sheaths; blade elliptic, about 2-7 cm long, 5-10
mm wide, acute. Inflorescence apparently single per leaf
axil; peduncle 4-7 cm long; somewhat scurfy ovary
with pedicel about 1.7-1.9 cm long at anthesis, subtend-
ed by a somewhat shorter, obtuse floral bract. Flowers
not opening widely, brick red or greenish-stained with
maroon. Sepals lanceolate-attenuate, acute; dorsal con-
cave at the base, 1.6-2 cm long, 3.5-4 mm wide; lateral
sepals somewhat falcate, 1.8-2.2 cm long, 5 mm wide.
Petals lanceolate, falcate, attenuate, 1.4-1.7 cm long,
2.5 mm wide, acute. Lip elliptic to rhombic, attenuate,
obscurely 3-lobate at about the middle, 1.4-1.5 cm long,
5-6 mm wide when spread, the apex acute and keeled;
callus simple, ligulate, rounded and thickened in front,
terminating at or below the middle. Column stout, about
4.5-5 mm long without the anther; foot about 5 mm
long. Capsule more than 3 cm long.
Epiphytic in wet, premontane rain forests at
1000-1300 m, often partially concealed by the
thick mats of epiphytic mosses in which it grows.
Flowering August to December. Costa Rica; also
reported from South America.
Maxillaria attenuata is recorded from Venezue-
la and Bolivia and may be conspecific with the
earlier name M. guadalupensis Cogn. as accepted
by Garay and Dunsterville (Venez. Orch. 111. 6:
250). A Bolivian plant illustrated as M. attenuata
(Icones Plantarum Tropicarum, Series II, Plate
251. 1989) is probably the same species, but the
lip has a proportionately longer midlobe. Plants
from Venezuela are recorded from 2285 m.
Maxillaria bicallosa (Rchb. f.) Garay, Caldasia
8:527. 1962. Zygopetalum bicallosum Rchb. f.,
Otia Bot. Hamb. 1:9. 1878. TYPE: Ecuador,
near Guayaquil, Lehmann (holotype: w). Kefer-
steinia bicallosa (Rchb. f.) Rchb. f. Otia Bot.
Hamb. 1:31. 1878. Figure 18D.
Plant epiphytic, cespitose, psygmoid, 20-40 cm tall.
Roots filiform, to 1.5 mm in diameter. Leaves equitant,
with unspotted, papery-margined sheaths 3-6 cm long
at the base; blade thin, spreading, linear-oblanceolate, to
45 cm long, 7-13 mm wide. Inflorescences 2 or more
within each leaf axil; scapes 2-4 cm long; pedicellate
ovaries (17)2.5-3 cm long, much exceeding the sub-
tending acute bract. Flowers wine red. Sepals similar,
elliptic lanceolate, 9-10 mm long, 2.5-3 mm wide,
acute. Petals elliptic, about 8 mm long, 2 mm wide,
acute. Lip narrowly ovate, slightly constricted above the
middle, 8-9 mm long, 3-4 mm wide, with rounded
apex; with ligulate callus from the base to about % the
lip, apex rounded to somewhat 2-lobate. Column cylin-
dric, about 5.5 mm long excluding the anther (1 mm
long); foot nearly obsolete. Capsule linear, about 2.5 cm
long.
Epiphytic in cloud forests at 1 100-1200 m. Flow-
ering September and October. Costa Rica (southern)
and Panama; also western Ecuador (600 m).
Maxillaria bicallosa is most closely related to
M. chartacifolia Ames & C. Schweinf. but differs
in its narrower leaves, which lack spots at the base
and have much shorter scapes and wine-red flow-
ers. The type sheet at Vienna shows broader
leaves than those from Costa Rica.
Maxillaria biolleyi (Schltr.) L. O. Williams., Ann.
Missouri Bot. Card. 28:425. 1941. Ornithidium
biolleyi Schltr., Repert. Spec. Nov. Regni Veg.
9:29-30. 1910. TYPE: Costa Rica, near San
Jose, Biolley 1052 (drawing of type: AMES). Ca-
maridium biolleyi (Schltr.) Schltr., Beih. Bot.
Centralbl. 36:498. 1918. Figure 5A.
Plant an erect to pendent epiphyte lacking pseudo-
bulbs (juveniles?); mature stems apparently monopodial,
rather thick and concealed by imbricate leaf sheaths.
Roots to about 1 mm in diameter. Leaves conduplicate,
lacking distinct petioles, deciduous, especially on the
lower stem; blades above the articulation 5-28 cm long,
1.5-3.2 cm wide, obtuse and unequally 2-lobate. Inflo-
rescences 2-many per leaf axil, shorter than the sub-
tending leaf; scape 5—7 cm long; ovary with pedicel 6-
7 mm long, subtended by a somewhat membranaceous
floral bract about 1 . 1 cm long. Flowers white or pinkish
with pink or salmon lip. Sepals similar, oblong-lanceo-
late, 1.3-1.5 cm long, 3-3.5 mm wide, acute. Petals
elliptic-lanceolate, 1.1-1.2 cm long, 2.5-3 mm wide,
acute to attenuate. Lip somewhat flexible with the col-
umn foot, 3-lobate below the middle, about 5 mm long
and 5 mm across when spread; lateral lobes basal and
erect around the column, rounded; midlobe biblike,
rounded to emarginate; callus between the lateral lobes,
simple, raised, heart-shaped. Column stout, 1.5-2 mm
long without the anther; foot less than 1 mm long.
Epiphytic and terrestrial at 1600-2200 m.
Flowering October to February. Costa Rica and
Panama.
Maxillaria biolleyi has long been confused with
M. adolphii (Schltr.) Ames & Correll, M. falcata
Ames & Correll, and M. quadrata Ames & Cor-
rell, but the smaller flowers on long scapes and
the relatively simple lip with heart-shaped callus
distinguish it from these.
Maxillaria brachybulbon Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:55. 1923. TYPE:
Costa Rica, San Jeronimo, Werckle 23 (drawing
of type: AMES). Figure 25A.
Plant a small cespitose epiphyte 4-8 cm tall. Roots
white, to about 0.75 mm in diameter. Pseudobulbs min-
ute, compressed, about 0.5 cm tall, apically 1 -foliate,
46
FIELDIANA: BOTANY
and subtended by usually 2 leaf-bearing sheaths. Leaves
oblong-oblanceolate, shortly petiolate, and obtuse, 2-8
cm long and 0.4-0.8 cm wide. Inflorescence apparently
arising on immature growth only, the scapes single per
rhizome bract, covered by up to 4 tubular, chartaceous
sheaths; ovary with pedicel shorter than the floral bract,
1.1-1.6 cm long. Flowers pale yellow or greenish,
sometimes striped with purple, the lip basally lined with
purple and the midlobe yellow. Sepals similar, acute;
dorsal elliptic, 1.4-1.6 cm long, 3-4 mm wide; lateral
sepals oblong-lanceolate, 1.6-2 cm long, 3-4 mm wide.
Petals narrowly elliptic, 1.5 cm long, 2.5 mm wide,
acute. Lip hinged to the column foot, about 1 cm long,
the midlobe fleshy and 3.3 mm long; callus ligulate on
lower %, parabolic and thickened apically. Column cy-
lindric, flattened beneath, 7 mm long including the foot
(2 mm long); anther cap 1.5 mm long; pollinia 4, lacking
a stipe but supported on a saddle-shaped viscidium.
Capsule ellipsoid, about 2 cm long.
Apparently rare epiphyte in rain forests at 50-
1000 m. Flowering throughout the year. Honduras
to Panama; also reported from Venezuela and Bra-
zil.
This species has been recognized by authors as
the South American M. xylobiiflora Schltr., a
somewhat larger species with linear leaves. Max-
illaria brachybulbon has a generally larger flower
and a proportionately longer midlobe than does
the closely related M. pterocarpa Barb. Rodr., of-
ten misidentified as M. brachybulbon.
Maxillaria bracteata (Schltr.) Ames & Correll,
Bot. Mus. Leafl. 11:14, 1943. Ornithidium
bracteatum Schltr., Repert. Spec. Nov. Regni
Veg. 9:217-218. 1911. TYPE: Costa Rica, La
Palma, Tonduz 12344 (lectotype: us). Camari-
dium bracteatum (Schltr.) Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:57. 1923. Maxillaria
vagans Ames & C. Schweinf., Sched. Orch. 8:
65-66. 1925. TYPE: Panama, Chiriqui, Powell
348 (holotype: AMES). Figure 16C.
Plant a somewhat ascending, cane-forming epiphyte
to about 50 cm tall comprised of shoots 3-15 cm long,
each terminated by a pseudobulb. Roots white, to about
1 mm in diameter. Pseudobulbs parallel with the con-
tinuing rhizome, fusiform to conic, compressed, 1 -foli-
ate, 2.5-5(7) cm long, 1-2(4) cm wide, subtended by 1-
4 strongly rugose sheaths with soon deciduous blades.
Leaf blades shortly petiolate, chartaceous, linear-oblong,
3-20 cm long, 1-3 cm wide, the apex unequally 2-lo-
bate. Inflorescences 1-3 per foliaceous sheath axil, 3-8
cm long; ovary with pedicel 8-12 mm long. Flowers
with somewhat spreading segments, white to pale yellow
with red or pink marks, especially the lip. Sepals similar,
oblong to oblanceolate, 1.7-2.2 cm long, 5-6 mm wide,
acute; dorsal somewhat keeled dorsally and apically.
Petals oblong-oblanceolate, 1.5-1.7 cm long, about 4
mm wide, acute. Lip ovate in general outline, 3-lobate
below the middle, 1.1-1.2 cm long, 6-7 mm wide when
spread; lateral lobes rounded, somewhat erect and em-
bracing the column; midlobe fleshy, rounded, abaxially
keeled; callus fleshy, truncate in front at about the same
level as the divergence of the lateral lobes, with a blunt
central keel. Column stout, arcuate, 4-5 mm long ex-
cluding the anther; foot about 2 mm long.
Common epiphyte of larger branches at 1 300-
2700 m. Flowering throughout the year. Costa
Rica and Panama.
Maxillaria bracteata is often seen on forest
remnants in pastures. In Costa Rica it is most sim-
ilar to M. gomeziana J. T. At wood, a species with
trigonous ovaries and statistically narrower
leaves. It strongly resembles M. imbricata Lindl.
from South America, but none of the available
illustrations of that species show the blunt keel of
the callus. For the moment these are best kept
separate.
This has long been known by the later syn-
onym, M. vagans Ames & C. Schweinf. The ap-
pearance of an isotype of M. bracteata (us) con-
sistent with Schlechter's original description
shows that this name has been badly misapplied
to other species (see Atwood, Lindleyana 9:239-
242, 1994).
Maxillaria bradeorum (Schltr.) L. O. Williams,
Ann. Missouri Bot. Card. 28:425. 1941. Ca-
maridium bradeorum Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:141-143. 1923.
TYPE: Costa Rica (photo of type: AMES). Max-
illaria semiorbicularis Ames & C. Schweinf.,
Sched. Orch. 8:64-65. 1925. TYPE: Costa
Rica, Province of San Jose, La Hondura, Stand-
ley 36247 (holotype: AMES). Figure 14D.
Plant a straggly terrestrial or epiphyte to at least 1.5
m long forming erect to decumbent canes comprised of
successively borne shoots, each shoot terminated by a
pseudobulb. Roots forming only on the lowest shoots,
to 2 mm in diameter. Pseudobulbs ovoid to suborbicu-
lar, compressed, to about 5 cm long; base subtended by
several leaves; apex l(-2?)-foliate. Leaves distinctly and
shortly petiolate, conduplicate; blade narrowly elliptic,
increasing in size toward apex within each shoot, to
about 20 cm long, 4.5 cm wide, acute. Inflorescence 1
per bract axil, subtended by a foliaceous bract with dis-
tinct articulation; scapes sheathed in acute bracts, to
about 7 cm long; ovary with pedicel (1)1.2-1.7(2.2) cm
long, exceeded by the subtending floral bract. Flowers
variable in color, white to light yellow, the tepals stained
with pink to lavender apically or spotted; lip yellow and
with reddish transverse bars. Sepals oblong to lanceo-
late; dorsal 2.2-3.1 cm long, 8-9 mm wide; lateral se-
pals somewhat oblique 2.1-3.2 cm long, 8-10 mm wide,
acute. Petals elliptic-lanceolate, 1.9-3 cm long, 7-10
mm wide; apical margins somewhat involute; apex acute
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
47
to attenuate. Lip hinged to the column foot, 3-lobate,
when spread 7-9 mm long, 1-1.3 cm wide; lateral lobes
clasping the column, the outer margin rounded, acute,
the midlobe subtriangular with acuminate apex; callus a
raised ligule ending below the sinuses of the lobes, 3-
dentate in front, papillose on the lower %. Column ar-
cuate, 8-10 mm long excluding the anther; foot (2.5)3-
4 mm long; anther about 2 mm in diameter.
Epiphytic and terrestrial in premontane and
lower montane rain forests at 700-1800 m. Flow-
ering at least July to November, January, and
March. Costa Rica and Panama.
Maxillaria bradeorum is closely related to sym-
patric M. ampliflora C. Schweinf. but has consis-
tently smaller flowers, usually with more pink or
lavender. The lateral lobes of the lip are propor-
tionately longer and more acute than in M. am-
pliflora.
Apparently Schlechter's type had a pair of
leaves at the pseudobulb apex, a feature not oth-
erwise seen among plants interpreted here as M.
bradeorum. The extant type drawings at AMES of
Camaridium bradeorum showing the lip with
smaller, more acuminate midlobe nicely distin-
guish this species from M. ampliflora C.
Schweinf. Ames and Schweinfurth argue in their
original description that their M. semiorbicularis
differs from C. bradeorum by the "smaller leaves,
monophyllous pseudobulbs, and different mid-
lobe." However, leaf size is generally extremely
variable in Maxillaria, and Schlechter appears to
have had an oddity in the 2-foliate pseudobulbs.
There is nothing in the original description of M.
semiorbicularis to support differences in the mid-
lobe of the lip with that of M. bradeorum.
Maxillaria brevilabia Ames & Correll, Bot. Mus.
Leafl. 11:15, fig. 1. 1943. Ornithidium alfaroi
Ames & C. Schweinf., Sched. Orch. 10:98.
1930. TYPE: Costa Rica, La Fuente, Alfaro 227
(holotype: AMES). Not Maxillaria alfaroi Ames
& C. Schweinf., Sched. Orch. 10:83-84. 1930.
Figure 9B.
Plant a somewhat creeping or lax cane-forming epi-
phyte to 50 cm tall; shoots forming an elongate scale-
covered rhizome to 7 mm in diameter. Roots to 1 mm
in diameter. Pseudobulbs ancipitous, elliptic-cylindric,
3-6 cm long, 1-3 cm broad, spaced 2-5 cm apart,
sheathed at the base when young by a pair of enlarged
nonfoliaceous bracts, apically 1 -foliate. Leaf shortly pet-
iolate; blade chartaceous in dried specimens, narrowly
elliptic, 4-17.5 cm long, 1.3-3.5 cm wide, obtuse or
rounded, rarely bluntly acute. Inflorescences developed
along the entire mature rhizome and usually densely so;
scapes less than 5 mm long, hidden by the rhizome
scales; ovary with pedicel 3-3.5 mm long concealed by
inflated scape bracts. Flower white sometimes with pink
or maroon flush at base of the segments, the lip orange-
yellow. Sepals ovate-lanceolate; dorsal 6.5-8 mm long,
about 2.5-3 mm wide; lateral sepals about Vi connate,
8-9.5 mm long. Petals oblong, 5.5-6.5 mm long, 1.5-
2 mm wide, subacute. Lip subovate when spread, 5-5.5
mm long, 3.5-4.5 mm wide, forming an acute angle
with the column foot, the apical third 3-lobate; lateral
lobes broadly rounded; midlobe triangular-ovate; callus
fleshy, transverse across the base of midlobe. Column
stout, about 2 mm long excluding the anther; foot about
2 mm long.
Epiphytic in montane rain forests, often in as-
sociation with oaks, Lauraceae and Chusquea, at
1000-2350 m. Flowering mostly August to Jan-
uary. Costa Rica (Volcan Poas) to Panama (Bo-
quete).
Maxillaria brevilabia is easily distinguished
from other members of the M. neglecta (Schltr.)
L. O. Williams complex by the relatively large,
cylindric pseudobulbs and broadly elliptic leaves.
Inflorescences develop along the entire length of
the most recently matured rhizome segment, and
the lip forms a sharp angle with the column foot,
unlike related species.
Maxillaria brunnea Linden & Rchb. f., Bonplan-
dia 2:281. 1854. TYPE: Colombia, Ocana,
Schlim 1161 (holotype: w). ? Maxillaria porrec-
ta Lindl., Bot. Reg. 24: misc. 92. 1838. TYPE:
Brazil(?), "Rio Janeiro," Loddiges s.n. (holo-
type: K). Maxillaria trinitatis Ames, Sched.
Orch. 2:34-35. 1923. TYPE: Trinidad, British
West Indies, heights of Aripo, W. E. Broadway
9890 (holotype: AMES). Maxillaria brenesii
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
56, 231-232. 1923. TYPE: Costa Rica, San Pe-
dro de San Ramon, Brenes 78 (lectotype: CR;
isolectotype: AMES). Figure 21 A.
Plant a densely cespitose epiphyte sometimes pro-
ducing large clumps. Roots white, to about 1 mm in
diameter. Pseudobulbs ovoid, little compressed lateral-
ly, 2.5-6 cm long, subtended by short, spotted, acute
sheaths; apex 1 -foliate. Leaves distinctly petiolate; pet-
iole 1-20 cm long; blade narrowly elliptic, usually glau-
cous beneath, 16-34 cm long, 2-5.5 cm wide, acute.
Inflorescences appearing in the bract axils of the re-
cently completed shoot; scapes 8-17 cm long, concealed
by 5-8 bracts of about the same length or increasing
gradually; ovary with pedicel 1.8-3.7 cm long. Flowers
creamy yellow flushed with rose, the lip apex purple
beneath. Sepals rather dissimilar, somewhat concave;
dorsal ligulate, 2.1-3 cm long, 5-6 mm wide, acute;
lateral sepals lanceolate-ligulate, oblique, 2.2-3.1 cm
long, 5-6 mm wide, acute to obtuse. Petals falcate-lig-
ulate, 2-2.5 cm long, 3-4 mm wide, acute. Lip generally
48
FIELDIANA: BOTANY
obovate when spread, 1.4-1.7 cm long, 7.5-10 mm
wide, 3-lobate on the apical %-V4; side lobes with mar-
gins embracing the column, rounded to truncate; mid-
lobe suborbicular, thickened centrally, the margins cren-
ulate; callus sparsely pubescent, ligulate, thickened and
somewhat V-shaped apically. Column stout, 7-9 mm
long excluding the anther; foot about 5 mm long; anther
about 3 mm in diameter. Capsule ellipsoid, 5-6 cm
long.
Locally common and widespread epiphyte on
large branches in rain or cloud forests at
(300)600-1800 m. Flowering June to March. Nic-
aragua (expected in Honduras) to Colombia and
Trinidad.
Maxillaria brunnea as accepted here may rep-
resent more than one species in Costa Rica, one
with yellowish flowers with larger bracts (perhaps
M. powellii Schltr., described from central Pana-
ma), the other with much more reddish flowers
with smaller bracts and shorter, more abrupt mid-
lobe (M. brenesii Schltr.). As interpreted for this
treatment, M. brunnea is characterized by the ces-
pitose habit, single leaf at the pseudobulb apex,
the creamy yellow to whitish flowers sometimes
tinged with red on the sepals, and dark red abaxial
side to the lip apex. Its close relative, M. longi-
loba, has a fleshier and proportionately longer
midlobe that is usually sharply reflexed. Speci-
mens from lower elevation are usually smaller,
with flowers less tinged with red. Flowers of re-
lated M. ringens Rchb. f. have longer and pro-
portionally narrower sepals colored white to yel-
low with the lip much shorter than Vz the lateral
sepals, while the lip of M. brunnea exceeds Vi.
The earliest name for members of the M. brun-
nea complex may be M. porrecta Lindl. (G. Car-
nevali, pers. comm.), an entity based presumably
on Brazilian material. Neither Ames nor Schlech-
ter compared their new species with M. brunnea
or M. porrecta.
Maxillaria burgeri J. T. Atwood, Lindleyana 9:
233-236. 1994. TYPE: Costa Rica, Province of
Heredia, southern slope of Volcan Barba at
2200 m, W. C. Burger & R. G. Stolze 6062
(holotype: F; isotype: MO). Figure 12C.
Plant an erect (when young), pendent, or decumbent,
much branched, cane-forming epiphyte to about 30 cm
tall, rooted only on the lower shoots; pseudobulbs to
about 5 cm apart on lower but closer in successive
shoots. Roots less than 1 mm in diameter. Pseudobulbs
suborbicular to obovoid, 7-12 mm long, 4-9 mm wide
often subtended when young by a foliaceous sheath, the
apex 1 -foliate. Leaves and foliaceous sheaths similar;
apical leaf with short petiole; blades thin, elliptic-ob-
long, rounded to emarginate, 1.5-9 cm long, 3-8 mm
wide, abaxially keeled. Inflorescence single per rhizome
bract of both mature and immature shoots, less than 1
cm long; ovary with pedicel 5-7 mm long, exceeded by
the subtending, acute, floral bract. Flowers not opening
widely, with lateral sepals and petals recurved, yellow;
lip yellow or with purple-red callus; column white. Se-
pals subequal, elliptic-lanceolate, 10-13 mm long, 2-
2.5 mm wide, acute; dorsal somewhat concave; lateral
sepals subfalcate. Petals elliptic-lanceolate, falcate,
acute, 9-12 mm long, 2.25-2.75 mm wide. Lip hinged
with the column foot, convex, recurved in natural po-
sition, ovate when spread, 4-5 mm long, about 3 mm
wide, 3-lobate on the lower V4; lateral lobes shorter than
the callus; midlobe ovate, reflexed at the base, obtuse or
rounded to emarginate; callus fleshy, rounded, and
somewhat elevated in front, terminating below the mid-
dle. Column stout, arcuate, somewhat tapering, 2-3 mm
long excluding the anther; foot about 1 mm long.
Poorly known epiphyte of montane rain forests
at 1 100-2500 m. Flowers May and June. Endemic
to Costa Rica.
Maxillaria burgeri is at once distinguished
from M. flava and M. microphyton by the subor-
bicular to ovoid pseudobulbs, which usually lack
a subtending foliaceous bract. Specimens have
been identified as M. microphyton, but the lip with
large midlobe is more similar to that of M. flava.
Maxillaria caespitifica Rchb. f., Linnaea 41:73.
1877. TYPE: Costa Rica, Endres (holotype: w).
Figure 25D.
Plant a cespitose to shortly rhizomatous epiphyte to
about 15 cm high. Roots to about 1 mm in diameter.
Pseudobulbs fusiform to cylindric, subtended by non-
foliaceous sheaths, apically 1 -foliate, 1.5-2.5 cm long.
Leaves thin, usually shortly petiolate, the blade lanceo-
late, to about 10 cm long, the apex unequally 2-lobate.
Inflorescences formed in the flush of new growth, to
about 2 cm long. Flowers yellow or greenish yellow,
the lip with red callus. Sepals similar, oblong, 5-7 mm
long, about 4 mm wide, acute to obtuse; dorsal concave;
lateral sepals slightly falcate. Petals oblong, 5-6 mm
long, about 3 mm wide; apex acute and recurved. Lip
simple to lightly 3-lobate, more or less straight with the
sides somewhat embracing the column, elliptic-ovate, 5-
6 mm long, about 3 mm wide, the apex truncate to
rounded or retuse; callus a fleshy and shiny ligule on
the lower Vi. Column arcuate, 5-6 mm long excluding
the anther; foot about 1 mm long.
Epiphytic but very poorly known; probably
from 100 to 1200 m as in Nicaragua. Costa Rica
and Nicaragua, possibly Honduras, and reported
from South America. Flowering in Costa Rica un-
known; January in Nicaragua. The above descrip-
tion is based in part on that by Hamer (Icon. PI.
Trop. 9: pi. 845).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
49
Some authors include this species as a synonym
of M. variabilis Bateman ex Lindl., but the more
cespitose plant and much smaller, usually greenish
flower of M. caespitifica are rather easy features
to recognize in live plants.
Maxillaria camaridii Rchb. f., Hamburger Gar-
ten-Blumenzeitung 19:547. 1863. Basionym:
Camaridium ochroleucum Lindl., Bot. Reg. 10:
t. 844. 1824. TYPE: Trinidad, Woodford. Cym-
bidium ochroleucum Lindl., Gen. Sp. Orch. PI.
168. 1833. Camaridium affine Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 17:72-73. 1922.
TYPE: Panama, on hills near Panama City,
Powell 7 (isotype: AMES). Not Maxillaria och-
roleucum Lodd. ex Lindl.
Plant a straggly, usually decumbent or pendent epi-
phyte. Stems elongate, the rhizome 2-15 cm between
pseudobulbs. Roots less than 1 mm in diameter. Pseu-
dobulbs elliptic-ovate, strongly compressed laterally,
2.5-5.5 cm long, 1.5-3 cm wide; base concealed by sev-
eral soon-deciduous, foliaceous sheaths; apex 1-2-foli-
ate. Leaves short-oblong to long-linear, short-petiolate
or not, the blades rarely to 33 cm long, 8-18 mm wide.
Inflorescence 1 (or more?) per foliaceous bract; scapes
to about 5 cm long, mostly included within the rhizome
bract; scape bracts acute; ovary with pedicel 1.6 cm
long. Flowers ephemeral, lasting about a day, white; lip
yellow with red or purple bars on lower half and at base
of column. Sepals dissimilar, elliptic, acute; dorsal
somewhat concave, 3.2 cm long, 1.2 cm wide; lateral
sepals 3.1 cm long, 1.2 cm wide. Petals elliptic-oblong,
somewhat falcate, 2.9 cm long, 8.5 mm wide, acute. Lip
strongly attached to but well distinguished from the col-
umn foot, 3-lobate on about the apical 3/5, 1.4 cm long,
1 .3 cm wide when flattened; lateral lobes rounded; mid-
lobe rounded in front; callus with rusty hairs on the low-
er half, the front part broken up into papillae. Column
cylindric, arcuate, about 1 cm long excluding anther;
foot about 2 mm long. Capsule ellipsoid, about 3 cm
long.
Epiphytic in lowland tropical forests at 0-250
m. Flowering August to November. Most of trop-
ical America north and west to at least Central
Panama; reported from Guatemala.
It is distinguished by the canes with pseudo-
bulbs and white, short-lived, fragrant flowers with
yellow lip marked with red. The rare M. suaveo-
lens Barringer is similar but is a much larger plant
with proportionately larger flowers. No specimens
of M. camaridii have been seen to confirm reports
from Costa Rica, but it may occur there.
Maxillaria campanulata C. Schweinf., Bot. Mus.
Leafl. 5:94-95. 1938. TYPE: Costa Rica, Ji-
menez 315 (holotype: AMES). Figure 15B.
Plant a large rhizomatous epiphyte with several
branching stems all rooted at the base, each to about 1.5
m long, comprised of successively borne shoots with each
terminated by a pseudobulb. Roots to about 1 mm in di-
ameter. Pseudobulbs ovoid, compressed, to about 40 cm
apart, to 10 cm long and 5 cm wide, mostly concealed
by several foliaceous bracts, the apex 1 -foliate. Leaves
coriaceous, keeled abaxially, articulate, petiolate; blades
oblong to linear or narrowly elliptic, to 25 cm long, 3.8
cm wide, with sheath blades relatively shortly petiolate,
the apical leaf with much more slender petiole, the apex
acute to obtuse and mucronate. Inflorescence 1 per foli-
aceous bract axil of the developing shoot; scape to about
10 cm long, longer than the subtending leaf, concealed
by acute bracts; ovary with pedicel 1.5-1.8 cm long ex-
ceeded by the subtending acute floral bract. Flowers cam-
panulate, somewhat nodding with recurved sepals and
petals, greenish yellow conspicuously marked transverse-
ly with brick red bars. Sepals oblong or oblanceolate;
dorsal about 2.2 cm long, acute; lateral sepals 2.3-2.5 cm
long, attenuate. Petals oblong, 1.9-2.1 cm long, 7-8 mm
wide, apex bluntly acute to obtuse. Lip hinged to the
column foot, obovate when spread, 9-10 mm long, 7-8
mm wide, 3-lobate on the upper V5, the sides embracing
the column; lateral lobes obtuse; midlobe fleshy, subor-
bicular, obtuse; callus a truncate or emarginate ligule end-
ing above the middle, extending toward the base where 3
or more low, warty keels are located. Column stout, near-
ly straight, 7-8 mm long excluding the anther; foot 1.5-
2 mm long. Capsule about 2.5 cm long.
Epiphytic in evergreen forests at 400-1500 m.
Conceivably the upper vertical distribution re-
flects locally warmer microclimates of pastures.
Flowering at least August to December. Endemic
to Costa Rica.
Maxillaria campanulata belongs to a group of
species characterized by the half-lyre-shaped fo-
liaceous bracts subtending the inflorescences, a
group that includes M. ampliflora, M. bradeorum,
and M. umbratilis. This species usually masquer-
ades as M. bradeorum, but the sepals and petals
are recurved, the lip is narrower than long when
spread, and the callus lacks the shaggy pubes-
cence at the base.
Maxillaria cedralensis J. T. Atwood & D. E.
Mora, Selbyana 18:31-32. 1997. TYPE: Costa
Rica, Province of San Jose, Cedral, 2000 m,
Apr. 1990, D. E. Mora B-35 (holotype: usj).
Figure 22D.
Plant cespitose forming large clumps, to 30 cm tall.
Stems approximate, roots to 1 mm in diameter. Pseu-
dobulbs ovate, compressed, 5-7 cm long, 2.5-4 cm
wide, subtended by inarticulate, triangular, nonfolia-
ceous bracts; apex 1-foliate. Leaf narrowly elliptic, to
45 cm long, 2.5-4 cm wide, apex bluntly acute. Inflo-
rescence 8-10 cm, slightly fractiflex, concealed by
bracts; ovary with pedicel 1.5-2 cm long, exceeded by
the acute, cucullate, floral bract. Flowers creamy white,
50
FIELDIANA: BOTANY
the petals spotted with maroon near the base of the te-
pals; lip cream, spotted with maroon; column spotted
with maroon on ventral side. Sepals elliptic, acute, with
revolute margins, 3.5 cm long, 1.3 cm wide. Petals el-
liptic-lanceolate, acute, about 3 cm long, 0.9 cm wide,
the margins revolute. Lip 3-lobate, 1.5 cm long, 0.8 cm
wide when spread; lateral lobes embracing the column,
truncate to somewhat emarginate apically; midlobe
about % lip length, recurved; callus 2-keeled, canalicu-
late, exceeding the lateral lobes in front. Column arcu-
ate, 1.25 cm long including the anther; foot very short;
pollinia 4, supported on a short stipe and viscidium.
Ecology poorly known, apparently epiphytic at
about 2000 m. Flowering at least March. Endemic
to Costa Rica, where it is known only from Cedral
de Aserri.
Maxillaria cedralensis is unique in the M. cu-
cullata complex with its large leaves, cream-col-
ored flowers, and spotted lip with rather narrow
midlobe. Maxillaria cedralensis differs from re-
lated M. punctostriata Rchb. f. and M. obscura
Lind. & Rchb. f. in its sepals with revolute mar-
gins.
Maxillaria chartacifolia Ames & C. Schweinf.,
Sched. Orch. 10:92. 1930. TYPE: Costa Rica,
Guanacaste, La Tejona, north of Tilaran, Stand-
ley & Valeria 45989 (holotype: AMES). Figure
18C.
Plant a psygmoid, large clumping, cespitose epiphyte
to about 40 cm tall, the shoots approximate. Roots to
1.5 mm in diameter. Pseudobulbs absent, obsolete, or
at least concealed by leafy rhizome bracts. Leaves 5 or
more per shoot at anthesis, distichous, with long sheaths
and petioles, the sheaths strongly conduplicate, spotted,
5-12 cm long and 1.2-2.5 cm from margin to midvein,
petiole narrowed just above the articulation, strongly
conduplicate, the blade 12-55 cm long and 1.7-4 cm
wide, the midvein sulcate above and keeled beneath. In-
florescence a fascicle of several usually successively
borne single-flowered scapes per rhizome bract axil, 7-
13 cm long, concealed by 4-6 acute, scape bracts 1.5-
3 cm long; ovary with pedicel 3-4 cm long, much ex-
ceeding the subtending bract. Flowers short lived, with
putrid odor similar to that of strong cheese, yellow at
anthesis, aging brownish red. Sepals lance-oblong, 1.8-
2.2 cm long, 6-8 mm wide, acute. Petals elliptic-oblan-
ceolate, 1.3-1.4 cm long, 4.5-5.5 mm wide, acute. Lip
hinged to the column foot, rhombic-ovate, obscurely 3-
lobulate, 1.3-1.4 cm long, 5.5-6.5 mm wide; callus
fleshy, extending from the base to apex, with an addi-
tional ovate, hard callus superposed at the middle. Col-
umn arcuate, broadened above, 8 mm long; foot 3-4
mm long, the anther cap 2.5 mm long. Capsule linear,
5-6 cm long, expanding into 6 valves and remaining
united at the apex.
A large clumping epiphyte of wet forests at
500-1700 m. Flowering September to November,
May, and to be expected throughout the year. Cos-
ta Rica to Ecuador.
This species and the closely related M. bical-
losa (Rchb. f.) Garay have a psygmoid (fanlike)
habit with at least 5 thin chartaceous leaves. Max-
illaria bicallosa has much narrower leaves (rarely
exceeding 1.2 cm) than M. chartacifolia, and its
sheaths are unspotted.
Maxillaria chionantha J. T. Atwood, Selbyana
16:242. 1995. TYPE: Panama, Province of Ver-
aguas, vicinity of Continental Divide, third
branch of Rio Santa Maria to drop-off to low-
lands, R. L. Dressier 4839 (holotype: SEL). Fig-
ure 24D.
Plant epiphytic, cespitose, sometimes densely so, 10-
30 cm high. Roots slender, to 1 mm in diameter. Pseu-
dobulbs ovoid, compressed, 2-3 cm long, usually con-
cealed by 1-3 subtending, foliaceous, and long petiolate
sheaths. Leaves long petiolate; petiole l/*-Vi the blade,
abruptly narrowed above the articulation; blade elliptic,
usually broadly so, 5-13 cm long, 2-5 cm wide, acute
and apiculate. Inflorescence 1 per rhizome bract; scape
3.5-8 cm long, concealed by acute scape bracts; ovary
1-2.2 cm long, somewhat warty, subtended, and much
exceeded by a cucullate, acute, floral bract. Flowers
campanulate, white aging tan with ivory to light yellow
lip. Sepals lanceolate, attenuate, 30-45 mm long, 6-8
mm wide, acute; laterals forming a chin 5-7 mm long.
Petals lanceolate, attenuate, 25-40 mm long, 4-6 mm
wide, acute. Lip oblong, simple to lightly 3-lobate on
the upper V6, about 12 mm long, 5 mm wide when
spread; lateral lobes slightly embracing the column; mid-
lobe deltoid, verrucose, with recurved apex; callus a lig-
ule from the base to above the middle, swollen in front.
Column 7 mm long, 1 1 mm including foot; foot 4-5
mm long; anther 2 mm long, somewhat hidden from the
broadened, dentate anther bed; pollinia 4, supported on
a stipe and saddle-shaped viscidium. Capsule 4.5-6 cm
long.
Epiphytic in premontane evergreen forests at
600-1000 m. Flowering August to February. Cos-
ta Rica and Panama.
Maxillaria chionantha closely resembles M.
pachyacron Schltr., which has usually longer in-
florescences, larger flowers with more attenuate
segments, and a much more distinctly 3-lobate lip
on the upper !/6. Maxillaria pachyacron also has
shorter pseudobulbs and grows at higher eleva-
tions. Maxillaria chionantha is also closely relat-
ed to M. attenuata Ames & C. Schweinf., a spe-
cies that also has distinctly petiolate leaves, atten-
uate floral segments, and a nearly simple lip, but
is easily distinguished by the larger plant and larg-
er and white flowers, rather than the red to ma-
roon flowers of the latter species.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
51
Maxillaria concavilabia Ames & Correll, Bot.
Mus. Leafl. 11:15, fig. 2. 1943. Ornithidium
stenophyllum Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19:59. 1923. TYPE: Costa Rica, San
Jeronimo, Werckle 109 (drawing of type: AMES).
Not Maxillaria stenophylla Rchb. f. 1854. Not
Maxillaria stenophylla F. Lehm. & Kraenzl.
1899. Figure 9E.
Straggly, rhizomatous epiphytes rooted only at the
basal shoots; bract-covered rhizome elongate, to 4 mm
in diameter. Roots fibrous, to 0.5 mm in diameter. Pseu-
dobulbs cylindric to ovoid, somewhat compressed, 1-
foliate, (8)10-30 mm tall, 6-12 mm wide, spaced 1-5
cm apart, subtended and concealed when young by a
pair of subulate bracts. Leaves shortly petiolate; blade
lance-linear, strongly acute, 4-14 cm long, 7-15 mm
wide. Inflorescence fascicled at the base of the pseu-
dobulb, each scape less than 5 mm tall and concealed
by the rhizome bracts; ovary 2-3 mm long. Flowers
membranaceous, white or lip with yellow midlobe. Se-
pals dissimilar, not much spreading, apiculate; dorsal el-
liptic, (3?)6-7 mm long, 2 mm wide; lateral sepals ob-
long-ovate, 6-6.5 mm long, about 2.5 mm wide, about
!/5 connate. Petals linear, 4.5-6 mm long, about 1 mm
wide, apiculate. Lip relatively fleshy, concave, 3-lobate,
2-2.5 mm long excluding the foot; lateral lobes subor-
bicular when viewed laterally; midlobe reflexed, bluntly
acute; callus at the base apparently transverse connecting
the lateral lobes. Column stout, 1-1.2 mm long exclud-
ing the anther, extended at the base into a foot about 1.5
mm long, strongly curved forward.
Epiphytic at 1400-2300 m, often in oak forests
with Chusquea understories. Available specimens
with flowers in good condition were all collected
in August. Known only from Costa Rica from the
Cordillera de Talamanca south of Cartago to the
Panama border, but expected in Panama.
Maxillaria concavilabia has been confused
with vegetatively similar M. neglecta (Schltr.) L.
O. Williams, a low-elevation species with similar
flower color. However, M. concavilabia has small-
er flowers that are more membranaceous in dried
specimens and therefore are difficult to rehydrate.
The lip has pronounced rounded lateral lobes
when viewed laterally, and the column foot is
more antrorsely curved from the base. Similar M.
parviflora (Poeppig & Endl.) Garay is a low-ele-
vation species lacking the rounded lateral lobes.
Maxillaria conduplicata (Ames & C. Schweinf.)
L. O. Williams, Ann. Missouri Bot. Gard. 29:
348. 1942. Ornithidium conduplicatum Ames &
C. Schweinf. Sched. Orch. 8:66-68. 1925.
TYPE: Panama, Chiriqui, Powell 341 (holo-
type: AMES). Figure 5D.
Plant poorly known, probably with canes to at least
50 cm tall. Canes mostly hidden from view by the some-
what inflated leaf sheaths (in dried material). Roots on
canes to about 2 mm in diameter. Pseudobulbs appar-
ently restricted to the base of the plant and formed only
in the juvenile stage, ovoid, 5.5 cm long (n = 1), api-
cally 1 -foliate. Leaves congested or not (n = 2), the
lower abscising early; sheaths rugose, somewhat inflated
in the dried specimen; blade subcoriaceous, oblong, to
20 cm long and 2.7 cm wide, acute to obtuse or 2-lobate.
Inflorescence a fascicle of several single-flowered
scapes in leaf axils; scapes to 4 cm long, concealed by
acute bracts; ovary with pedicel 8-9 mm long, subtend-
ed by a much shorter bract. Flowers globose, light yel-
low, with light pink on outside of sepals and petals; lip
yellow. Sepals somewhat fleshy apically, similar, ovate,
acute; dorsal somewhat concave, 5.9-6.5 mm long, 2.5-
3 mm wide; lateral sepals about 7 mm long, 3 mm wide.
Petals elliptic-ovate, acuminate, about 6 mm long, 2.5-
3 mm wide. Lip somewhat hinged to the column foot,
4-lobate when spread, subquadrate, the basal lobes
somewhat embracing the column and forming a shallow
cup, the apical J/2 somewhat conduplicate in front; callus
at the base a simple thickening connecting the lateral
lobes in front; also an apical, cushion-like callus in front.
Column stout, about 1 .5 mm long excluding the anther;
foot about 1 mm long; anther about 1 mm long.
Maxillaria conduplicata is known from Pana-
ma and Costa Rica at 1200-1500 (3000) m. A live
plant collected in a cloud forest at Cerro de la
Muerte at about 3000 m survived two summers
at Sarasota, Florida, where it flowered in Novem-
ber 1996. Based on this and two available her-
barium specimens, it appears to flower from Sep-
tember to November. It may be much more com-
mon than available specimens would indicate, as
the rather large plant with small, inconspicuous
flowers probably does not catch the eye of collec-
tors.
Maxillaria confusa Ames & C. Schweinf.,
Sched. Orch. 8:57-58. 1925. TYPE: Costa
Rica, Peralta, Lankester 908 (holotype: AMES).
Figure 23C.
Epiphytes rarely forming large clumps. Roots to
about 1 mm in diameter. Pseudobulbs apically 1 -foliate,
truncate, compressed, ovoid, 1-3 cm long, provided ba-
sally with 1-2 foliate bracts. Leaves petiolate, those sub-
tending the pseudobulb constricted above the articula-
tion; petioles to about Va the blade length; blades elliptic -
oblanceolate, cuneate, 3-17 cm long, 1.3-3 cm wide,
acute to obtuse. Inflorescences lateral, several per bract
axil, each covered with 4-5 pale green bracts; scapes 3-
7 cm long; ovary with pedicel 1.5-1.7 cm long. Flowers
white to yellowish aging brownish, the lip with purple
markings. Sepals dissimilar; dorsal elliptic-lanceolate,
concave, 1.8-2.2 cm long and about 5 mm wide, the
lateral sepals triangular-lanceolate, somewhat attenuate,
2-2.5 cm long and 5 mm wide at the base forming a
52
FIELDIANA: BOTANY
conspicuous, pointed chin. Petals lanceolate-falcate,
1.5-2 cm long and about 4 mm wide. Lip articulated
with the column foot, obovate, cuneate, strongly 3-lo-
bate on the apical '/,, 12-15 mm long and about 6 mm
wide when spread; lateral lobes embracing the column;
midlobe verrucose, suborbicular; callus ligulate, thick-
ened and rounded in front, ending just above the middle,
somewhat pubescent. Column stout, about 5 mm long
excluding the anther; foot 6-9 mm long; anther rough-
papillose, about 2 mm long, the anther bed conspicu-
ously fringed; pollinia with short, squarrose stipe and
horseshoe-shaped viscidium. Capsule about 4 cm long.
Rather common yet somewhat inconspicuous
epiphyte of premontane rain forests at (150)500-
1450 m. Flowering January to September, proba-
bly throughout the year. Belize to Panama.
Maxillaria confusa is similar to M. ramonensis
Schltr. but has larger flowers with fringed anther
bed.
Maxillaria costaricensis Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:232-233. 1923.
TYPE: Costa Rica, District of Volis in San Ra-
mon, 1050 m, Brenes 237 (lectotype: CR, not
seen; isotypes: AMES, NY). Figure 1 ID.
Plant a somewhat straggly epiphyte to about 25 cm
high, sometimes forming large clumps. Roots to about
1 mm in diameter. Pseudobulbs fusiform to cylindric;
to 3 cm apart, subtended by nonfoliaceous sheaths, api-
cally 1 -foliate, 1.5-3 cm long, to 1 cm wide. Leaves
thin, usually shortly petiolate, the blade linear-lanceo-
late, 5-14 cm long, 5-9 mm wide, the apex obtuse to
unequally 2-lobate. Inflorescence formed in the flush of
new growth, to about 2 cm long; ovary with pedicel 1.5-
2 cm long. Flowers green to cream or rose (old flow-
ers?) tinted with red, the lip dark maroon. Sepals simi-
lar; dorsal concave oblong, 1-1.3 cm long, about 3-4
mm wide, acute to obtuse and apiculate; lateral sepals
lanceolate to falcate, 1.1-1.4 cm long, 4-4.5 mm wide,
acute. Petals oblanceolate, 1-1.2 cm long, about 3 mm
wide, apex acute to obtuse and somewhat recurved. Lip
simple to 3-lobate, more or less straight with the sides
somewhat embracing the column, elliptic-ovate, 1-1.2
cm long, 4.5-5.5 mm wide; apex truncate to rounded;
callus a fleshy and shiny ligule on the lower Vi. Column
arcuate, 7 mm long excluding the anther, the foot 2 mm
long; anther about 2 mm in diameter. Capsule ellipsoid,
about 1.5-2 cm long.
Epiphytic in rain and cloud forests at 500-1600
m. Flowering January to May and November. En-
demic and apparently restricted to Costa Rica
north of the Central Valley.
Maxillaria costaricensis is recognized by the
rhizomatous habit, usually greenish flowers with
pink or red stain, and maroon lip. Morphologi-
cally it is most similar to M. variabilis Bateman
ex Lindl., but the leaves of M. costaricensis are
usually narrower, the flowers have a different col-
or, and in areas of sympatry the plant grows in
wetter zones. The petals of M. variabilis are more
recurved apically. In dried plants the pseudobulbs
of M. costaricensis tend to curl, while those of
sympatric M. variabilis tend to remain stiff. How-
ever, in Panama, plants with flowers of M. var-
iabilis are seen on plants similar to those of M.
costaricensis. For the moment these are included
in M. variabilis, but the name, M. chiriquensis
Schltr., may be applicable should any segregates
be recognized as distinct.
Maxillaria crassifolia (Lindl.) Rchb. f., Bonplan-
dia 2:16. 1854. Heterotaxis crassifolia Lindl.,
Edwards's Bot. Reg. 12: t. 1028. 1826. TYPE:
Jamaica, J. Lee (holotype: K). Dicrypta crassi-
folia (Lindl.) Lindl. ex Loud., Hort. Brit. Suppl.
3:536. 1839. Epidendrum sessile Sw., Prodr.
Ind. Occ. 122. 1788. TYPE: Jamaica (holotype:
BM, not seen). Maxillaria sessilis (Sw.) Fawc.
& Rendle, Fl. Jamaica 1:120. 1910. Not Max-
illaria sessilis Lindl. 1845. Dicrypta baueri
Lindl., Gen. Sp. Orch. t. 44. 1830. TYPE: Ja-
maica. Figure 17B.
Plant a cespitose epiphyte, sometimes forming large
clumps; pseudobulbs essentially obsolete. Roots to 2
mm in diameter. Leaves 4-5 per shoot, fleshy, shortly
petiolate at the base, conduplicate, the blade linear to
very narrowly elliptic, unequally bilobed at the apex,
somewhat carinate abaxially and canaliculate adaxially
on the midvein, 7-27 cm long, 1.3-3 cm wide, the ad-
axial surface appearing somewhat sunken along major
veins. Inflorescences 1-few single-flowered scapes per
leaf (rhizome bract) axil produced successively; ovary
with pedicel about 1.5 cm long. Flowers barely emerg-
ing from the leaf axil, autogamous in some populations
(Florida), white to yellow, the lip laterally stained or
spotted with purple and with yellow callus. Sepals ellip-
tic to ovate, thickened apically, 13-15(20?) mm long,
4-6 mm wide, acute. Petals oblanceolate to spatulate,
11-14 mm long, 3-4 mm wide, subobtuse. Lip articu-
late with the column foot, constricted below, the apical
l/3 appearing somewhat 3-lobate, the sides inflexed to-
ward the column, 12-15 mm long, 5-6 mm wide; callus
farinose (mealy), sticky, extending from the base to near
the apex, linear, interrupted above the middle and broad-
ened above. Column lightly arcuate, cylindric, broad-
ened above, 9-10 mm long, forming a poorly denned
foot 1-2 mm long.
An epiphyte of larger branches and trunks of
trees in primary or secondary wet forests, rarely
on rocks or embankments, at 5-1300 m. Flower-
ing throughout the year. Widespread throughout
the neotropics; Central America, tropical South
America, Jamaica, Cuba, and south Florida.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
53
This widespread, lowland species is easily dis-
tinguished by the fleshy, conspicuously veined
leaves produced in fans and by the erect, upward-
facing flowers. Schlechter's M. gatunensis (Re-
pert. Spec. Nov. Regni Veg. Beih. 17:68. 1922) is
usually interpreted as a rather robust example
from Panama with sepals 2 cm long, but the rel-
atively short callus indicated in his floral diag-
nosis may be inaccurate. He probably did not ob-
serve the continuation of the callus to the lip apex,
where it is less conspicuous in dried specimens.
Maxillaria cryptobulbon Carnevali & J. T. At-
wood, Novon 1:159-162. 1991. TYPE: Ecua-
dor, Morona-Santiago, Serrania de Cutucii, Ack-
erman 1041, based on Madison s.n. (holotype:
SEL). Figure 20D.
Plant a cespitose epiphyte 30-50 cm tall. Roots to 1
mm in diameter. Pseudobulbs ellipsoid to suborbicular
to oblong, somewhat depressed, 8-12 mm long, 9-15
mm wide, concealed by 2-3 subtending, broad sheaths,
the uppermost of which may bear a blade, the apex 1-
foliate. Leaves long petiolate; petiole (4)8-15(24) cm
long, %-% the blade length; blades broadly elliptic,
somewhat decurved, (20)25-35(42) cm long, (2.3)4.5-7
cm wide, acute, the abaxial surface often glaucous. In-
florescences 1-3 per shoot, borne in the flush of new
growth; scapes 5-10 cm long; ovary with pedicel 3-3.3
cm long mostly or entirely concealed by the floral bract.
Flowers spreading; sepals dull red-brown to maroon-
red; petals pink or yellow-orange within, tinged with
red-brown without; lip creamy yellow with yellow cal-
lus. Sepals similar, oblong to somewhat lanceolate, 2.6-
2.8 cm long, 6-7.5 mm wide, the apex obtuse to round-
ed; dorsal concave; laterals somewhat oblique. Petals
obliquely oblong-lanceolate to elliptic, 2.2-2.5 cm long,
5-6 mm wide, apex acute to obtuse. Lip sparsely pu-
bescent within, ovate in general outline when spread,
1.6-1.8 cm long, 8-9 mm wide, base cuneate, 3-lobate
in the apical V*\ lateral lobes erect around the column
with erose, rounded, or subtruncate apices; midlobe
ovate, obtuse, thickened and fleshy, abaxially keeled, the
margins thin and undulate; callus a thickened ligule,
rounded in front just above the middle. Column arcuate,
7-8 mm long excluding the anther; foot 4-4.5 mm long;
anther 2.5-3 mm long, keeled dorsally.
Epiphytic in premontane rain or cloud forests
at 1200-1400 m. Flowering at least June and July.
Costa Rica, Ecuador, and Venezuela (450 m).
The broad, long petiolate leaves and small, de-
pressed pseudobulb distinguish the species. It is
most similar to M. brunnea Lind. & Rchb. f. but
has much broader leaves and smaller compressed
pseudobulbs, and the flowers are stained with
darker red.
Maxillaria ctenostachya Rchb. f., Card. Chron.
39. 1870. Costa Rica, Veitch (holotype: w). Ca-
maridium ctenostachys (Rchb. f.) Schltr.; Fed-
des Repert. Spec. Nov. Regni Veg. Beih. 19:
238. 1923. ICamaridium arachnites Schltr., Re-
pert. Spec. Nov. Regni Veg. Beih. 17:73-74.
1922. TYPE: Panama, Chiriqui, Powell 210 (is-
otypes (5 sheets): AMES). Figure 8B.
Plant a somewhat straggly, cane-forming epiphyte to
about 1 m tall; shoots of each cane to 20 cm long, each
terminated by a pseudobulb, diminishing in size in succes-
sive shoots. Pseudobulbs ovoid to conic, 1.5-10 cm long,
subtended by 1 or more foliaceous sheaths; apex 2-foliate.
Leaves linear-lanceolate, obtuse, to 25 cm long, 1-1.2 cm
wide, apiculate. Inflorescences forming in the flush of im-
mature growth, l(-more?) per axil of the rhizome bracts;
scape to about 3 cm long; ovary with pedicel about 5-7
mm long, subtended and exceeded by an attenuate bract
1.5-1.7 cm long. Flowers white, sometimes striped or suf-
fused with purple, campanulate. Sepals similar, attenuate,
more than 2 cm long, 3 mm wide. Petals similar to the
sepals, attenuate, 1 .7-2 cm (probably more) long, about 2.5
mm wide. Lip hinged to the column foot, ovate, about 7
mm long and 5 mm wide when spread, 3-lobate on the
upper %; lateral lobes obtuse and exceeding the callus; mid-
lobe recurved, ovate; callus complex, consisting of 5 sharp
keels from the base to about the middle, the outer of which
terminate as sharp teeth, and a fleshy rounded callus near
the divergence of the lateral lobes. Column 5-6 mm long
excluding the anther; foot about 2 mm long.
Epiphytic in cloud forests at 900-1220 m.
Flowering at least September and October. Costa
Rica and Panama.
The spidery flowers borne on long canes with
apically 2-foliate pseudobulbs are unique in Costa
Rica. Maxillaria ctenostachya is poorly known
and seldom collected.
Maxillaria dendrobioides (Schltr.) L. O. Williams,
Ann. Missouri Bot. Card. 27:283. 1940. Camar-
idium dendrobioides Schltr., Beih. Bot. Centralb.
36:415. 1918. TYPE: Costa Rica, San Ramon,
1500-1600 m, Tonduz 17620 (holotype: CR). Ca-
maridium jimenezii Schltr., Beih. Bot. Centralb.
36:416. 1918. TYPE: Costa Rica, La Palma,
Werckle 855 (holotype: CR). ?Camaridium simile
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
239. 1923. TYPE: Costa Rica, San Ramon, 1025
m, Brenes (VII) 64 (isotype: NY). Figure 3C.
Plant a shrubby epiphyte to about 40 cm tall on usu-
ally large branches; rhizomes forming indeterminate
canes, entirely concealed by sheaths, strict when small,
decumbent when large; sometimes with a few branches
diverging at acute angles with the main stem. Roots fi-
brous, white, mostly forming at the base of the plant and
infrequently on the canes, rarely to 1 mm in diameter.
54
FIELDIANA: BOTANY
Pseudobulbs absent (juveniles?). Leaves densely distich-
ous; sheath orangish to greenish and spotted with purple
in dried material, with hyaline margins; blade coria-
ceous, 1-6 cm long, 5-10 mm wide, elliptic when short,
lanceolate when long, the base clasping the stem, the
apex unequally 2-lobate. Inflorescence single per leaf
axil, mostly concealed by subulate bracts; scapes 1-2.5
cm long; ovary and pedicel 4-5 mm long, exceeded by
the subtending floral bract. Flowers yellow to greenish,
shading to red or salmon, the segments not spreading
broadly. Sepals 8-13 mm long, 2-3.5 mm wide; dorsal
elliptic-lanceolate, acute; lateral sepals falcate-lanceolate
and acute. Petals elliptic to oblanceolate or obovate,
6.5-12 mm long, 2-3 mm wide, acute. Lip 3-lobate to
broadly pandurate in outline, 4-7 mm long, 2.5-3 mm
wide, obtuse; callus a fleshy ligule on the lower Vi, trun-
cate to rounded. Column 3-3.5 mm long excluding the
anther; foot 1-2 mm long. Capsule globose, less than 1
cm long.
Epiphytic and common on larger branches of
trees in cloud forests at 800-2400 m. Often seen
on remnants in pastures. Flowering year around.
Costa Rica and western Panama.
As represented by the composite description
above, M. dendrobioides is probably a complex
of species. Maxillaria dendrobioides (sensu stric-
to) has a distinctly three-lobed lip with the mid-
lobe more or less ligulate in shape. Plants with
more or less pandurate lip, i.e., the base and apex
each with a pair of angled lateral lobes and shorter
leaves have been segregated as Camaridium sim-
ile. Camaridium jimenezii is supposed to have the
lip apex constricted, but this feature was not ob-
served in any of the specimens examined and is
probably an artifact. Vegetatively, M. valerioi
Ames & C. Schweinf. is identical with the com-
plex but has crystalline white flowers with a dis-
tinctly spatulate lip.
Maxillaria dichotoma (Schltr.) L. O. Williams,
Bot. Mus. Leafl. 9:15. 1940. Camaridium di-
chotomum Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 8:98. 1921. TYPE: Pincincha, in re-
gione subtropica vallis Nanegal, Sodiro 94 (not
found). Maxillaria paleata of authors. Not
(Rchb. f.) Ames & Correll. Figure 14B.
Plant a straggly, pendent or decumbent epiphyte to
about 1 m tall comprised of several successive shoots
anchored only at the base; each shoot to 18 cm long,
terminated by a pseudobulb. Roots white, to less than 1
mm in diameter. Pseudobulbs ovoid, compressed, 1.5-
4 cm long, 1-2.5 cm wide, subtended when young by
1-3 foliaceous sheaths; apex 1 -foliate. Leaves both api-
cal and along the rhizome; foliate bracts broad, narrow-
ing abruptly above the articulation to form a slender
petiole; apical leaf petiolate; all blades elliptic, 2.5-20
cm long, 1.5-5 cm wide, acute or acuminate. Inflores-
cence short, lateral, single flowered, produced preco-
ciously on the canes of developing snoots; scapes to
about 1.5 cm long, concealed by rhizome bracts; ovary
with pedicel 7-9 mm long, subtended by a cucullate,
acute or subulate floral bract 1.3-2.1 cm long. Flowers
variable in color, mostly yellow or less often white
tinged with purple or pink. Sepals similar; dorsal ellip-
tic, 1.4-1.7 cm long, about 5.5 mm wide, acute; lateral
sepals obliquely ovate-lanceolate, 1.4-1.8 cm long,
about 6 mm wide. Petals elliptic, 1.3-1.6 cm long, 3.5-
5.5 mm wide. Lip hinged to the column foot, 3-lobate
below the middle, strongly arched, 8-10 mm long, about
5 mm wide with side lobes spread; midlobe fleshy,
ovate; callus fleshy at the base terminating at the sinuses.
Column stout, 4-5 mm long excluding anther; foot
about 3 mm long; anther about 2 mm in diameter.
Epiphytic on the larger branches and trunks of
trees in premontane and lower montane evergreen
forests at 500-1700 m. Flowering at least August
to April and probably throughout the year. Costa
Rica and Panama to Colombia, Ecuador, and Peru;
doubtfully in Nicaragua as reported.
This little-known species has the broadest range
among its relatives, M. minor (Schltr.) L. O. Wil-
liams, M. paleata (Rchb. f.) Ames & Correll, M.
schlechteriana J. T. Atwood, M. sigmoidea (C.
Schweinf.) Ames & Correll, and M. vaginalis
Rchb. f.
Maxillaria diuturna Ames & C. Schweinf.,
Sched. Orch. 8:58. 1925. TYPE: Panama, C. W.
Powell 376 (holotype: AMES). Figure 16B.
Plant a straggly, usually pendent epiphyte with suc-
cessively borne snoots developing 3-10 cm apart, with
only the lowermost shoots forming roots. Roots white,
rather large, to 1 .5 mm in diameter. Pseudobulbs com-
pressed and sharp edged, 2-3.5 cm long and 1.5-2.5 cm
wide, apically truncate to emarginate 1 -foliate, sheathed
at the base with 2-4 leaf-bearing sheaths. Leaves shortly
petiolate, elliptic-oblong, acute to obtuse, rarely round-
ed, the apical leaf 5.5-12 cm long and 1.5-3.5 cm wide.
Inflorescences appearing singly within each rhizome
bract of the immature shoot, to about 4 cm long, covered
by few bracts below the ovary but mostly hidden from
view by the rhizome bracts; ovary with pedicel much
longer than the subtending floral bract, 2-2.5 cm long.
Flowers clear yellow to greenish yellow, the lip some-
times red at the base and the column with a few red
spots on the lower side. Sepals similar, 1.4-1.5 cm long,
6-7 mm wide, acute; dorsal ovate-lanceolate; lateral se-
pals obliquely ovate, forming a mentum at the column
base. Petals ovate, 1.1-1.2 cm long and 5-7 mm wide,
acute to obtuse. Lip hinged to the column foot, ovate-
pandurate, vaguely 3-lobulate, apically rounded and
constricted at about the apical Vi, 10-12 mm long, 5-6
mm wide; callus narrowly ovate and obtuse, fleshy,
lightly 3-ridged on the lower lh. Column arcuate, to 8
mm long including the 3-mm foot. Capsule ellipsoid to
2.5 cm long.
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
55
Fairly common epiphytic or rarely terrestrial
herb in rain forests at 50-1000(1600?) m. Flow-
ering throughout the year. Nicaragua to Panama
and probably Colombia.
The ascending shoots rooted only at the base
of the plant and sharp-edged, apically truncate to
emarginate, 1 -foliate pseudobulbs are unique fea-
tures among Central American maxillarias. It is
perhaps most closely related to M. nagelii L. O.
Williams, a species from Mexico.
Maxillaria dressleriana Carnevali & J. T. At-
wood, Lindleyana 11:29-31. 1996. TYPE: Pan-
ama, Chiriqui, flowered in cultivation at the
Marie Selby Botanical Gardens, 3 Jan. 1995,
Carnevali 3858 (holotype: SEL; isotypes: AMES,
INB, MO). Figure 2 ID.
Plant a cespitose, epiphytic herb 25-30 cm tall. Rhi-
zome short, clothed by scarious sheaths. Pseudobulbs
dull dark green, tinged with dull dark maroon basally in
older pseudobulbs, clustered, 2.5-4 cm tall, 1-1.5 cm
wide and thick, ovoid or ovoid-ellipsoid to somewhat
conic; subtended by nonfoliate sheaths; apex 1 -foliate.
Leaves erect on the pseudobulb, coriaceous, elliptic or
rarely ovate -elliptic, 24-27 cm long, 2-3 cm wide, pet-
iolate, acute. Inflorescence erect, shorter than pseudo-
bulb; scape 8-12 mm long, terete; ovary with pedicel
1.4-1.9 cm long, subtended by a floral bract 1.7-2.1 cm
long, 1.1-1.5 mm wide. Flowers deep dull red-maroon,
petals paler with a yellowish cast, lip dark red-maroon,
apex dull deep yellow, column dull pale cream-yellow.
Sepals somewhat fleshy; dorsal slightly concave and
subparallel to the column, oblong-elliptic, 1.5-1.6 cm
long, 6.5-7 mm wide; apex obtuse-rounded; margins
somewhat revolute; lateral sepals widely spreading,
somewhat oblique, 1.5-1.7 cm long, 5-6 mm wide; apex
obtuse or obtuse-rounded; margins revolute. Petals sub-
parallel to column, narrowly obovate or narrowly obo-
vate-oblong, 1.4-1.5 cm long, 4.55 mm wide; apex
acute or obtuse-rounded; margins somewhat revolute.
Lip 3-lobate, 1.4-1.6 cm long, 7.5-8.5 mm wide when
expanded; lateral lobes falcate-uncinate, acute, emerging
just below the middle portion of the lip, 1-1.5 mm long,
ca. 1 mm wide at base; midlobe subquadrate-oblong or
subquadrate-pandurate, somewhat broadened apically,
7.5-9 mm long, 4.5-5.5 mm wide; apex subtruncate,
shallowly emarginate; disc with a farinose yellow callus
reaching midlength of the midlobe. Column arcuate,
broadly winged apically, 8-10 mm long, 4-5 mm wide;
column foot 4-5 mm long; anther smooth.
Poorly known ecologically; 80 m. Flowering
November. Endemic to Costa Rica and western
Panama.
Within the M. rufescens Lindl. complex in Cos-
ta Rica, the dull, dark maroon flowers are unique.
Maxillaria elatior (Rchb. f.) Rchb. f., Ann. Bot.
Syst. 6:532. 1863. Dicrypta elatior Rchb. f.,
Linnaea 18:403. 1844. TYPE: Guatemala, Que-
zaltenango, Leibold 14 (not seen). ICamari-
diwn xylobiichilum Kraenzl., Vidensk. Medd.
Nat. Copenhagen 71:174. 1920. TYPE: Local-
ity unknown (drawing of type: AMES). Maxil-
laria triangularis Lindl., Edwards's Bot. Reg.
31: Misc., p. 9. 1845. TYPE: Guatemala Hart-
weg s.n. (holotype: K). Maxillaria albertii
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
229. 1923. TYPE: Costa Rica, Alajuela, Rio Je-
sus de San Ramon, Brenes 257 (lectotype: CR;
isolectotype: AMES). Figure 7F.
Plant large to 1 m tall, forming conspicuous canes,
straggly, pendent when large; shoots comprising each
cane forming 2-10 cm apart. Roots white, large, to 2
mm in diameter. Pseudobulbs subtended by 1 -several
leaf-bearing rhizome bracts, apically 1-2-foliate, ovate,
smooth, compressed, 4-8 cm long, 2-3 cm wide. Leaves
petiolate, the blade coriaceous, 5-30 cm long, 2-4 cm
wide. Inflorescence 1 per rhizome bract axil of imma-
ture shoots, to about 5 cm long, excluding the pedicel
and ovary, covered by 3 or more acute, imbricating
sheaths; ovary with pedicel 3.2-5 cm long, far exceed-
ing the subtending floral bract. Flowers orange to red
or greenish yellow flushed with red. Sepals ovate-lan-
ceolate, 2-2.6 cm long, 7-10 mm wide, acute; lateral
somewhat oblique. Petals falcate-lanceolate, about 1.8-
2.2 cm long, 4-5.5 mm wide. Lip simple when spread,
elliptic-ovate, slightly 3-lobulate just below the apical
V3, about 1.7 cm long and 9-10 mm wide, the sides
somewhat infolded around the column; apex tapering,
somewhat obtuse, recurved; callus a broad, thickened,
biblike ligule on lower V2-2/3, rounded to slightly V-
shaped in front. Column arcuate, about 1.2 cm exclud-
ing the anther; foot 3-4 mm long; pollinia 4, supported
on a saddle-shaped viscidium and a short, squarrose
stipe. Capsule ellipsoid, 4-5 cm long.
Epiphytic on larger branches of evergreen for-
ests, apparently tolerant of various elevations
(400-1500 m). Flowering September to April.
Mexico to Costa Rica.
The type of Dicrypta elatior Rchb. f. (Leibold
14} was not found, but a specimen so labeled in
Reichenbach's handwriting (Leibold 44) indicates
that the collection number may have been mis-
printed.
Costa Rican M. elatior was originally described
as M. albertii by Schlechter, who compared it with
M. houtteana Lindl., not M. elatior. Over the
whole range, pseudobulbs may have one or two
apical leaves, and specimens from Belize show
the short rhizomes between pseudobulbs as seen
in Costa Rica. The only remaining difference be-
tween Costa Rican plants and those to the north
seems to be the tendency for narrower leaves in
Costa Rica.
56
FIELDIANA: BOTANY
Maxillaria endresii Rchb. f., Gard. Chron. n.s.
26:680. 1886. TYPE: Costa Rica, Edward Low
s.n. (holotype: w). Figure 20B.
Plant a densely cespitose epiphyte forming large
clumps. Roots white, large to 2 mm in diameter. Pseu-
dobulbs sessile, 3.5-6 cm long and 2.5-4 cm wide,
shiny but strongly ridged in the dried specimen, com-
pressed, ovoid to orbicular, and apically obtuse, truncate
or emarginate, 1 -foliate, subtended and covered when
young by a pair of acute sheaths to 7 cm long. Leaves
large, the long petiole to 1A the blade length; blade ellip-
tic-oblong, 15-35 cm long and 4-7 cm wide, acute to
obtuse. Inflorescence to about 15 cm, 1 -flowered per
rhizome bract forming on mature shoots, sheathed by 4-
7 bracts; ovary with pedicel 2.5-3 cm long. Flowers
strongly scented, variable in floral segment attenuation,
the perianth yellow within, lighter on the abaxial side,
the lip bright yellow with a white margin, the lateral
lobes striped with purple. Sepals similar, acute; dorsal
oblong, erect 5.5-7 cm long, 9 mm wide, the lateral
sepals lanceolate, falcate, 5.5-8 cm long, 1 cm wide.
Petals light yellow, smaller than the sepals, broadened
at the base, linear-lanceolate, 4.5-6.5 cm long, 7 mm
wide, attenuate. Lip abruptly 3-lobate at the middle, pu-
berulent centrally, 2.5-3 cm long, 1.2 cm wide, the rath-
er narrow lateral lobes overlapping the large, orbicular,
recurved midlobe; callus simple, extending to just below
the middle, more or less V-shaped in front. Column
white, lined with purple, cylindric, somewhat arcuate, 2
cm long including the 8-mm-long foot; anther bed ma-
genta, coarsely ciliate; anther cap brownish, 3 mm long.
Capsule ellipsoid, to 3 cm long.
Epiphyte of larger trunks in evergreen primary
forests at 50-700 m. Flowering March to May
and September. Nicaragua to Panama; possibly
into South America, where specimens may be
misidentified as M. luteoalba Lindl.
The truncate, rounded pseudobulbs with large
leaves supported on long petioles and large, fra-
grant flowers with attenuate floral segments are
good features for field recognition.
Maxillaria endresii has been confused with the
South American M. luteoalba Lindl., which has a
lip with broader side lobes and a nonorbicular
pseudobulb. It is also similar to M. angustiseg-
menta Ames, Hubbard & C. Schweinf., a higher
elevation species from Costa Rica and Panama
with subconic pseudobulbs, shorter perianth seg-
ments, and broader side lobes.
Maxillaria exaltata (Kraenzl.) C. Schweinf., Hot.
Mus. Leafl. 11:272. 1945. Camaridium exalta-
tum Kraenzl., Bot. Jahrb. Syst. 37:386-387.
1906. TYPE: Peru: Department of Loreto,
mountains east of Moyobamba, Weberbauer
4620 (photo of type: AMES). Figure 6C.
Plant a somewhat straggly epiphyte to about 2 m tall,
forming upright or decumbent canes to about 1 cm in
diameter, concealed by rugose sheaths. Roots to about 1
mm in diameter. Pseudobulbs reduced to slight swelling
near the stem apex or essentially absent. Leaves sub-
coriaceous, linear-lanceolate, increasing in size near the
stem apex, to about 20 cm long and 2.5 cm wide, acute.
Inflorescences several per leaf axil, the peduncles about
5 cm long and concealed by bracts; ovary with pedicel
1.5-1.8 cm long, exceeded by the subtending somewhat
inflated floral bract. Flowers white or cream, lip pink to
red-violet with yellow midlobe; segments not broadly
expanded. Sepals dissimilar, the dorsal oblong, 1.8-2 cm
long, 5-6 mm wide, acute; lateral sepals obliquely in-
serted on the column foot, oblong to triangular, 2.2-2.4
cm long, about 5 mm wide, acute. Petals oblong, some-
what falcate, oblique at the base, 1.6-1.8 cm long, 3-
3.5 mm wide, acute. Lip somewhat obovate when
spread, 1.2-1.4 cm long, 5.5-6.6 mm wide, 3-lobate on
the apical V4; side lobes somewhat clasping the column;
midlobe fleshy, verrucose, bluntly acute to obtuse, about
2.5 mm long and wide. Column 6-7 mm long excluding
the anther; foot 7-7.5 mm long; anther verrucose, 2.5
mm long.
Epiphytic in cloud forests at 650-1500 m.
Flowering in Central America September to
March and July. Peru (type) to Costa Rica.
Maxillaria exaltata is a member of the M. mer-
idensis Lindl. complex, characterized by the up-
right canes, which are somewhat swollen termi-
nally and concealed by rugose sheaths. This spe-
cies differs from M. meridensis by the much larg-
er plant size and by the lip, which has a nearly
orbicular midlobe. It is also a species of lower
elevations, and M. meridensis flowers March to
August (December). Both species have been seen
growing within a few meters of one another at
Tapantf.
Maxillaria falcata Ames & Correll, Bot. Mus.
Leafl. 11:15. 1943. Ornithidium costaricense
Schltr., Repert. Spec. Nov. Regni Veg. 8:456.
1910. TYPE: Costa Rica, Pittier (Tonduz?)
2177 (lectotype selected here: AMES 24213).
Not Maxillaria costaricensis Schltr. Figure 7C.
Plant an erect to decumbent epiphyte or terrestrial to
about 1 m high, lacking pseudobulbs (juveniles?); canes
monopodial, erect, branched, sometimes fasciculately so,
concealed by leaf sheaths. Roots to about 1 mm in di-
ameter. Leaves usually deciduous along the lower cane,
with rugose sheaths; blade above the articulation thin,
conduplicate, oblong-lanceolate, abaxially keeled, 3.5-
16 cm long, 7-23 mm wide, the apex unequally 2-lobate.
Inflorescences 2 or more per leaf axil, shorter than the
subtending leaf; scape about 3 cm long; ovary with ped-
icel 1-1.2 cm long, subtended by a somewhat cucullate
floral bract to about 1.7 cm long and with hyaline mar-
gins. Flowers white or pinkish. Sepals similar, oblong,
1.8-2.4 cm long, 4-5.5 mm wide, acute. Petals elliptic-
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
57
oblanceolate to somewhat falcate and attenuate, 1 .7-2.4
cm long, 5-6 mm wide. Lip rigidly attached to the col-
umn, 3-lobate on the apical 1A-1A, 5.5-6.5 mm long,
about 4 mm across the lateral lobes when spread; lateral
lobes falcate, rounded, longer than the subacute, subov-
ate midlobe; callus from the base to above the middle,
low and lamellate at the base, high and 3-keeled in front.
Column longer than the lip, cylindric, slightly arcuate,
7.5-8 mm long excluding the anther; foot about 1 mm
long.
Epiphytic or terrestrial on steep embankments
at 1800-2400 m. Flowering February and March.
Costa Rica and Panama.
This species is easily distinguished from similar
ones by the relatively large lateral lobes of the lip,
which are exceeded by the column. Lips are often
lacking, probably having been eaten by insects.
The drawing at AMES of Ornithidium costari-
cense is labeled as based on Tonduz 2177, not
Pittier 2177. A notebook at us includes the col-
lections of both Tonduz and Pittieri (and Biolley)
within the same institutional numbering system;
therefore, the confusion is understandable. The
original Latin description agrees well with the
drawing, which is all that has been found of the
original materials.
Maxillaria flava Ames, Hubbard & C. Schweinf.,
Bot. Mus. Leafl. 3:41. 1934. Ornithidium ra-
monense Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 19:243-244. 1923. TYPE: Costa Rica,
San Pedro de San Ramon, Brenes 137 (photo-
graph of type: AMES). Not Maxillaria ramonen-
sis Schltr. Figure 12D.
Plant an erect or pendent, cane-forming epiphyte to
about 30 cm tall, rooted only on the lower shoots; stems
much elongate, to about 7 cm long between pseudobulbs
m the lower shoots, diminishing in successive shoots.
Roots to about 1 mm in diameter. Pseudobulbs conic
and rather truncate, clearly and regularly furrowed in
dried specimens, 1-3 cm long, subtended by 1-2 foliate
sheaths, the apex 1 -foliate. Leaves similar, those sub-
tending the pseudobulb with sheath; apical leaf without
petiole; blades thin, elliptic, 1-7 cm long, 4-14 mm
wide, rounded to emarginate, abaxially keeled. Inflores-
cence 1 per rhizome bract of both mature and immature
shoots; 8-20 mm long; ovary with pedicel 5-8 mm long,
somewhat exceeded by the subtending acute, floral
bract. Flowers not opening widely or with sepals and
petals recurved, greenish yellow. Sepals subequal, ellip-
tic-lanceolate, 9-12 mm long, 1.5-2.5 mm wide; laterals
subfalcate. Petals elliptic-lanceolate, 7.5-10 mm long,
2-2.5 mm wide, acute. Lip hinged with the column foot,
convex, ovate when spread, 3-4.5 mm long, about 2 mm
wide, 3-lobate on the lower quarter, recurved in natural
position; lateral lobes shorter than the callus, short and
acute; midlobe ovate, rounded to emarginate; callus
fleshy, rounded and somewhat elevated in front, termi-
nating below the middle. Column stout, arcuate, some-
what tapering, 2.5-3 mm long excluding the anther; foot
about 1 mm long; anther less than 1 mm in diameter.
Epiphytic in cloud forests at 700-1350 m.
Flowering October and November. Endemic to
Costa Rica and Panama.
Maxillaria flava is similar to M. microphyton
but unique in the distinctly furrowed pseudobulbs.
The flowers are apparently always yellow, lacking
the red stripes or suffusion seen in related species;
hence the name. The short lateral lobes of the lip
exceeded by the callus readily distinguishes this
from M. microphyton Schltr.
Maxillaria friedrichsthalii Rchb. f., Bot. Zeitung
(Berlin) 10:858. 1852. TYPE: Guatemala,
Chontales, in Monte Aragua, Friedrichsthal
(isotype: AMES). Maxillaria turialbae Schltr.,
Beih. Bot. Centralbl. 36:414-415. 1918. TYPE:
Costa Rica, Turialba, F. C. Lehmann 1098
(drawing of type: AMES). Figure 8C.
Plant a loosely straggly epiphyte, or if cespitose then
formed of a cluster of successively borne shoots. Roots
fibrous, white, to 1 mm in diameter. Pseudobulbs ob-
long, 1-3 cm apart, 3-5 cm tall and 6-15 mm wide,
somewhat laterally flattened and with several vertical
ribs more conspicuous when dried, subtended by 1-2
early deciduous blade-bearing bracts, apically 2-3-foli-
ate. Leaves linear to oblong, 3.5-15 cm long and 3.5-
12 mm wide, unequally 2-lobed apically. Inflorescence
lateral, borne singly per rhizome bract axil, the pedun-
cles 1.5-3 cm long covered by 5-7 bracts; ovary with
pedicel 10-15 mm long, entirely concealed by a sub-
tending pair of bracts, the uppermost 11-17 mm long.
Flowers with a slight urinous odor, coriaceous, not
opening fully and difficult to spread, white to yellowish,
turning greenish with age, the lip often yellow centrally
and flushed or rarely spotted lavender laterally. Sepals
similar, oblong-lanceolate, concave, 15-20 mm long,
3.5-5 mm wide, acute; lateral sepals falcate, adnate to
the column foot. Petals falcate, elliptic-oblong, 13-18
mm long, 2-3 mm wide, acute. Lip hinged with the
column foot, elliptic oblong, very slightly 3-lobate, 12-
15 mm long; callus linear, basal, extending to about the
middle. Column 9-11 mm long, including the short
foot; pollinia with long stipe, anther bed ciliate.
Epiphytic in wet forests at 50-1500 m. Flow-
ering mostly November to March. Mexico (Cam-
peche), Belize, and Guatemala to Panama.
This species is usually confused with M. acian-
tha Rchb. f. and M. scorpioidea Kraenzl. It has
the smallest flowers of the three species, is the
most frequently seen, and has a broader range and
perhaps ecological tolerance in Central America
than the other two. Schlechter moved his own
species, M. turialbae, into synonymy with M.
58
FIELDIANA: BOTANY
friedrichsthalii, and the floral dimensions given in
his original description fit this concept exactly, not
with M. aciantha or M. scorpioidea.
Maxillaria fulgens (Rchb. f.) L. O. Williams,
Ann. Mo. Bot. Card. 28:425. 1941. Ornithidium
fulgens Rchb. f., Beitr. Orch. Centr.-Am. 76.
1866. TYPE: Costa Rica, "Naranjo," Wend-
land (holotype: w; photo of type: AMES). Figure
6A.
Plant a robust, shrublike, or cane-forming epiphyte to
about 70 cm tall, with 1 -several branches anchored only
at the base, each branch consisting of 1 or more shoots
produced successively 10-20 cm apart and terminating
in a 1 -foliate pseudobulb. Roots large, to 3 mm in di-
ameter. Pseudobulbs to about 3 cm long, conic, sub-
tended and hidden by 2-5 blade-bearing bracts. Leaf
blades coriaceous, lanceolate, to narrowly elliptic, some-
what petiolate above the articulation, 9-20 cm long,
1 .5-4.5 cm wide, acute. Inflorescence a fascicle of 2-8
single-flowered scapes produced in the bract axils of the
developing vegetative bud, each scape 2-3 cm long;
ovary with pedicel 11-15 mm long. Flowers globose,
the segments not spreading widely, bright red, lip yel-
low. Sepals broadly triangular-ovate, 7 mm long, 4-4.5
mm wide, acute to obtuse. Petals elliptic-ovate, slightly
clawed at the base, 6 mm long, 2.5-3 mm wide, acute.
Lip rigidly attached to the column, 5 mm long in natural
position, difficult to spread, constricted in the middle,
saccate on the basal half, the apex deflexed. Column
stout, 2.5 mm long; foot about 2 mm long.
Epiphyte of the larger branches in wet forests
at 100-2000 m. Flowering June to December,
February, April, and May. Nicaragua to Panama,
and northern South America.
This species is one of the most remarkable for
its paedomorphic habit of producing several flow-
ers per bract axil of a vegetative shoot barely be-
yond the bud stage. The multiple flowers appear
superficially to arise from rhizome bract axils of
the already developed shoot.
Maxillaria gomeziana J. T. Atwood, Lindleyana
11:202-204. 1996. TYPE: Costa Rica, Province
of Puntarenas, Goto Brus, 1600 m. Mar. 1995,
Atwood, C. & J. Luer ex Atwood 5052 (holo-
type: SEL). Figure 16D.
Plant a somewhat ascending, cane-forming epiphyte
to about 30 cm tall, comprised of shoots 5-10 cm long
including the pseudobulb. Roots £ 1 mm in diameter.
Pseudobulbs fusiform, compressed, 1 -foliate, 3.5-5 cm
long, 1-1.5 cm wide, subtended by 1-4 strongly rugose
sheaths, the uppermost with soon deciduous blades.
Leaf blades shortly petiolate, subcoriaceous, linear-ob-
long, 12-28 cm long, 1-1.5 cm wide, the apex unequally
2-lobate. Inflorescences 1-3 per foliaceous sheath axil;
scapes 2.5-4 cm long; ovary with pedicel 11-15 mm
long, trigonous, subequal to the subtending floral bract.
Flowers with somewhat spreading segments, pale yel-
lowish green, lip marked with red apically and on side
lobes. Sepals similar, oblong to oblanceolate, 1.7-2.3
cm long, 4-5 mm wide, acute; dorsal somewhat keeled
dorsally and apically; lateral sepals recurved. Petals el-
liptic-oblanceolate, 1.6-1.8 cm long, 3-4 mm wide,
acute. Lip hinged to column foot, 3-lobate below the
middle, 1-1.2 cm long, 6-8 mm wide when spread; lat-
eral lobes rounded, somewhat erect; midlobe fleshy, ob-
long-obovate with recurved lateral margins in natural
position, rhombic when spread, rounded, abaxially
keeled; callus fleshy, thickened and rounded in front,
ending below sinuses between lobes, with a blunt central
and 2 less distinct lateral keels. Column stout, some-
what arcuate, 5-6 mm with anther, 3.5-4 mm without
anther, foot about 2 mm long. Capsule 3-winged.
Poorly known epiphyte at (200)400-1620 m.
Flowering at least August, October, and February;
flowering in cultivation at SEL October to Decem-
ber. Native to Panama and adjacent Costa Rica.
Only recently has a living plant of M. gomeziana
been observed in flower, hence the delay in its de-
scription. It is closely related to M. bracteata
(Schltr.) Ames & C. Schweinf., with which it shares
the central keel of the callus, but has narrower
leaves, more slender pseudobulbs, more slender
perianth segments, and trigonous ovaries that de-
velop into trialate capsules. (See Addendum, p.
173.)
Maxillaria haberi J. T. Atwood, Selbyana 16:
245. 1995. TYPE: Costa Rica, Province of He-
redia, forest between Rio Peje and Rio Sardi-
nalito, Atlantic slope of Volcan Barva, 700-750
m, 4 Apr. 1986, M. H. Grayum 6769 (holotype:
CR). Figure HE.
Plant a conspicuously rhizomatous epiphyte to at
least 50 cm tall, probably pendent when large; rhizome
6-8 mm in diameter, 7-10 cm long between pseudo-
bulbs, concealed when young by acute, nonfoliaceous
bracts. Roots slender, to 1 mm in diameter. Pseudobulbs
ovate, compressed, subtended by 1-3 leaves, 3.5-5 cm
long, 1.5-2 cm wide, with 1 -foliate apex. Leaves ellip-
tic-oblanceolate, conspicuously petiolate, the blade 12-
22 cm long, 4-6 cm wide, acute. Inflorescence appear-
ing in the flush of new growth, scapes 4-5 cm long,
ovary 10-13 mm long, somewhat exceeded by the sub-
tending floral bract. Flower somewhat spreading, yellow
to orange or tan, the lip maroon-red near base, white
near apex. Sepals similar, elliptic-lanceolate, 3.8-4.1 cm
long, 1 cm wide, acute. Petals oblanceolate, about 3.7
cm long, 6 mm wide, acute. Lip rather fleshy, obovate
in general outline, 2.5 cm long, 1.8 cm wide when
spread, 3-lobate on the upper W, lateral lobes embracing
the column; midlobe subquadrate, obtuse; callus of 3
keels from the base to below the sinuses of the lateral
lobes, the middle keel lower and less distinct than the
ATWOOD & MORA DE RET AN A: FLORA COSTARICENSIS
59
outer. Column slightly arcuate, 1 .8-2 cm long excluding
unknown anther, with a swelling on the lower 1A ventral
side.
Epiphytic at 850 m. Flowering at least April,
July, and September. Endemic to Costa Rica;
known only from the Penas Blancas Valley near
Monte verde.
Maxillaria haberi is known only from three
collections made between 1986 and 1989: Gra-
yum 6769 (CR), Bello & Cruz 1023 (INB), and Ha-
ber & Cruz 7474 (MO). Apparently a rare species;
recent attempts to relocate it have failed.
It is most readily distinguished by the combi-
nation of long rhizomatous habit with 1 -foliate
pseudobulbs terminating each shoot, the large el-
liptic-oblanceolate leaves, and the relatively large
yellow flowers. The plant appears similar to that
of M. planicola C. Schweinf. but has foliaceous
sheaths at the base of the pseudobulbs. The flow-
ers are also large as in M. planicola, but the lip
is proportionately longer and has a callus of three
keels rather than of stiff hairs and a ligule.
Maxillaria hedwigiae Hamer & Dodson, Ic. PI.
Trop. 8: t. 800. 1982. TYPE: Guatemala, Rio
Dulce, near Lake Isabal, Hamer A 161 (holo-
type: SEL). Figure 2 1C.
Plant a cespitose epiphyte, sometimes forming large
clumps. Roots large, to 2 mm in diameter. Pseudobulbs
rugose, ovoid-conic, 1 -foliate, to about 4.5 cm long and
2.5 cm wide, sheathed basally when young by 4-6
acute, nonfoliate bracts. Leaf thin, petiolate, the blade
narrowly elliptic, to 36 cm long and 3.5 cm wide (usu-
ally smaller), acute. Inflorescence 3-5 cm long with 3
or 4 acute bracts about 1 .5 cm long; ovary with pedicel
2.2-3 cm long, exceeding the subtending floral bract.
Flowers white to cream or greenish, or probably turning
greenish with age; lip bright orange. Sepals elliptic, 17-
19 mm long, 7-9 mm wide. Petals elliptic, 15-17 mm
long, 6-7 mm wide. Lip 3-lobate at about the middle;
lateral lobes erect, lanceolate; midlobe very fleshy,
somewhat lanceolate but with a truncate to retuse apex;
callus spatulate, extending to about the middle. Column
arcuate, clavate, about 9 mm long, with a short foot
about 1 mm long and winged on the ventral side of the
upper half. Capsule ellipsoid, 2 cm long.
Epiphytic at 0-650 m. Flowering (October)
November to March. Locally common from
(Mexico?) Guatemala to Costa Rica.
The white to greenish white or cream flowers
with distinct orange lip are the best features for
recognition. Also, the midlobe is usually lanceo-
late and more or less acute, unlike its relatives,
M. acutifolia Lindl. and M. morales ii Carnevali &
J. T. Atwood, which have truncate apices.
Maxillaria horichii Senghas, Orchidee (Ham-
burg) 28:13. 1977. Figure 3. TYPE: Costa Rica,
"near Sisica, between La Palma de San Isidro
del General and Play a Dominica," Horich s.n.
(holotype: HEID, not seen). Sepalosaccus hori-
chii (Senghas) Senghas, Orchideen 29:1745.
1994. Figure 10A.
Plant a straggly, rhizomatous epiphyte rooted only on
the lower shoots; rhizome elongate, to 5(6) mm in di-
ameter, concealed by bracts, pendent, forming shoot sys-
tems to about 50 cm long. Roots fibrous, rarely more
than 1 mm in diameter. Pseudobulbs cylindric to ovoid,
compressed, subtended by 2-3 subulate, nonfoliaceous
bracts, 2-3 cm long, 1.2-2 cm wide, apically 1 -foliate,
spaced 3-9 cm apart on the rhizome. Leaves shortly
petiolate; blade lanceolate, 8-19 cm long, 1.2-2.8 cm
wide, acute. Inflorescences in fascicles borne at the base
of the recently matured pseudobulb; scapes to about 1.5
cm long, mostly concealed by the rhizome bracts; ovary
and pedicel about 5 mm long, concealed by inflated,
ovate scape bracts. Flowers bright orange, column
sometimes (always?) white. Sepals dissimilar, not open-
ing widely; dorsal elliptic-ovate, concave, 7-9 mm long,
3-4 mm wide; lateral sepals ovate, about % connate,
7.5-10 mm long, 4-5 mm wide, acute. Petals ligulate,
6-7 mm long, about 2 mm wide, acute. Lip rigidly at-
tached to the column foot, reflexed at about the middle
when viewed laterally, 5-6 mm long when straightened,
the basal half canaliculate, the apical half 3-lobate; lat-
eral lobes reduced to auricles; midlobe very bluntly V-
shaped, provided with a similarly bluntly V-shaped cal-
lus at the midlobe base. Column stout, about 2 mm long
excluding the anther; anther with a beak; pollinia 4, sup-
ported on a distinct stipe about 1 mm long and a saddle-
shaped viscidium; rostellum prominent.
Epiphytic in premontane rain forests at 500-
1500 m. Flowering material has been collected
September to November and July. Costa Rica
from the region of San Isidro del General, Prov-
ince of San Jose to Boquete, Panama.
Maxillaria horichii is unique in the more or less
urseolate red flowers resembling those of species
of Mediocalcar from New Guinea. Although sim-
ilar to M. pseudoneglecta (see below), the lip (es-
pecially when spread) and column are quite dif-
ferent. The lip of M. horichii is narrowly cuneate
at the base with a relatively long, broadly lanceo-
late midlobe, and the lateral lobes are reduced to
auricles. When viewed laterally, the lip appears
boomerang shaped. The callus is bluntly V-shaped
in front and not transversely straight and contin-
uous with the lateral lobes as in M. pseudone-
glecta. Although the orange-colored forms seen in
M. pseudoneglecta suggest hybridization with M.
horichii, available herbarium specimens reveal no
flowers with intermediate morphology.
60
FIELDIANA: BOTANY
Maxillaria inaudita Rchb. f., Beitr. Orch. Centr.-
Am. 76. 1866. TYPE: Costa Rica, Cartago,
Wendland s.n. (holotype: w). Figure 2C.
Plant epiphytic, cespitose, as juveniles consisting of
tightly clustered shoots with pseudobulbs; at maturity
producing a cluster of elongate, monopodial, decumbent,
leafy canes to 1 m tall. Roots large, to 2 mm in diameter.
Pseudobulbs of juveniles ovoid, apically 1 -foliate, to 5
cm long. Leaves coriaceous, distichous on the canes, 3-
7 cm apart, shortly petiolate above the articulation, the
sheaths rugose, the petiole narrowing just above the ar-
ticulation, the blade 5-20 cm long, 2-5 cm wide, ob-
long-elliptic, often striate, the apex variable, usually
rounded to obtuse, sometimes emarginate. Inflores-
cences lateral on the canes, single per leaf axil, short,
rarely reaching the subtending blade apex; scapes 5-8
cm long, mostly concealed by acute sheaths and the sub-
tending leaf sheath; ovary with pedicel 14-18 mm long,
usually exceeded by the subtending inflated floral bract.
Flowers white to light yellow, the lip darker yellow or
spotted with dark yellow near the apical margin. Sepals
elliptic to lanceolate, 3-6.5 cm long, 7-12 mm wide,
acute. Petals narrowly elliptic, falcate, 2.5-6 cm long,
5-10 mm wide. Lip cuneate, apically 3-lobate, 1.5-2 cm
long, 1.2-1.7 cm wide when spread; callus 3-carinate to
just above the middle. Column arcuate, 1.2-1.5 cm long
including the short foot; pollinia 4, supported on short,
squarrose stipe with viscidium. Capsule narrowly ob-
ovoid, 5-6 cm long.
Epiphytic in cloud forests at 800-1800 m.
Flowering throughout the year in Costa Rica; Jan-
uary to June in Panama. Costa Rica and Panama.
Sometimes confused with the much smaller-
flowered M. trilobata Ames & C. Schweinf., M.
inaudita is easy to recognize by the large light
yellow to white flowers. Maxillaria inaudita var.
minor Ames & C. Schweinf. is described as hav-
ing flowers about half the size of those of typical
M. inaudita, and most of the available samples
from Panama show this smaller tendency. Most
herbarium specimens include only upper stem
portions lacking pseudobulbs, which probably ex-
plains why Reichenbach's description indicates
that the species lack pseudobulbs. Approximate
shoots with apically 1 -foliate, ovoid pseudobulbs
do develop in juveniles, and the mature canes de-
velop from these. Juveniles appear very different
from the adults.
Maxillaria lankesteri Ames, Sched. Orch. 7:11-
12. 1924. TYPE: Costa Rica, Lankester 508
(holotype: AMES). Ornithidium aurantiacum
Schltr, Repert. Spec. Nov. Regni Veg. Beih. 19:
241. 1923. Costa Rica, San Pedro de San Ra-
mon, Brenes 129 (drawing of type: AMES). Not
Maxillaria aurantiaca A. Rich. & Gal. Figure
13B.
Plant a somewhat straggly epiphyte to 20 cm tall;
rhizome erect or decumbent, rooted mostly at the base;
rhizome segments to about 8 cm long between pseudo-
bulbs. Roots to 0.5 mm in diameter. Pseudobulbs cylin-
dric to somewhat tapered, 1-3 cm long, about 7 mm
wide at the base; subtended by 1-2-foliate sheaths; apex
1 -foliate. Leaves coriaceous to fleshy, sessile, elliptic,
1-3.5 cm long, 7-13 mm wide, with unequally 2-lobate
apex. Inflorescence 1 per rhizome bract axil of the de-
veloping shoot; scape to about 1 .5 cm long; ovary with
pedicel 7 mm long, nearly concealed by the subtending
floral bract. Flowers translucent tan with maroon stripes.
Sepals similar, elliptic to oblong-lanceolate, 1.2-1.4 cm
long, 3-3.5 mm wide, acute; laterals inserted on the col-
umn foot. Petals elliptic-lanceolate, about 1 1 mm long,
3 mm wide, acute. Lip hinged to the column foot, ellip-
tic-ovate in general outline, 9-9.5 mm long, 3.5 mm
wide in natural position, 5.5 mm wide when spread, 3-
lobate at the base; lateral lobes short, not exceeding !/5
the lip; midlobe elliptic, obtuse, the side margins much
recurved; callus tonguelike, exceeding and connecting
the lateral lobes. Column stout, 2-3 mm long excluding
the anther; foot about 2 mm long; anther about 1.5 mm
in diameter.
Maxillaria lankesteri is a poorly collected epi-
phyte in premontane rain forests at 600-1500
(1900?) m. Flowering July to February. Costa
Rica and western Panama.
It appears to be uncommon, because few du-
plicate specimens are available. It has been con-
sidered as a synonym of M. \vercklei (Schltr.) L.
O. Williams, a species with somewhat smaller
shoots and flowers nearly identical in morphology
but much smaller in size.
Maxillaria linear! folia Ames & C. Schweinf.,
Sched. Orch. 10:95-96. 1930. TYPE: Costa
Rica, San Ramon, A. M. Brenes (19) 314 (ho-
lotype: AMES). Figure 3A.
Plant a straggly, somewhat vinelike, essentially mon-
opodial, pendent or decumbent epiphyte forming wiry,
sparsely branched canes to about 4 mm in diameter.
Roots white to grayish, to 0.5 mm in diameter. Pseudo-
bulbs lacking, stems covered by leaf sheaths. Leaf
blades narrowly lanceolate, coriaceous, shiny, 2-10 cm
long, 3-8 mm wide, acute. Inflorescence single per leaf
axil; scape about 1.5-2 cm long; ovary with pedicel 3-
3.5 cm long, far exceeding the subtending floral bract.
Flowers white or cream marked with purple spots, the
lip purple or spotted with purple. Sepals narrowly ovate,
13-14 mm long, 4 mm wide, acute; dorsal somewhat
concave. Petals lanceolate, falcate, 12 mm long, 3 mm
wide. Lip hinged to the column foot, cuneate, 11-12
mm long, 3-4 mm wide, shallowly 3-lobate on the upper
W, midlobe somewhat broadened above, broadly round-
ed at the apex. Column arcuate, much broadened above,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
61
8 mm long, extended into a foot 3 mm long at the base.
Capsule ellipsoid, 2.5-3 cm long.
An epiphyte of larger branches, grasslike,
sometimes forming large, hanging masses at
1200-1700 m. Flowering September to February
and July. Costa Rica and Panama.
In Costa Rica, M. linearifolia is most similar to
M. appendiculoides C. Schweinf. but differs in the
distinctly lanceolate, not elliptic, leaves. It is sim-
ilar to the South American M. graminifolia
(Kunth) Rchb. f. and may prove to be conspecific.
Maxillaria longiloba (Ames & C. Schweinf.) J.
T. Atwood, Ic. PI. Trop. 16: t. 1562. 1993. Bas-
ionym: Maxillaria brenesii Schltr. var. longilo-
ba Ames & C. Schweinf., Sched. Orch. 10:90-
91. 1930. TYPE: Costa Rica, Province of He-
redia, Yerba Buena, northeast of San Isidro.
Standley & Valeria 49104 (holotype: AMES; iso-
type: us). Figure 19D.
Plant a cespitose epiphyte, sometimes forming large
clumps. Roots about 1 mm in diameter. Pseudobulbs
not much flattened laterally, conic-ovoid, 2.5-4 cm long,
1.5-2 cm wide, hidden by short, subulate nonfoliaceous
sheaths fraying with age, apically 1 -foliate. Leaf petio-
late, the blade coriaceous, keeled beneath, narrowly el-
liptic, 17-27 cm long, 2.5-4.5 cm wide, acute. Inflo-
rescences 2 or more per bract axil at the base of the
pseudobulb, produced successively; scape 8-25 cm long,
mostly concealed by 4-7 acute sheaths; ovary and ped-
icel 2.5-4 cm long, subtended by a usually somewhat
shorter, acute, laterally flattened floral bract. Flowers
fragrant, not spreading widely, greenish yellow, the lip
usually black-purple. Sepals similar, somewhat concave,
oblong-lanceolate, acute; dorsal 2.2-2.5 cm long, about
7 mm wide; lateral sepals adnate to the column foot,
2.4-2.7 cm long, about 8 mm wide. Petals linear-lan-
ceolate, 2-2.2 cm long, 4 mm wide, acute. Lip articulate
to the column foot, parallel with the column, about 1.5
cm long, 1.5 cm wide when spread, 3-lobate; side lobes
more or less truncate apically with involute margins
forming a canaliculate lip base; midlobe strongly re-
flexed to about 90°, exceedingly fleshy and terete, warty,
the margins somewhat revolute; callus a fleshy plate,
apparently pubescent (difficult to see in some herbarium
specimens). Column stout, arcuate, 7-10 mm long in-
cluding the anther; column foot 2-3 mm long; pollinia
4, supported on a saddle-shaped viscidium. Capsule
about 6 cm long.
Epiphytic in cloud forests at 1100-2000 m;
probably restricted to the dryer areas of rain shad-
ows. Flowering October to March. Costa Rica and
Panama.
Maxillaria longiloba is readily distinguished
from M. brunnea Lind. & Rchb. f. by flower color
and the sharply reflexed midlobe of the lip. No
hybrids have been seen in Monteverde, where
both species may be seen growing together.
Maxillaria longipetiolata Ames & C. Schweinf.,
Sched. Orch. 8:61-62. 1925. TYPE: Panama,
hill east of Corozal, Powell 307 (holotype:
AMES). Figure 19 A.
Plant a cespitose epiphyte 15-35 cm tall. Roots gray-
ish, to 1 .5 mm in diameter. Pseudobulbs ovoid, to 3 cm
long, 1 -foliate. Leaves with distinct petiole more than 1A
the blade; blade narrowly elliptic to linear, chartaceous
to subcoriaceous, 15-27 cm long, 1-2.5 cm wide, acute.
Inflorescence shorter than the pseudobulbs, in speci-
mens seen about 1 cm long; ovary with pedicel 1.5 cm
long. Flowers somewhat translucent, dull red, lip darker
red and column orange with white tip (fide Ingram 832,
SEL). Sepals ovate-lanceolate, 1.8-2 cm long, 5.5-7.5
mm wide, acute; lateral sepals somewhat oblique. Petals
linear to narrowly elliptic, acute, 1.5-1.8 cm long, 3-5.5
mm wide. Lip hinged to the column foot, 1.4-1.6 cm
long, about 8 mm wide when spread, distinctly 3-lobate
at about the middle; lateral lobes rounded and forming
an acute but shallow sinus with the midlobe; midlobe
oblong, apically rounded to bluntly acute; callus ligulate,
somewhat raised apically and terminating at the middle,
apex rounded to somewhat V-shaped. Column stout, 7-
8 mm long excluding the anther (1.1 cm long including
the foot); foot 4-5 mm long; anther about 2 mm in di-
ameter; stipe short and viscidium concave.
Apparently a rare canopy epiphyte in lowland
rain forests at 0-550 m. Flowering at least Janu-
ary, June, October, and December. Costa Rica and
Panama.
Maxillaria longipetiolata is easily distinguished
by the long petiolate leaf and the much abbrevi-
ated scape with flowers produced at the base of
the pseudobulbs. It is clearly related to South
American M. auyantepuiensis Foldats.
Maxillaria lueri Dodson, Icon. PI. Trop. 2: t. 155.
1980. TYPE: Ecuador, Pichincha, 300 m, Dod-
son & Luer 6800 (holotype: SEL). Figure 2A.
Plant a pendent, somewhat straggly epiphyte to 1 m
long (Ecuador); stem concealed by black-warty imbri-
cate sheaths. Roots to 1 mm in diameter. Pseudobulbs
terete, surrounded by leaf sheaths, apically 1 -foliate and
surrounded basally by 3-4 well-developed leaves.
Leaves narrowly linear, the larger at least 30 times as
long as wide, articulate to the sheath, linear, to 40 cm
long, less than 5 mm wide. Inflorescences borne on
short peduncles concealed by the leaf bases; ovary
length about 1.5-2 cm exceeded by the floral bract.
Flowers poorly known in fresh condition, pinkish white.
Sepals dissimilar; dorsal elliptic, 1.8 cm long, 7 mm
wide, acute; lateral sepals obliquely ovate, to 2.5 cm
long, 1 cm wide, adnate to the column foot. Petals
obliquely elliptic, to 1.5 cm long, 6 mm wide. Lip about
62
FIELDIANA: BOTANY
2 cm long, apically 3-lobate, the midlobe about 1A the
length of the lip, its blade with a central ligulate callus.
Column 7-8 mm long with a conspicuous foot of about
the same length.
Uncommon epiphyte of rain forests at 50-500
m. Flowering October and December. Ecuador,
Panama, and Costa Rica.
Maxillaria lueri is easily recognized by the
long, narrow, pendent leaves and spotted sheaths.
Maxillaria maleolens Schltr. Repert. Spec. Nov.
Regni Veg. Beih. 19:233. 1933. TYPE: Costa
Rica, Alajuela, Brenes 281 (lectotype: CR). Fig-
ure 17C.
Plant a large, coarse, cespitose epiphyte. Roots,
white, large, to 3.5 mm in diameter. Pseudobulbs ovate,
smooth, compressed, 5-9 cm long, 2.5-3.5 cm wide,
apically 1 -foliate, and subtended by 2 or more large leaf-
bearing sheaths. Leaves subcoriaceous and rather tough,
16-42 cm long, 2.2-5.5 cm wide, oblong, variously pet-
iolate, apically obtuse to unequally 2-lobate. Inflores-
cence a fascicle of successively flowering scapes; each
scape 4-5 cm long, covered by somewhat flared, obtuse,
apiculate bracts; ovary with pedicel furfuraceous, 1.2-
1.8 cm long, much exceeding the subtending, much ab-
breviated, collar-like floral bract. Flowers leathery, pre-
sented perpendicularly to the line of growth, with fra-
grance similar to that of coconut, greenish yellow, with
paler lip marked with red and with yellow calli. Sepals
narrowly ovate, 2.3-2.6 cm long, 1-1.1 cm wide, cus-
pidate. Petals elliptic-oblong to lance-ovate, broadly
acute, 1.8-2.1 cm long, 5-6 mm wide. Lip hinged to
the column foot, rounded at the base, 1.8-2.2 cm long,
1.4 cm wide, 3-lobate just beyond the middle; midlobe
triangular; callus thickened, biblike, extended to the
middle, on which is superposed an additional linear
raised callus, plus an additional ovate callus on the mid-
lobe. Column arcuate, 1.4-1.5 cm long including the
minutely papillose anther cap and the obsolete foot; pol-
linia 4, supported on a squarrose stipe and a saddle-
shaped viscidium. Capsule ellipsoid, 3.5 cm long.
Apparently a rare, large, clumping epiphyte of
premontane rain forests to 1150 m. Flowering
throughout the year. Mexico (Veracruz) and Hon-
duras to Panama; probably also in Guatemala.
Maxillaria maleolens is easily distinguished by
the large plants with shoots producing several
leaves at the pseudobulb base and yellow and red
flowers with obtuse floral segments. The odor, re-
sembling something between naphthalene and co-
conut, could be interpreted as "pleasant" despite
the specific epithet meaning "bad odor." Similar
species, M. discolor (Lodd.) Rchb. f. and M. vil-
losa (Barb. Rodr.) Cogn., may also occur in Costa
Rica. Maxillaria discolor, known from adjacent
Panama, has dark leaves with a purple abaxial
surface and yellow flowers that elsewhere in the
range (Surinam) have an odor similar to that of
very sharp cheese. A living plant apparently of
M. villosa presumably collected in Costa Rica has
appeared at Lankester Gardens. The species lacks
the purple pigment in the leaves, but the flowers
are yellow and lack perceptible fragrance. These
species are excluded from the flora until they can
be verified with field collections.
Maxillaria meridensis Lindl., Orch. Linden 19:
No. 100. 1846. TYPE: Venezuela, near Merida,
6000 ft, Linden s.n. (holotype: K). Figure 6D.
Plant erect, forming canes; shoots 7-11 cm apart and
terminated just above the swelling or indistinct pseudo-
bulb, each shoot developing from the base of the swell-
ing of the previous shoot, a feature obscured by closely
imbricated, rugose-verrucose rhizome bracts. Roots to 1
mm in diameter. Leaves 4-6 per shoot, the lowermost
abscising with age, the blades 5-28 cm from the artic-
ulation, and 3-9 mm wide. Inflorescence 1-2 per rhi-
zome bract; scapes 4-5 cm long, covered by about 5
imbricate, acute sheaths to 2.3 cm long; ovary with ped-
icel shorter than the subtending bract, 1.5-1.8 cm long
at anthesis. Flowers light yellow, sometimes tinged with
pink; lip with two reddish spots. Sepals narrowly ellip-
tic-lanceolate, 2-2.5 cm long, 5-7 mm wide; dorsal
somewhat concave; lateral sepals forming a mentum
around the column foot. Petals linear-lanceolate, arcu-
ate, 1.8-2.4 cm long, 2-3 mm wide. Lip 1.2-1.4 cm
long and about 7 mm wide, 3-lobate on the apical VS;
midlobe fleshy, triangular, verrucose, about 5 mm long;
callus ligulate, reaching to just above the middle, thick-
ened apically and rounded or somewhat parabolic in
front. Column arcuate, 7-8 mm long excluding the an-
ther; anther about 2 mm in diameter. Capsule 3.5 cm
long.
Terrestrial on open embankments, often on
windy and cloud-swept ridges; in Costa Rica
known from Cordillera de Talamanca at about
1400-1800 m. Flowering March, May, August,
and September. Northern South America south to
Bolivia(?) and Costa Rica.
Maxillaria meridensis is similar to M. exaltata
(Kraenzl.) C. Schweinf., a species of mostly lower
elevations, although they may occur sympatrically
(e.g., Tapantf). That species forms a much larger
plant, and the ratio of the midlobe length to the
entire lip length is <J/4. The corresponding ratio
in M. meridensis is ^¥3.
Maxillaria microphyton Schltr., Repert. Spec.
Nov. Regni Veg. 8:457. 1910. TYPE: Costa
Rica, in forests of La Palma, Tonduz 9670 (iso-
type: us). Ornithidium parvulum Schltr., Repert.
Spec. Nov. Regni Veg. 9:206. 1911 (based on
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
63
same collection as M. microphytori). Ornithi-
dium pallidiflavum Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:143, 242-243. TYPE: Cos-
ta Rica, San Pedro de San Ramon, 1075 m, Bre-
nes 135 (isotype: AMES). Maxillaria pallidiflava
(Schltr.) Senghas, Orchidaceen 29:1751. 1994.
Figure 12B.
Plant an erect or pendent, cane-forming epiphyte to
about 30 cm tall, rooted only on the lower shoots; stems
much elongate, to about 7 cm long between pseudobulbs
in the lower shoots, diminishing in successive shoots.
Roots to about 1 mm in diameter. Pseudobulbs conic to
suborbicular, never regularly furrowed in dried speci-
mens, 0.7-2.5 cm long, subtended by 1-2-foliaceous
sheaths, the apex 1 -foliate. Leaves similar, those sub-
tending the pseudobulb with sheath; apical leaf shortly
petiolate; blades thin, elliptic, 1.5-6 cm long, 4-10 mm
wide, rounded to emarginate. Inflorescence 1 per blade-
bearing or non-blade-bearing sheath of both mature and
immature shoots; ovary with pedicel 3-4 mm long,
somewhat exceeded by the subtending acute floral bract.
Flowers not opening widely or with sepals and petals
recurved, sometimes minutely pubescent, variously col-
ored, greenish yellow, often suffused or lined with dark
red, apparently turning dark with age. Sepals subequal,
elliptic, the laterals subfalcate, 6-12 mm long, 1.5-2.5
mm wide. Petals elliptic-lanceolate, sometimes attenu-
ate, 6-10 mm long, 2-3 mm wide, acute. Lip straight
in natural position, somewhat hinged to the column foot,
ovate when spread, 3.5-4.5 mm long, to about 2 mm
wide, 3-lobate below the middle; lateral lobes embracing
the column and exceeding the callus, bluntly acute; mid-
lobe ovate, obtuse to rounded; callus at the base fleshy,
rounded and somewhat elevated in front. Column stout,
arcuate, somewhat tapering, 2-3 mm long excluding the
anther; foot about 0.5 mm long; anther less than 1 mm
in diameter; pollinia 4, with somewhat squarrose stipe.
Locally abundant epiphyte or terrestrial of
cloud forests at 1000-2000 m. Often found grow-
ing on shrubs. Flowering mostly June to Novem-
ber, also March and April. Costa Rica; expected
in Panama.
The straight lip with lateral lobes exceeding the
callus distinguishes this from similar species, es-
pecially M. flava and M. wercklei.
Although both M. microphyton and Ornithi-
dium parvulum are based on the same collection,
conceivably the names are based on separate
specimens that might not have been the same spe-
cies. In the absence of holotypes, there is no
choice but to regard the second published name
as a later synonym of the first.
Maxillaria minor (Schltr.) L. O. Williams, Amer.
Orch. Soc. Bull. 10:273. 1942. Camaridium mi-
nus Schltr., Beih. Bot. Centralbl. 36, Abt. 2:417.
1918. TYPE: Costa Rica, La Palma, C. Werckle
(holotype: CR?; drawing of type: AMES). Figure
13C.
Plant a straggly, pendent epiphyte to about 1 m long
comprised of several successively borne shoots to 15 cm
long, each consisting of a slender, wiry, scaly rhizome
(cane) to about 3 mm in diameter terminated by a pseu-
dobulb. Roots white, to barely 0.5 mm in diameter.
Pseudobulbs ovoid to orbicular, biconvex, 1-2 cm long,
to 1 cm wide, subtended when young by 1-2 leaf-bear-
ing sheaths; apex 1 -foliate. Leaves 4-8 cm long, 7-12
mm wide; blades of rhizome bracts with a broad sheath,
narrowing abruptly above the articulation to a slender
petiole. Inflorescences 1 per rhizome bract, produced
precociously on the canes of developing shoots; scapes
to about 5 mm long; ovary with pedicel 3-4 mm long,
subtended by a cucullate, acute, or subulate floral bract.
Flowers variable in color, mostly white to salmon pink,
the lip yellow. Sepals similar; dorsal oblanceolate, 7 mm
long, 3 mm wide, acute; lateral sepals obliquely ovate,
5 mm long, 2 mm wide. Petals oblanceolate-obovate, 5
mm long, to 2 mm wide. Lip hinged to the column foot,
3-lobate just above the middle, 3 mm long, about 2 mm
wide, the midlobe fleshy, papillose, sharply deflexed,
with a fleshy callus at the base and a second U-shaped
callus at the base of the midlobe. Column stout, arcuate,
1.5 mm long, with a very short foot about 0.5 mm long.
Capsule globose, about 6 mm long.
Epiphytic on the larger branches and trunks of
trees in wet forests at 1000-2500 m. Flowering
July to March. Costa Rica and Panama.
The small flowers formed precociously on im-
mature canes and the straggly habit are immediate
field characters for recognition. Maxillaria minor
vegetatively resembles M. paleata (Rchb. f.)
Ames & Correll, M. schlechteriana J. T. Atwood,
M. sigmoidea (C. Schweinf.) Ames & Correll, and
M. vaginalis Rchb. f., but of these has the smallest
and most dully colored flowers. The entire com-
plex of species is found at middle to high eleva-
tions, often growing sympatrically.
Maxillaria monteverdensis J. T. Atwood & G.
Barboza, Lindleyana 9:241-242. TYPE: Costa
Rica, intersection of Provinces of Alajuela,
Guanacaste, and Puntarenas, Monteverde Re-
serve, Atwood 89-33 (holotype: SEL; isotype:
CR). Figure 7A.
Plant a large, erect, monopodial, cane-forming epi-
phyte in adults to at least 35 cm tall (probably much
taller) with leafy canes lacking pseudobulbs; juveniles
cespitose and with pseudobulbs. Roots mostly restricted
to the base, to about 2 mm in diameter. Pseudobulbs of
juveniles about 4.5 cm tall and 2.5 cm wide, com-
pressed, sessile, apically 1 -foliate with shortly petiolate
leaf. Leaves thin, articulate; blade keeled below, 4-1 1
cm long, 1.5-2 cm wide; apex obtuse to rounded and
unequally 2-lobate. Inflorescence several per leaf axil.
64
FIELDIANA: BOTANY
long for the genus and sometimes exceeding the sub-
tending leaf; scapes to about 8 cm long, persistent; ovary
with pedicel 1.3-1.5 mm long, subtended by a somewhat
longer subulate, acute floral bract. Flower large for the
genus, campanulate, to about 3.2 cm long, white to pink-
ish with lip yellow to orange, fragrant. Sepals similar,
oblong, 2.5-3.1 cm long, about 5-7 mm wide, apex
acute. Petals falcate-oblanceolate, about 2.5-2.7 cm
long, 6.5-9 mm wide, acute to acuminate. Lip somewhat
rigid with the column foot, canaliculate on the lower %,
3-lobate on the upper Vs, about 1.5-1.8 cm long, 3.5-4
mm wide in natural position, to 2 cm long when spread;
lateral lobes embracing the column; midlobe ovate, ob-
tuse, and with crenulate margin; callus a truncate to
rounded lamina terminating at about the lobe sinuses
with a single papilla at the base. Column nearly straight,
cylindric, 1.2-1.5 cm long excluding foot and anther;
foot short, about 3 mm long.
Epiphytic in lower montane rain forests at
1500-1800 m. Flowering at least January to
March. Costa Rica (Monteverde) and Panama.
Maxillaria monteverdensis is most closely re-
lated to M. amabilis but has longer ovaries and
larger flowers, and the lip has a single papilla at
the base instead of a distinct keel. The flowers
resemble those of M. vaginalis Rchb. f., but the
latter species distinctly has pseudobulbs terminat-
ing adult shoots. When described, M. montever-
densis was thought to be endemic to Monteverde,
but specimens have now surfaced from Panama.
Maxillaria moralesii Carnevali & J. T. Atwood,
Lindleyana 11:31-32. 1996. TYPE: Costa Rica,
Alajuela, Rio Angel, flowered in cultivation at
Missouri Botanical Garden 25 Dec. 1994, Car-
nevali & Morales 3841 (holotype: INB; isotype:
SEL). Figure 22A.
Plant a cespitose or shortly creeping epiphytic herb
8-15 cm tall. Rhizome short; roots to 2 mm in diameter.
Pseudobulbs pale yellow-green, clustered, ellipsoid or
oblongoid, 2-3 cm tall, 0.8-1.5 cm wide and thick, but
always wider than thick; subtended by non-foliaceous
sheaths; apex 1 -foliate. Leaves without petiole, erect on
the pseudobulb, coriaceous, elliptic or lanceolate-ellip-
tic, 6.5-25 cm long, 1.3-1.7 cm wide, acute. Inflores-
cence erect or spreading, shorter than pseudobulb, pe-
duncle 8-12 mm long; ovary with pedicel 1.6-1.7 cm
long, subtended by an obovate-elliptic floral bract 1.1-
1.2 cm long, 6-7 mm wide. Flowers dull yellow-green,
lip yellow with orange-maroon speckles, column pale
green. Sepals somewhat fleshy, acute; dorsal ovate-el-
liptic, concave, 12.5-13.5 mm long, 6-7 mm wide; lat-
eral sepals ovate, 12.5-13.5 mm long, acute, mucronu-
late. Petals lanceolate, 12.5-13.5 mm long, 3.8-4.2 mm
wide, acute, mucronulate. Lip 3-lobate, 10.5-12 mm
long, ca. 7 mm wide when expanded; lateral lobes tri-
angular-ovate, obtuse, erect-uncinate, emerging from the
lower J/3 of the lip; 1.2-1.5 mm long; midlobe ovate -
oblong-subquadrate, evenly tapering from base to apex,
5.8-6.2 mm long, 2-2.5 mm wide, apex subtruncate-
rounded; disc bearing a narrowly triangular farinose cal-
lus narrowing toward the midportion then thickening
again. Column arcuate, 7.5-8 mm long, 3-4 mm wide;
broadly winged near apex; column foot 2-2.5 mm long;
anther smooth, 2.3-2.5 mm long. Capsule about 2.5 cm
long.
Poorly known ecology, tropical lowland rain
forests (e.g., La Selva Biological Station) to 900 m.
Flowering November to February. Central Valley of
Costa Rica; probably also Nicaragua and Honduras.
Four distinct members of the M. rufescens com-
plex are treated separately here. Maxillaria acu-
tifolia Lindl. is a small member with nearly cylin-
dric pseudobulbs and a yellow to green flower
with the lip containing a red stripe up the center.
Maxillaria hedwigiae Hamer & Dodson is a spe-
cies with a white to green flower and an orange
lip. Finally, M. dressleriana is a species with
ovoid pseudobulbs and brownish maroon flowers.
Maxillaria moralesii has been accepted variously
as M. rufescens Lindl., a larger-flowered species
from South America. It is possible that plants of
M. moralesii and M. acutifolia as accepted here
are indistinguishable.
Maxillaria muscoides J. T. Atwood, Lindleyana
9:236. 1994. TYPE. Costa Rica, Cerros el
Humo de Pejivalle in the zone of Turrialba,
Dodson 2526 (holotype: SEL). Figure 12A.
Plant small, somewhat moss- or brushlike, to about
10 cm tall, consisting of a tuft of small, few-branched
canes appearing somewhat cespitose; rhizome elongate,
to 2 cm between pseudobulbs, rooted only at the base,
concealed by scale bracts. Roots less than 0.5 mm in
diameter. Pseudobulbs conic, somewhat grooved, to
about 1.2 cm long and 5 mm wide, subtended by 3-4
foliaceous bracts; apex 1 -foliate. Leaves acicular, terete,
articulate, 1-3.5 cm long, about 1.5 mm in diameter, the
apex obtuse and apiculate. Inflorescence 1 per folia-
ceous or nonfoliaceous rhizome bract appearing on the
nearly mature shoot; scapes to about 1 cm long; ovary
with pedicel 2-2.5 mm long, exceeded by the subulate
floral bract. Flowers pale yellow, apparently not opening
broadly. Sepals dissimilar; dorsal ligulate, somewhat
concave, 5-5.5 mm long, 1.2-1.8 mm wide, acute, dor-
sally keeled at the apex; lateral sepals triangular-falcate,
attached to the column foot, 5 mm long, less than 2 mm
wide, acute. Petals elliptic-lanceolate, slightly falcate,
4-4.5 mm long, about 1.2 mm wide, acute. Lip hinged
to the column foot, recurved in natural position, when
spread (difficult) 3-3.5 mm long, 2-2.2 mm wide, shal-
lowly 3-lobate at about the middle; lateral lobes rounded
and somewhat embracing the column; midlobe ovate,
acute, dorsally keeled; callus lamellate connecting the
lateral lobes, ending at or below the sinuses. Column
short, less than 2 mm long excluding the anther; foot
about 1 .8 mm long; anther 1 .5 mm in diameter; pollinia
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
65
4, supported on a short, squarrose stipe with dark vis-
cidium (dried material).
Epiphytic in premontane rain forests at 1350 m.
Flowering August to November. Endemic to Cos-
ta Rica; known only from Tapanti.
In habit M. muscoides is unlike any other spe-
cies known from Central America, and the needle-
like leaves are reminiscent of certain Brazilian
taxa; however, these other taxa have two leaves
at the pseudobulb apex. Flower morphology in-
dicates a distinct relationship with Costa Rican M.
microphyton Schltr. Its small size renders it unable
to compete with more showy orchids for the eye
of all but the most ardent collectors.
Maxillaria nasuta Rchb. f., Beitr. Orch. Centr.-
Am. 104. 1866. TYPE: Colombia, Lasita (ho-
lotype: w; isotype: K). Maxillaria nasalis Rchb.
f., Saund. Ref. Bot. 2: t. 102. 1872. TYPE: Cos-
ta Rica, Endres 266 (holotype: w). Maxillaria
brevipedunculata Ames & C. Schweinf. Sched.
Orch. 10:91. 1930. TYPE: Costa Rica, Toro
Amarillo, A. Alfaro 190 (holotype: us; photo of
type: AMES). Figure 17D.
Plant a robust, cespitose epiphyte of larger branches
and trunks in evergreen forests. Roots large, to 5 mm in
diameter. Pseudobulbs smooth, appearing varnished, not
forming ridges when dry, ovate, compressed, 5-7 cm
long, 2.5-3.5 cm wide, subtended by several leaf-bear-
ing rhizome bracts, apically 1 -foliate. Leaves linear,
with a long conduplicate petiole, 20-65 cm long from
the articulation, 2-3.5 cm wide, acute to obtuse. Inflo-
rescence 2 or more per rhizome bract, successively
borne, covered by several blunt flaring sheaths; ovary
about 3 cm long. Flowers with leathery segments, green
to yellow, the lip mostly red. Sepals lanceolate to ellip-
tic, 3.5-4.5 cm long, 1-1.2 cm wide, acute. Petals el-
liptic, oblique at the base, 2-3 cm long, 5-7 mm wide,
acuminate. Lip basally rounded, 2 cm long, 1.5 cm
wide, 3-lobate from just above the middle; midlobe tri-
angular, reflexed, callus on the lower half oblong but
poorly defined, papillose, and cushion-like. Column ar-
cuate, 1—1.4 cm long, the anther cap with a dark spot
on each side. Capsule ellipsoid, 5 cm long.
A large, clumping epiphyte of larger branches
in wet forests at 150-1500 m in Central America;
to 2000 m in Ecuador. Flowering April to October
in Central America. Mexico to Bolivia.
Specimens from higher elevations in Costa
Rica and Ecuador often exhibit larger-scape bracts
than do those from lower elevations, but the vari-
ation is probably continuous.
Maxillaria neglecta (Schltr.) L. O. Williams,
Ann. Mo. Bot. Card. 29:348, Fig. 4. 1942. Or-
nithidium neglectum Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:242. 1923. TYPE: Costa
Rica, A. M. Brenes 164 ex R. Schlechter
(pressed from cultivated material) (drawing of
type: AMES). Not Scaphy glottis pendula Poeppig
& Endl. sensu Brieger. Figure 9C.
Plant a straggly, usually somewhat pendent epiphyte
often forming large clumps to about 50 cm across; rhi-
zome elongate, concealed by bracts. Roots slender to 0.5
mm in diameter. Pseudobulbs cylindric, somewhat com-
pressed, 1.8-3.5 cm long, 4-12 mm wide, spaced 1-5
cm apart, 1 -foliate. Leaf chartaceous, linear to lanceo-
late, 7-20 cm long, 5-13 mm wide. Inflorescence a fas-
cicle of scapes borne at the base of the most recently
developed pseudobulb, the scapes much less than 1 cm
long and concealed by the rhizome bracts; ovary 3-4
mm long. Flowers fleshy, white, the lip with yellow
midlobe. Sepals concave and not opening widely, dorsal
5-6 mm long, 4-5 mm wide when flattened; lateral se-
pals about l/2 connate, each obliquely ovate, 5-7 mm
long, 4.5-5.5 mm wide. Petals narrowly oblong-ovate,
slightly constricted on the upper half 4-5.5 mm long,
1-2 mm wide, the apex rounded and apiculate. Lip 3-
lobate, 4.5-6 mm long, 3-4 mm across when spread
(difficult); lateral lobes broad, subquadrate, partially
connected to the column foot and with straight or con-
cave lateral margins; midlobe fleshy, ovate, and obtuse,
V4-Y3 the lip; with a transverse ridgelike callus at the base
of the midlobe connecting the lateral lobes. Column
stout, straight, 1.2-2 mm long excluding the anther; foot
at the base about 2.5 mm long; anther globose, about 1
mm in diameter. Capsule globose about 6 mm long, 5
mm wide.
Epiphytic in tropical evergreen forests; often
over water at 0-900(1600?) m. Flowering August
to November; fruiting by January. Honduras to
Panama; possibly also Colombia.
Maxillaria neglecta (sensu L. O. Williams,
Ann. Missouri Bot. Card. 29:348. 1942) has tra-
ditionally included plants segregated below as M.
pseudoneglecta (based on Ornithidium anceps
Rchb. f.). Maxillaria neglecta is a more wide-
spread, usually lowland species and has generally
much more narrowly linear leaves. It is readily
distinguished from M. pseudoneglecta (occasion-
ally white-flowered) by the narrower lateral lobes
of the lip with essentially straight outer margin.
When spread, the lip is a maximum of 4 mm
across in M. neglecta and always (4.5)5-7 mm
across in M. pseudoneglecta. Maxillaria neglecta
is distinguished from M. brevilabia by its gener-
ally smaller, linear, and acute leaves; the latter
species has elliptic, obtuse leaves. Inflorescences
of M. brevilabia develop along the entire length
of the recently matured rhizome, not just at the
base of the pseudobulb as in M. neglecta.
66
FIELDIANA: BOTANY
Maxillaria nicaraguensis (Hamer & Garay) J.
T. Atwood, Selbyana 10:60-62. 1988. Neour-
bania nicaraguensis Hamer & Garay, Icon.
PI. Trop. 13: t. 1238. 1985. TYPE: Nicaragua,
Department of Chontales, A. H. Heller 7057
(lectotype: SEL; isolectotypes: SEL, AMES). Fig-
ure 4B.
Plant a straggly decumbent epiphyte lacking pseudo-
bulbs but often forming large clumps; stems ascending,
to about 20 cm tall, unbranched above and sheathed by
the bases of 6-12 distichous leaves to 5 cm apart. Roots
white, to 1 mm in diameter. Leaves 4-7.5 cm long and
5-8 mm wide, oblanceolate-oblong, somewhat unequal-
ly 2-lobate at the obtuse apex, articulated to the some-
what inflated rugose sheath. Inflorescence appearing
singly or in pairs in the leaf axils, the peduncle plus
ovary 1-2 cm long. Flowers creamy yellow marked
with red. Sepals oblong-lanceolate, 8-9 mm long, 1.5
mm wide. Petals obliquely oblong-lanceolate, 6-7.5 mm
long, 1.5-2 mm wide. Lip rigidly attached to the base
of the column, oblong-pandurate, constricted in the mid-
dle, 5-8 mm long, 2-2.5 mm wide; basal half concave
and parallel to the column; apical l/2 reflexed and with a
crenulate margin. Column arcuate, about 4 mm long;
foot nearly absent. Capsule ellipsoid, about 1.25 cm
long.
Epiphytic in larger branches of tree crowns of
wet lowland forests at 50-900 m. Flowering No-
vember and December. Nicaragua and Costa Rica.
Although represented at La Selva Biological
Station by a single collection, it is quite common
there but probably rarely seen in flower. A short
flowering period would account for the few col-
lections having been made.
Maxillaria cf. obscura Lind. & Rchb. f., Beitr.
Orchid.-K.C.Amer. 31-32, t. 6. 1866. TYPE:
Colombia (including Panama). Figure 22B.
Plant a cespitose epiphyte to perhaps 20 cm tall.
Roots white, to about 1.25 mm in diameter. Pseudo-
bulbs compressed, ovoid to suborbicular, 1.5-4.5 cm
long, 1-2.5 cm wide; base subtended by 1-2 subulate
bracts with articulation but without definite blade; apex
usually somewhat emarginate, 1 -foliate. Leaves shortly
petiolate (petiole < !/5 blade); blade narrowly elliptic, 5-
29 cm long, 1.2-2.9 cm wide, bluntly acute to obtuse
and unequally 2-lobate. Inflorescence apparently 1 per
bract axil of the mature shoots, to about 5 cm long;
ovary with pedicel (1.6)1.8-2 cm long, trigonous, ex-
ceeded by a cucullate, acute floral bract. Flowers ma-
roon to brown, or with sepals and petals yellow stained
with maroon. Sepals similar, broadly oblanceolate to el-
liptic, acute; dorsal somewhat concave, 1 .7-2. 1 cm long,
5-6.5 mm wide; lateral somewhat oblique, 1.8-2.1 cm
long, 5-6 mm wide. Petals broadly oblanceolate, 1.5-
1.7 cm long, 4-5 mm wide, acute. Lip hinged to the
column foot, recurved, 3-lobate below the middle, 1-1.2
cm long, 6-7 mm wide when spread; lateral lobes round-
ed and often slightly indented near the apex; midlobe
ovate, fleshy, 6-7 mm long, about 4-4.5 mm wide, acute
to obtuse and thickened near the apex; callus terminating
at about the divergence of the lateral lobes, trapezoid,
thickened in front, somewhat canaliculate toward the
base. Column stout, arcuate, 5.5-7 mm long excluding
the anther; foot 2-3 mm long.
Maxillaria cf. obscura grows in upper montane
cloud forests at 1500-3000 m. Flowering Novem-
ber to May. Costa Rica and Panama.
This species, variously accepted as M. cucul-
lata Lindl. (sensu lato) or M. atrata var. brach-
yantha Rchb. f., is restricted to southern Costa
Rica and Panama. Maxillaria cf. obscura is
unique among the M. cucullata Lindl. complex in
southern Central America in the trigonous ovary,
relatively short column, and trapezoid callus of
the lip. It is also unique in the complex for its
narrower petioles and smaller flowers, which are
either dark maroon or have sepals and petals that
are yellow stained with maroon. They are never
lined with maroon dots as in M. punctostriata
Rchb. f. Maxillaria cf. obscura is perhaps most
closely related to M. lexarzana Soto & Chiang
from Mexico but has smaller flowers with a short-
er column. Also, that species flowers at the end
of the rainy season (August to October), while this
species flowers during the dry season.
The name M. obscura is based on a plant with
its origin given as "Columbia," which included
Panama at the time of publication. The relatively
long midlobe of the lip apparent in the plate ac-
companying the original description matches the
plants described here. No type specimen has been
located.
Maxillaria oreocharis Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 17:69. 1922. TYPE: Panama,
Chiriquf, Powell 256 (isotype: AMES). Figure
15C.
Plant a large, somewhat scandent epiphyte with as-
cending shoots 1-6 cm apart, with only the lowermost
forming roots. Roots white, to about 1 mm in diameter.
Pseudobulbs ovate, apically 1 -foliate, strongly com-
pressed laterally, 1.5-4 cm long, 1-2 cm wide, subtend-
ed by bladeless rhizome bracts (at maturity), the leaf
articulation oblique. Leaves linear, shortly petiolate, the
blade 13-30 (39) cm long, 5-15 mm wide, acute. Inflo-
rescences 1 per each of several concealing rhizome
bracts of the mature shoot, 1 .5-2.5 cm long, with 3 or
more acute sheaths; ovary with pedicel 2-2.5 cm long,
far exceeding the subtending floral bract. Flowers com-
plex in color, the sepals red exteriorly, pink interiorly,
the petals white to yellow (yellowing with age?); lip red,
the apical Vs yellow. Sepals narrowly ovate-lanceolate.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
67
1.5-1.8 cm long, 4-5 mm wide, acute. Petals obliquely
lanceolate, 1.2-1.6 cm long, 3-5 mm wide. Lip about
1.2 cm long, 6 mm wide when spread, hinged to the
column foot, oblong, somewhat constricted at the base
of the apical V6; apex fleshy, reflexed; callus ligulate,
reaching to just beyond the middle. Column to about 10
mm long with anther; foot 2 mm long.
Epiphytic in wet forests at 100-1100 m. Flow-
ering August to December (February). Nicaragua
to Panama.
This species is easily distinguished from close-
ly related M. alba (Hook, f.) Lindl. and M. anceps
Ames & C. Schweinf. by the narrower, acute
leaves and by the purple color in the somewhat
smaller flowers, which are borne within the rhi-
zome bract axils of mature shoots.
Maxillaria pachyacron Schltr., Repert. Spec.
Nov. Regni Veg. 9:165. 1911. TYPE: Costa
Rica, La Palma, A. Tonduz 9681 (isotype:
AMES), Ingram & Ferrell 770 (epitype selected
here: SEL). Figure 24B.
Plant epiphytic, cespitose, forming small clumps to
about 12 cm tall. Roots to 1 mm in diameter. Pseudo-
bulbs narrowly to broadly ovoid, 1-2 cm long, subtend-
ed by 1(2) leaf-bearing rhizome bracts, apically 1 -foli-
ate. Leaves distinctly petiolate, the petiole to about %
the blade length and narrowed at the articulation; blade
elliptic-ovate to linear, 5-20 cm long, 1.5-3.5 cm wide,
acute, the apex often recurved. Inflorescence often
reaching above the leaves, to 25 cm high; scapes wiry,
nearly covered by 4-5 acute to obtuse, involute sheaths
1.5-2.5 cm long; ovary with pedicel lepidote, usually
longer than the subtending bract, 2.3-3.5 cm long. Flow-
ers spidery, the segments tan apically, whitish basally
turning brown with age. Sepals lance-attenuate, 3.5-5
cm long; dorsal 3-4 mm wide; lateral sepals 4-5 mm
wide near the base. Petals lanceolate, falcate, 2-3 cm
long, 3 mm wide. Lip obovate, 1.3 cm long and 6-7
mm wide, 3-lobate on the apical !/6; midlobe very fleshy,
with a ligulate callus extended to just below % the lip,
parabolic, thickened and elevated in front. Column ar-
cuate, 1 cm long, including the 5-mm-long foot, anther
bed papillose, pollinia 4, on a saddle-shaped viscidium.
Capsule somewhat obovoid.
Common in cloud forests at 1000-1900 m.
Flowering July to March. Nicaragua to Panama.
This species was synonymized by Ames under
M. reichenheimiana Endres & Rchb. f., but that
species has a longer mentum (chin) formed
around the longer column foot besides the more
obvious broader, white-spotted, and usually emar-
ginate leaves. Maxillaria arachnitiflora Ames &
C. Schweinf. is also similar but has more acumi-
nate sepals, often spiraled near their apices, and a
much longer column foot, and its floral bract ex-
ceeds the ovary. The similar M. chionantha J. T.
Atwood has longer pseudobulbs, proportionately
broader leaves, and a longer midlobe and grows
at lower elevations.
The identification of M. pachyacron Schltr. is
based on the original description and on the only
known isotype (drawing), at Harvard. The draw-
ing shows not only the small midlobe of the lip
but also a broad leaf, suggesting that Schlechter
constructed a monstrosity from different speci-
mens, possibly representing different species. Be-
cause it could be confused with M. chionantha,
an epitype is designated for clarification in the
absence of other original material. As a further
complication, a specimen with a label consistent
with the collection data in the original description
of M. pachyacron has surfaced at us, but the flow-
er is clearly much smaller than that indicated in
the description and has a very dissimilar lip. Fur-
thermore, the plant has long rhizomes, whereas
the original description indicates "rhizomate val-
de abbreviate. " This plant is M. schlechteriana J.
T. Atwood, and the label seems to have been com-
bined with the specimen in error.
Maxillaria paleata (Rchb. f.) Ames & Correll,
Bot. Mus. Leafl. 11:15. 1943. Ornithidium pa-
leatum Rchb. f., Linnaea 41:36. 1877. TYPE:
Central America (holotype: w). Camaridium
wrightii Schltr., Repert. Spec. Nov. Regni Veg.
16:448. 1920. TYPE: Nicaragua, Wright s.n.
(isotype: AMES). Maxillaria wrightii (Schltr.)
Ames & Correll, Bot. Mus. Leafl. 11:18. 1943.
Figure 13E.
Plant a straggly, pendent epiphyte, somewhat
branched, to about 1 m long, anchored only at the base;
successively borne shoots to 25 cm long, diminishing
above, each consisting of a slender, wiry, scaly rhizome
(cane) to about 4 mm in diameter terminated by a pseu-
dobulb. Roots white, less than 1 mm in diameter. Pseu-
dobulbs ovoid to orbicular, biconvex, 1-5 cm long, to
2.5 cm wide, subtended when young by 1-4 leaf-bearing
rhizome bracts. Leaves 2-23 cm long, 0.5-4.5 cm wide;
blades of rhizome bracts with a broad sheath, narrowing
abruptly above the articulation to a slender petiole, the
apex acute. Inflorescence short, lateral, single flowered,
produced precociously on the canes of developing
shoots; scapes to about 2 cm long; ovary with pedicel
6-9 mm long, subtended by a subulate bract 1.2-1.7 cm
long. Flowers reddish purple. Sepals similar; dorsal ob-
lanceolate, 1.6-2.1 cm long, 4-4.5 mm wide, acute; lat-
eral sepals obliquely ovate, 1.6-2.1 mm long, 5 mm
wide. Petals elliptic-lanceolate to oblanceolate, 1.5-1.8
cm long, to 5-5.5 mm wide. Lip more or less fixed to
the column foot, 3-lobate just above the middle, 9-10
mm long, about 2.5 mm wide; base subsaccate; midlobe
fleshy, papillose, sharply deflexed; callus fleshy, tongue-
68
FIELDIANA: BOTANY
like at the base, ending at about the sinuses between
lateral lobes. Column stout, straight, 3.5-4.5 mm long
excluding anther; foot nearly obsolete, to about 1 mm
long.
Epiphytic and terrestrial in montane forests at
(900)1200-3000 m. Flowering July to March;
probably throughout the year. Southern Nicaragua
(Volcan Maderas) to Panama.
Maxillaria paleata is similar to M. minor
(Schltr.) L. O. Williams but has much larger, ma-
genta flowers. Most authors have accepted M.
schlechteriana J. T. Atwood (as M. wrightii van
imbricatum (Schltr.) Ames & Correll) as a variety
of what is accepted here as M. paleata. However,
no hybrids are known in areas of sympatry be-
tween that entity with smaller, lighter lavender
flowers and M. paleata.
A search for the type of Ornithidium paleatum
at the Reichenbach Herbarium has shown that the
name M. paleata has been misapplied to a similar
species, M. dichotoma (Schltr.) L. O. Williams.
The misapplication is based on a poor drawing of
the lip, which is grossly similar to that of M. di-
chotoma, but reexamination shows the character-
istic sigmoid shape consistent with M. wrightii.
Maxillaria parviflora (Poeppig & Endl.) Garay,
Bot. Mus. Leafl. 21:258. 1967. Scaphyglottis par-
viflora Poeppig & Endl., Nov. Gen. ac Sp. 1:58,
t. 1836. TYPE: Peru, Cuchero, Poeppig (holo-
type: w). Pseudomaxillaria parviflora (Poeppig &
Endl.) Brieger, Bot. Jahrb. Syst. 97:555. 1977. Or-
nithidium confertum Griseb., Fl. Br. W. hid. Isl.
626. 1864. TYPE: Trinidad; Cuba, Wright 650
(isotypes: GOET, USF). Maxillaria conferta (Gri-
seb.) C. Schweinf. ex Leon, Contrib. Ocas. Mus.
Hist. Nat. Coleg. de la Salle 8:395. 1946. Orni-
thidium chloroleucum Barb. Rodr., Gen. et Spec.
Orch. Nov. 2:208. 1882. TYPE: Brazil. Pseudo-
maxillaria chloroleuca (Barb. Rodr.) Hoehne, Ar-
quiv. Bot. Estad. S. Paulo 2:72. 1947. Maxillaria
simulans Ames & C. Schweinf. Sched. Orch. 10:
99, t. 1930. TYPE: Guatemala, Tuerckheim 8302
(holotype: AMES). Figure 9D.
Plant a straggly rhizomatous epiphyte to about 40 cm
across, rooted only at the lowermost shoots; rhizomes
elongate, concealed by bracts, to about 4 mm in diam-
eter. Roots white, rarely exceeding 0.5 mm in diameter.
Pseudobulbs cylindric to ovoid, compressed, 1.5-4 cm
long, 6-12 mm wide, spaced 1-7 cm along the rhizome,
subtended by a pair of subulate nonfoliaceous bracts,
apically 1 -foliate. Leaf subcoriaceous, shortly petiolate,
the blade lanceolate to linear, 9-20 cm long, 9-20 mm
wide, acute. Inflorescences fascicled at the base of the
most recently matured pseudobulb and usually with the
flush of new growth; scapes < 1 cm long and hidden
by the rhizome bracts; ovary and pedicel 3.5-4.5 mm
long. Flowers white to light yellow (apparently purplish
in Jamaican populations) with orange-yellow lip. Sepals
ovate, concave, carinate apically, acute; dorsal 4.5-7
mm long, 2-3 mm wide; lateral sepals Vi connate, 4.5-
6 mm long, 2-3 mm wide. Petals ovate-elliptic, 3-4 mm
long, 1.2-2.5 mm wide, acute. Lip fleshy, continuous
with the column foot; blade deltate to 3-lobate, 2.5-4
mm long, 1.5-2.5 mm across when spread; side lobes
somewhat erect; midlobe ovate and obtuse; callus at the
base emarginate and a second, larger, emarginate callus
in front continuous with the lateral lobes. Column stout,
about 1.5 mm long excluding the globose anther. Cap-
sules globose, 7-8 mm long.
An epiphyte of seasonally dry forests at 0-
1 100 m. Phenological data are lacking for Costa
Rica, but in Chiapas to Honduras flowering spec-
imens have been collected from September to No-
vember. As here interpreted, M. parviflora ranges
from Southern Chiapas, Guatemala, Belize, and
Honduras to Florida (USA), the Caribbean arc,
and probably much of tropical South America. It
seems oddly absent from mainland Nicaragua to
Panama, but specimens exist representing Isla del
Coco (Costa Rica), an island with floristic affini-
ties in South America.
This is the smallest-flowered low-elevation spe-
cies of the M. neglecta complex, which is char-
acterized by partially connate lateral sepals. It is
easily distinguished by the nearly absent mentum
and by the cuneate lip, which lacks the broad lat-
eral lobes seen in M. neglecta and M. pseudone-
glecta.
Maxillaria parviflora from South America
probably includes Ornithidium rhomboglossum
Schltr., M. exigua Regel, and M. surinamensis
Focke ex Rchb. f. as synonyms.
Maxillaria parvilabia Ames & C. Schweinf.,
Sched. Orch. 8:62-64. 1925. TYPE: Costa
Rica, Province of San Jose, La Palma, Standley
32939 (holotype: AMES; isotype: us). Figure 4D.
Plant an epiphyte often more than 1 m tall; rhizomes
forming upright or decumbent, often much-branched,
leafy canes to 1 cm in diameter lacking pseudobulbs
(apparently also as juveniles), concealed by imbricate
sheaths. Roots mostly at the base but also along the
canes, to about 1.2 mm in diameter. Pseudobulbs ap-
parently absent in juvenile shoots. Leaves distichous,
often with persistent blades restricted near the stem
apex, the sheath minutely rugose and yellowish in dried
specimens; blade deciduous, coriaceous, linear, 3.5-14
cm long, 8-15 mm wide, the base narrower than the
sheath, the apex unequally 2-lobate. Inflorescences 1-
several per leaf sheath; scapes 2-4 cm long, mostly con-
cealed by acute, involute sheaths; ovary with pedicel 6-
8 mm long exceeded by the subtending floral bract.
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
69
Flowers very fragrant, clear or light yellow, lip marked
with red. Sepals dissimilar; dorsal elliptic, 1.4-2 cm
long, 4-5 mm wide, acute; lateral sepals falcate-lanceo-
late, 1.3-1.9 cm long, 4-5 mm wide, acute. Petals el-
liptic-oblanceolate, 1.2-1.7 cm long, 4-4.5 mm wide,
acute. Lip hinged to column foot, 5-6 mm long, 3-3.5
mm wide when spread, cuneate at the base, 3-lobate
above the middle; lateral lobes rounded and clasping the
column; midlobe rounded to subtriangular; callus low,
fleshy, ligulate ending below the middle. Column
straight, 2-2.5 mm long excluding the anther; foot about
2 mm long.
Epiphytic and common in cloud forests at
1200-2300 m, particularly in wind gaps; other-
wise, little is known of its ecology. Flowering No-
vember to March. Mostly along Continental Di-
vide in Costa Rica and western Panama.
Among similar cane-forming species, this one
is unique in its small lip-to-dorsal sepal ratio (<
%) and the yellow and very fragrant flowers.
Maxillaria piestopus Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:302. 1923. TYPE: Costa
Rica, San Ramon. Tonduz s.n., May 1913 (de-
stroyed), Lankester 1213 (neotype selected
here: AMES 34211). Figure 19C.
Plant a cespitose, erect epiphyte to 1 1 cm tall. Roots
to 1.5 mm in diameter. Pseudobulbs suborbicular, about
1.2 cm tall, subtended by apparently nonfoliaceous
bracts; apex 1 -foliate. Leaves oblanceolate, erect, ob-
tuse, sessile, 8-9 cm long, 1.1-1.3 cm wide. Inflores-
cence erect, subequal to the flower, produced at the base
of immature shoots; peduncle about 5 cm long, com-
pressed, with 4-5 large, triangular, obtuse sheaths; ovary
with pedicel 2.5-2.8 cm long, triquetrous, about as long
as the floral bract. Flowers pale yellow; lip with red
spotting. Sepals dissimilar, dorsally carinate, 1.7-2 cm
long; dorsal oblong, with somewhat dilated, obtuse to
rounded apex; lateral sepals oblique at the base, narrow-
ly oblong, subacute. Petals oblanceolate to spatulate,
1.2-1.3 cm long, rounded. Lip hinged to the column
foot, 1.3-1.6 cm long, 7-10 mm broad when spread,
distinctly 3-lobate at about the middle; lateral lobes
small, obtuse; midlobe suborbicular to obovate, appar-
ently without callus. Column slender, semiterete, arcu-
ate, about 1 cm long, foot perhaps 2 mm long.
Apparently very rare; known only by a photo-
graph of the holotype and by the neotype. Prob-
ably epiphytic, but nothing is known of its ecol-
ogy. The type was collected in flower in May.
Endemic to Costa Rica; the Lankester specimen
was collected in Orosi.
Maxillaria piestopus is (or was) a distinct spe-
cies with a unique oblanceolate leaf and large,
triangular bracts of the scape (dried specimens),
which are characteristic of no other species
known from Costa Rica. Also, the triquetrous
ovary, oblanceolate petals, and large midlobe of
the lip are unique. This species seems most sim-
ilar to a Bolivian species described as Mormolyca
fuchsii J. T. Atwood, which has similar scape
bracts, a winged ovary, and otherwise similar
flowers. It is possible that Maxillaria piestopus
belongs with Mormolyca.
The above description is condensed from
Schlechter's original and augmented by measure-
ments taken from the neotype. The photograph of
the destroyed holotype is rejected as a neotype
because the photograph is not part of the original
collection and cannot be dissected. The photo-
graph, however, shows a drawn dissection show-
ing Schlechter's concept of M. piestopus that is
consistent with the neotype.
Maxillaria pittieri (Ames) L. O. Williams, Ann.
Missouri Bot. Card. 29:349. 1942. Ornithidium
pittieri Ames, Sched. Orch. 2:36-37. 1923.
TYPE: Costa Rica, San Isidro de Heredia, Pit-
tier 14048 (holotype: AMES; isotype: us). Figure
6B.
Plant a branched, somewhat straggly, mostly pendent
epiphyte to nearly 2 m long; each shoot 10-15 cm long;
stems somewhat curved. Roots mostly on the lower
shoots where anchored, to 5 mm in diameter. Pseudo-
bulbs lacking. Leaves 4-8 clustered near the stem apex,
articulate to the sheath; blade coriaceous, very narrowly
elliptic, 6-18 cm long, 1-2.2 cm wide, conduplicate at
the base, somewhat concave apically, acute. Inflores-
cences densely aggregated at the base of completed
shoots, the scapes about 2 cm long and hidden by the
leaf bases; ovary with pedicel 1-1.4 cm long. Flowers
mauve to pink, sometimes spotted with crimson, the lip
apex yellow to green. Sepals similar, ovate, somewhat
fleshy toward the apex, 7-9 mm long, 3.5-5 mm wide,
acute to obtuse. Petals obovate, similar to the sepals, 6-
7 mm long, 2.5-3.5 mm wide. Lip simple, rigidly adnate
to the column foot, 4-5 mm long, 2.5-3 mm wide; api-
cal half somewhat reflexed and quite fleshy, with a
fleshy, bifid callus at about the middle. Column stout,
straight, 2.5-3 mm long without the anther; foot 2.5-3
mm long, swollen apically; anther about 1 mm in di-
ameter.
Apparently an uncommon epiphyte of cloud
forests at 1500-2500 m. Flowering September,
November, January, and May. Costa Rica and
Panama.
Maxillaria pittieri differs from similar M. ful-
gens (Rchb. f.) L. O. Williams by its lack of pseu-
dobulbs, usually somewhat curved stems, and
somewhat larger, mauve to pink flowers (orange
to red in M. fulgens) produced at the base of the
most recently completed shoot. Flowers of M. ful-
70
FIELDIANA: BOTANY
gens are produced precociously within the bud
scales of a developing shoot.
Maxillaria cf. planicola C. Schweinf., Bot. Mus.
Leafl. 8:188. 1940. Based on Camaridium la-
tifolium Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 17:74. 1922. TYPE: Panama, on hills near
Panama City, Powell 8 (lectotype: AMES; isolec-
totype: AMES). Figure 11 A.
Plant poorly known, a rhizomatous epiphyte; rhizome
segments about 15 cm long between pseudobulbs, con-
cealed by imbricate, incurved bracts. Roots to about 1
mm in diameter. Pseudobulbs lance-ovoid, strongly
compressed laterally, about 7 cm long, 2 cm wide; base
concealed by nonfoliaceous sheaths; apex 1 -foliate. Leaf
shortly petiolate; blade thin, elliptic, 22-32 cm long, 5-
6 cm wide, acute. Inflorescences 2 or more per rhizome
bract of apparently immature shoots (mature fide Fl.
Pan.); ovary 1-1.5 cm long. Flower large, white, the lip
with yellow (spots?) on the lower half. Sepals narrowly
elliptic, 3-3.5 cm long, 7.5-9 mm wide, acute; lateral
sepals somewhat oblique. Petals ligulate, somewhat ob-
lanceolate, about 3 cm long, 6 mm wide, acute to obtuse.
Lip attached to the column foot, 3-lobate on the upper
half, the lower half curved in such a way as to form an
extra pair of more obscure lobes, about 1 .5 cm long and
1.5 cm wide when spread; lateral lobes embracing the
column, rounded; midlobe ovate, rounded, reflexed; cal-
lus a thick, fleshy ligule ending at about the middle, with
variously furcate hairs on the lower %. Column arcuate,
about 1 1 mm long excluding the anther, with nearly ob-
solete foot at the base. Capsule ellipsoid-ovoid, about 4
cm long.
Primary forests below 450 m. Panama and Cos-
ta Rica (Rio Agua Buena, Rincon de Osa).
The single Costa Rican collection with two
available duplicates (Liesner 2010 at MO, SEL) ex-
hibits fruit only but is most likely the above spe-
cies. Both specimens show the elongate rhizome
covered by incurved bracts, nonfoliaceous
sheaths, and laterally much flattened, 1 -foliate
pseudobulbs characteristic of M. planicola. Also,
the broad leaf at the pseudobulb apex, inflores-
cence size, and fruit with persistent column are
consistent with that species. The above descrip-
tion of the flower is based on analysis of the is-
olectotype (AMES).
Maxillaria ponerantha Rchb. f., Bonplandia 2:
17. 1854. TYPE: Venezuela, around Caracas,
Wagener s.n. (holotype: w). Figure 7E.
Plant small, to about 20 cm tall, somewhat straggly
with ascending, leafy shoots 1-2 cm apart and rooted
only at the base. Roots white, to about 0.5 mm in di-
ameter. Pseudobulbs apically 1 -foliate in Costa Rican
material (2-foliate in the type description), oblong, com-
pressed, to 1 .5 cm long and 5 mm wide, subtended when
young by 3 or more leaf-bearing, rugose sheaths. Leaves
oblong, 1-2.5 cm long, 2.5-6 mm wide, apically un-
equally bilobed. Inflorescence 1 per rhizome bract axil
of the immature shoot; scape about 5 mm long covered
by 2 or more acute scape bracts to 7 mm long and hid-
den from view by the rhizome bracts; ovary with pedicel
slightly exceeding the subtending floral bract, 7-8 mm
long. Flowers dark purple in Costa Rican material. Se-
pals elliptic-oblong, dorsally carinate apically, 8 mm
long, 2.5 mm wide, acute. Petals linear-oblong to ob-
lanceolate, 6 mm long, 1.5 mm wide, obtuse. Lip hinged
to the column foot, narrowly obovate, slightly 3-lobate,
the constriction at the final Vs forming the large midlobe,
margin minutely papillose; callus fleshy, oblong, thick-
ened and rounded in front, to about J/2 the lip. Column
arcuate, somewhat broadened apically, about 5 mm long;
foot about 1.5 mm long.
Epiphytes forming dense colonies in premon-
tane rain forests to about 700 m. Flowering June
to September. Northern South America and Costa
Rica (Puntarenas).
Plants conform to the type description of M.
ponerantha, but the flowers are much darker in
color and similar to a photograph of a plant from
Surinam published by M. C. M. Werkhoven (Or-
chids of Surinam, p. 156. 1986) under the same
name. Also, the pseudobulbs in Costa Rican ma-
terial are apically 1 -foliate, not 2-foliate. A low-
elevation species, it is expected to exhibit a broad
geographic range. Plants have often been mis-
identified as M. foliosa Ames & C. Schweinf. (=
M. acervata Rchb. f.), but that species is some-
what larger, has apically 2-foliate pseudobulbs (in
Costa Rican material), has slightly larger light
green flowers with a reddish lip, and occurs in
cloud forest regions at more than 1000 m. The
difference in elevation preference between these
two species doubtless provides them with little
opportunity for hybridization.
Maxillaria pseudoneglecta J. T. Atwood, Lin-
dleyana 8:30-31. 1993. Ornithidium anceps
Rchb. f., Beitr. Orchid. -K.C. Amer. 75-76.
1866. TYPE: Costa Rica, Cartago, Wendland
s.n. (holotype: w). Not Maxillaria anceps Ames
& C. Schweinf. 1930. Figure 10B.
Plant a straggly, somewhat pendent epiphyte forming
loose clumps, rooted only along the basal shoots; rhi-
zome elongate, to 6 mm in diameter. Roots rarely more
than 0.5 mm in diameter. Pseudobulbs cylindric to el-
lipsoid, rarely globose, compressed, 1 .5-3 cm long, 0.7-
1.8 cm wide, spaced 1-8 cm apart, subtended by a pair
of subulate bracts, apically 1 -foliate. Leaf shortly peti-
olate; blade lanceolate, 5-15 cm long, 1-2.8 cm wide,
acute. Inflorescence a fascicle of single-flowered scapes
at the base of the most recently matured pseudobulb;
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
71
scapes less than 1 cm long, concealed by the rhizome
bracts; ovary and pedicel 4.5-6 mm long, concealed by
inflated bracts. Flowers variously yellow (lip with or-
ange midlobe), entirely orange, or white and lip with
orange midlobe. Sepals concave and not well spread;
dorsal oblong-ovate, 5.5-7.5 mm long, 2.2-3.5 mm
wide, apiculate; lateral about Vz connate, more or less
obliquely ovate, apiculate, 5-9 mm long, 4-5 mm wide.
Petals oblong, constricted on the upper half, 4-6.5 mm
long, 1 .5-2 mm wide, obtuse. Lip continuous with and
not distinct from the column foot, 3-lobate, 5-7 mm
long, (4.5)5-7 mm across the lateral lobes when spread;
side lobes deep with convex lateral margins; midlobe
fleshy, triangular-ovate, occupying about 1A the lip, with
a transverse ridgelike callus at the base of the midlobe
connecting the lateral lobes. Column stout, 1.5-2 mm
long excluding the anther; anther globose, about 1 mm
in diameter.
Epiphytic in premontane and montane forests
at (5007)900-2200 m. Most seem to occur in ev-
ergreen forests, but plants growing in semideci-
duous forests with Plumeria rubra and Cattleya
skinneri are known. Flowering June to September.
Costa Rica (Cordillera de Tilaran) to Boquete,
Panama.
Maxillaria pseudoneglecta is most similar to M.
neglecta but has statistically larger flowers. Also,
the lip has much broader lateral lobes with convex
margins, and the leaves are broader and more lan-
ceolate.
Maxillaria punctostriata Rchb. f., Linnaea 41:
23. 1877. TYPE: Costa Rica, Low s.n. (holo-
type: w). Maxillaria rubrilabia Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:236. 1923.
TYPE: Costa Rica, San Ramon, 1050 m, Brenes
151 (destroyed); Todzia 334 (neotype: F). Fig-
ure 22C.
Plant a cespitose epiphyte to about 30 cm tall. Roots
white, to about 1 .5 mm in diameter. Pseudobulbs com-
pressed, ovoid to suborbicular, 2.5-6 cm long, 2-3 cm
wide; base subtended by nonfoliaceous, subulate bracts;
apex 1 -foliate. Leaves indistinctly to shortly petiolate
(<'/5 blade); blade narrowly elliptic, 10-25 cm long, 2-
4 cm wide; apex obtuse, emarginate, and apiculate. In-
florescence 1 or more per rhizome bract axil; to about
10 cm long; ovary with pedicel 1.8-2.4 cm long, terete,
exceeded by a cucullate, acute floral bract. Flowers yel-
low to orange with lines of small purple dots; lip deep
maroon. Sepals similar, broadly oblanceolate to elliptic,
acute; dorsal somewhat concave, 2.2-2.7 cm long, 6.5-
8.5 mm wide; lateral sepals somewhat oblique, 2.1-2.7
cm long, 6-8.5 mm wide. Petals broadly oblanceolate,
1.6-2.1 cm long, 6-7 mm wide, acute. Lip hinged to
the column foot, recurved, 3-lobate below the middle,
1.2-1.8 cm long, 7-8 mm wide when spread; lateral
lobes rounded; midlobe ovate, fleshy, 8-10 mm long, 6-
7.5 mm wide, acute to obtuse and thickened near the
apex; callus terminating at about the divergence of the
lateral lobes, oblong to somewhat trapezoid, thickened
in front. Column stout, arcuate, 9-1 1 mm long exclud-
ing the anther; foot 3-4 mm long. Capsule about 4.5
cm long.
Common in evergreen to semideciduous cloud
forests at 1100-2400 m. Flowering March to Au-
gust and November; fruiting in July. Costa Rica;
possibly in northern Nicaragua masquerading un-
der the name M. cucullata Lindl. owing to the
need for a revision of the M. cucullata complex.
At least three species of the M. cucullata Lindl.
complex occur in Costa Rica, this one with yel-
lowish sepals with maroon dots arranged in lines
and a nearly black lip, another (M. cf. obscura
Lind. & Rchb. f.) with much smaller maroon or
yellowish flowers tinted with maroon, and a third
recently described as M. cedralensis J. T. Atwood
& Mora de Retana with larger, cream flowers. If
the painting and presumably the type of M. cu-
cullata (from Mexico) showing a long peduncle
and a yellow flower stained with purple is accu-
rate, this species probably does not occur in Costa
Rica. There is little doubt that the name M. punc-
tostriata applies to Costa Rican material that
match the above description. The neotype of M.
rubrilabia matches M. puntostriata very well and
is here accepted as a synonym.
There may be a fourth species in the north with
larger yellow to orange flowers with a very large,
inflated floral bract concealing the ovary. A spec-
imen (Mora 60, INB) collected at Estacion Cacao
at 1100 m, Cordillera de Guanacaste, and Canton
de Liberia appears to be M. mombachoensis Hell-
er ex J. T. Atwood, which was originally de-
scribed from adjacent southern Nicaragua, or it
may be new. The dried plant has rather thick
leaves and a short scape, suggesting a relatively dry
or windy cloud forest habitat. (See Addendum.)
Maxillaria quadrata Ames & Correll, Bot. Mus.
Leafl. 6:16. 1943. Ornithidium lankesteri Ames,
Sched. Orch. 4:52-53. 1923. TYPE: Costa
Rica, San Cristobal, Lankester & Sancho 421
(holotype: AMES). Figure 5B.
Plant a large, monopodial, cane-forming epiphyte or
terrestrial to about 1 m tall, probably becoming pendent
with size; juveniles unknown, but probably cespitose
with each shoot terminating in a pseudobulb; mature
stems sheathed with closely spaced, conduplicate leaves,
mostly deciduous near the base with sheaths fragmenting
into fibers. Roots slender, to about 1 mm in diameter.
Pseudobulbs unknown; absent on mature stems. Leaves
articulate, coriaceous, keeled beneath, shortly petiolate
above the articulation, the blades 5-18 cm long, 1-2.8
72
FIELDIANA: BOTANY
cm wide, the apex unequally 2-lobate. Inflorescences
several-many per leaf axil; scapes to about 10 cm long,
not much exerted beyond the subtending leaf blade,
clothed in acute sheaths; ovary with pedicel about 1.2
cm long, usually exceeded by the subtending floral bract.
Flowers with somewhat recurved segments, pink to
white. Sepals similar, oblong-oblanceolate, 1.7-1.9 cm
long, 4-5 mm wide, the apex acute. Petals elliptic-ob-
lanceolate, 1.2-1.5 cm long, 3.5-4.5 mm wide, apex
acute to acuminate. Lip rigid with the column foot, more
or less straight and not saccate on the lower l/i, 3-lobate
above the middle, 4-5 mm long, 2.5-3 mm wide in
natural position; lateral lobes well extended beyond the
sinuses, rounded; midlobe ovate, rounded to emarginate,
recurved; callus a truncate ligule ending just below the
lobe sinuses, with an additional 3 or more papillae at
the base. Column stout, 3-4 mm long excluding the
anther; the anther about 1 .5 mm in diameter.
Maxillaria quadrata appears to be a rare epi-
phyte or terrestrial in montane cloud forests at
1500-1900 (2400) m. Flowering December to
March. Endemic to Costa Rica; to be expected in
western Panama.
From its closest relative, M. adolphii Ames &
Correll, it differs by its occurrence at lower ele-
vations and by its more or less straight, nonsac-
cate lip with more elongate lateral lobes, which
usually are concave, embracing the column.
Some herbarium specimens from upper eleva-
tions (to 2400 m) may actually represent hybrids
of M. adolphii with M. biolleyi (Schltr.) L. O. Wil-
liams. The latter species as a potential parent
would explain the relatively straight, simple lip in
flowers that otherwise appear similar to flowers
of M. quadrata.
Maxillaria ramonensis Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:235. 1923. TYPE:
Costa Rica, Santiago de San Ramon. Brenes
154 (lectotype: CR; isolectotype: AMES). Maxil-
laria brevipes Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19:302. 1923. TYPE: Costa Rica, La
Palma, Werckle 110 (photo of type: AMES). Fig-
ure 23D.
Plant a cespitose epiphyte forming small clumps 6-
20 cm tall. Roots filiform, white, to 0.5 mm in diameter.
Pseudobulbs approximate, ovoid to suborbicular, 1-2
cm long, to about 2 cm wide, subtended by nonfolia-
ceous bracts, or with a single foliaceous bract, the apex
1 -foliate. Leaves long-petiolate; sheath of foliaceous
bracts much broader than the petiole; petioles canalicu-
late, 1-4 cm long, to about W the blade length; blade
well defined, narrowly to broadly elliptic, rarely some-
what oblanceolate, acute to subattenuate apically. Inflo-
rescences 1 or more per rhizome bract, the scape 1-3.5
cm long; ovary with pedicel 1-1.2 cm long, exceeded
by the acute floral bract. Flowers apparently not opening
broadly; sepals and petals translucent cream-white with
white veins aging reddish or brownish; lip white strong-
ly suffused with red, the apex usually dark red. Sepals
dissimilar; dorsal oblong-ovate, 9-12 mm long, 4.5-5.5
mm wide with obtuse, mucronate apex; lateral sepals
inserted on the column foot, oblique, triangular, 1.5-1.8
cm long, 5-7.5 mm wide with bluntly acute to obtuse
apex. Petals oblique at the base, ligulate to ovate, acute,
1-1.3 cm long, 3.5-4.5 mm wide; apex acute to obtuse.
Lip hinged to the column foot, cuneate from the base,
the apical !/5-Vi 3-lobate, 1.1-1.3 cm long, 5-6.5 mm
wide when spread; lateral lobes rounded to somewhat
angular in front and embracing the column; midlobe
ovate, rounded in front. Column stout, arcuate, 4-5 mm
long excluding the anther; foot 8-9 mm long; anther 2-
2.5 mm in diameter; pollinia 4, supported on a squarrose
stipe.
Epiphytic in premontane and lower montane
rain forests at 500-1200(1500) m. Flowering May
to December. Nicaragua to Panama; also reported
from Honduras.
Maxillaria ramonensis lacks the fringed anther
bed seen in M. confusa and has a much more stout
column and a longer column foot than flowers of
either M. brachybulbon Schltr. or M. acostae
Schltr. Maxillaria brevipes is here interpreted as
a small M. ramonensis. Both names are based on
plants with stout columns and long column feet.
The above description includes forms from Pan-
ama with a long midlobe and may prove to rep-
resent a different species.
Maxillaria reichenheimiana Endres & Rchb. f.,
Card. Chron. 1678. 1871. TYPE: Costa Rica,
M. Endres (holotype: w). Figure 24A.
Plant epiphytic, cespitose, forming small clumps to
about 15 cm tall. Roots to about 1 mm in diameter.
Pseudobulbs about 1-2 cm long, orbicular to ovoid,
compressed, usually hidden by subtending bracts, 1(2)
of which may bear a leaf blade. Leaves shortly and dis-
tinctly petiolate, the blade round to elliptic, 4.5-1 1 cm
long, 2-4.5 cm wide, apically emarginate to obtuse and
very rarely acute. Inflorescences usually held above the
leaves, the scapes 5(2)-8 cm long, nearly concealed by
4-6 acute to obtuse bracts 1-2.5 cm long; ovary with
pedicel shorter than the subtending floral bract, lepidote,
1.5-2.5 cm long. Flowers spidery, pale yellow aging
brown. Sepals lanceolate, acuminate, 3-4 cm long; dor-
sal sepal 2.5 mm wide; lateral sepals 4 mm wide. Petals
falcate, lanceolate, about 2.5 cm long, 4 mm wide, at-
tenuate. Lip obovate, the apical !/6 3-lobate, 1 .3 cm long,
about 6 mm wide; midlobe fleshy; callus ligulate, to just
below % the lip, parabolic, thickened, and somewhat el-
evated in front. Column 1.2 cm long including the 8
mm-long foot; anther bed papillose, pollinia 4, with sad-
dle-shaped viscidium.
Epiphytic in premontane and lower montane
rain forests at 1000-1500 m. Costa Rica south to
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
73
Venezuela and Ecuador. Flowering April to Sep-
tember. Reports from Nicaragua are most likely
based on M. pachyacron.
Maxillaria reichenheimiana is readily distin-
guished from M. pachyacron by its broad, usually
emarginate leaves with white spots. The flowers
of both species are remarkably similar, but the se-
pals of M. reichenheimiana are usually shorter,
and the column foot is distinctly longer. Both spe-
cies are sympatric over parts of the range, e.g., at
Monteverde, Costa Rica. Both species are isolated
phenologically because M. pachyacron flowers
from October to March.
Maxillaria ringens Rchb. f., Ann. Bot. Syst. 6:
523. 1863. TYPE: Costa Rica, Endres 8 (lec-
totype selected here: w). ?M. yzabalana S. Wat-
son, Proc. Amer. Acad. Arts 23:266. 1888. M.
tuerckheimii Schltr, Repert. Spec. Nov. Regni
Veg. 10:295-296. 1912. TYPE: Guatemala,
Cubilguitz, von Ttirckheim 790 (drawing of
type: AMES). M. rouseauae Schltr., Beih. Bot.
Centralbl. 36(2):413-414. 1918. TYPE: Pana-
ma, "Im Gebiete des Panama-Kanals," Rou-
se au s.n., Powell 115 (neotype selected here:
MO 955962). M. pubilabia Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 17:70-71. 1922. TYPE:
Panama, "Auf Hiigeln bei Panama City," Pow-
ell 214 (lectotype: AMES; isolectotype: MO). M.
amparoana Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19:54-55. 1923. TYPE: Costa Rica,
Carillo, Werckle 7. M. lactea Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:233. 1923, not
1925. TYPE: Costa Rica, San Mateo, Brenes
143 (isotype: AMES). Figure 20A.
Plant an often densely cespitose epiphyte to about 50
cm tall. Roots to about 1 mm in diameter. Pseudobulbs
ovoid, not much compressed laterally, rugose or some-
times furrowed, 3-6 cm long, 1 .5-3 cm wide, subtended
by nonfoliaceous sheaths lacking articulations and soon
fragmenting with age, apex 1 -foliate. Leaves with dis-
tinct, canaliculate petiole to 10 cm long; blade elliptic-
oblong, carinate and somewhat glaucous beneath, 11.5-
38 cm long, 2-4.5 cm wide, the apex bluntly acute to
rounded. Inflorescences several per rhizome bract;
scape 3-13 cm long, concealed by bracts; ovary with
pedicel (1.2)2-3.7 cm long. Flowers white, the sepals
yellow within, aging pinkish or tan; lip white with yel-
low callus, usually with purple veined lateral lobes and
spots on midlobe. Sepals somewhat chartaceous and
canaliculate, straight, oblong, with apex acute to obtuse
or acuminate; dorsal 2.5-4 cm long, 4-6 mm wide; lat-
eral sepals 2.7-4.2 cm long, 4.5-7 mm wide. Petals nar-
rowly lanceolate, incurved, 2.5-4 cm long, 3-5 mm
wide. Lip ovate in general outline, (0.9)1.1-1.5 cm long,
(4)6-8.5 mm wide when spread, 3-lobate on the apical
%; side lobes embracing the column, obtuse; midlobe
fleshy, warty, and suborbicular. Column stout, clavate,
nearly straight, 6.5-8 mm long excluding foot or anther;
foot 4-5.5 mm long; anther about 2 mm long, with a
dorsal crest. Capsule ellipsoid, 3-4.5 cm long.
Epiphytic at (100)300-1600 m. Flowering Au-
gust to November in Costa Rica, July to Decem-
ber in Panama, and June to September in Nica-
ragua to Mexico. Mexico to Panama with doubtful
reports from Colombia (probably M. brunnea Lin-
den & Rchb.f.).
Maxillaria ringens is here interpreted as a vari-
able species. It is characterized by the often large,
cespitose plants with strongly fragrant clusters of
white flowers with narrow segments. Schlechter
recognized various entities based on scape length,
the presence or absence of hairs on the lip, and
flower color. Under apparently different cultural
conditions, singular clones at Selby Gardens
sometimes exhibit the long scapes of M. ringens
or the short scapes of M. amparoana. Finding
hairs on the lip probably depends on the quality
of the microscope because they were observed in
virtually all rehydrated flowers (n = 10) broadly
sampled among available specimens. Flower color
ages pinkish or tan, and color descriptions by col-
lectors are by no means uniform. In Costa Rica,
live specimens collected from upper elevations
are somewhat smaller, have smoother pseudo-
bulbs, and may be different. One specimen at CR
(Robinson & Vaughn 1243) with somewhat larger
flowers and long scapes had flowered in April and
may be M. fuerstenbergiana Schltr., a species un-
known from Central America.
There has been some confusion concerning the
application of the name M. ringens. Reichen-
bach's original description indicates "labello se-
palis ter breviore," or lip one-third as long as
(three times shorter than) the sepals. This phrase
excludes M. brunnea, in which the lip is at least
half as long as the sepals.
Although no original materials were found of
M. rousseauae, the designated neotype is so iden-
tified by Schlechter.
Maxillaria rodrigueziana Atwood & Mora-Re-
tana, Icon. PI. Trap. 14: t. 1361. 1989. TYPE:
Panama, El Cope, T. Antonio 2137 (holotype:
MO; isotype: SEL). Figure 23A.
Plant a large, cespitose epiphyte, to 35 cm high.
Roots white, to 1 mm in diameter. Pseudobulbs ovoid,
rugose, 1 -foliate, 4-5 cm long and 3-4 cm wide, sub-
tended and often hidden from view by the inflated blade-
bearing rhizome bract base, which is usually abruptly
74
FIELDIANA: BOTANY
narrowed at the articulation. Leaf blades coriaceous, el-
liptic, obtuse to rounded, 18-30 cm long, 4.5-8 cm
wide, supported by a long petiole 5-15 cm long. Inflo-
rescences 14-20 cm tall, mostly hidden by 4-7 in-
rolled, obtuse bracts 2-6.5 cm long; ovary with pedicel
about 2.5 cm long, concealed by the conspicuous, con-
duplicate, acute floral bract 5-8 cm long. Flowers
creamy white, fragrant, the sepals turning greenish to-
ward the apex, lip base white, apically yellow, and lined
on the sides with red. Sepals long attenuate, 12-16 cm
long with the dorsal slightly shorter than the lateral se-
pals, 1.0-1.3 cm wide at the base. Petals long-attenuate,
10-14 cm long, 7-9 mm wide at the base. Lip ovate,
3-lobate on the apical 1A, about 2 cm long and 1.5 cm
wide; midlobe broadly rounded and somewhat reflexed;
lateral lobes folded and embracing the column; callus
ligulate, puberulent, to about the lh the lip. Column cy-
lindric, about 1.5-1.8 cm long, forming a foot 8 mm
long at the base; pollinia 4, lacking a stipe but supported
on a saddle-shaped viscidium.
Large and apparently rare epiphyte in evergreen
forests at 700-750 m. Flowering June, October,
and November. Costa Rica and Panama.
This species is one of the most spectacular Cen-
tral American maxillarias and is unique for its
sheer size. It is easily distinguished from its South
American sister species, M. speciosa Rchb. f., by
the more leathery flowers and by the spotting on
the sepals and petals restricted to the lower half.
The midlobe of the lip is less than a quarter the
lip length in this species and nearly half the lip
length in M. speciosa. One might regard it as a
subspecies of M. speciosa, but the consistently
different features mentioned argue for recognition
of two distinct species. According to R. L. Dress-
ier, M. rodrigueziana has a heavy fragrance of
cheap perfume that is a bit much in close quarters.
Maxillaria sanguinea Rolfe, Bull. Misc. Inform.
1895:8. TYPE: Panama, Chiriqui, James
O'Brien s.n. (holotype: K). Figure 10E.
Plant epiphytic with short to somewhat elongate rhi-
zomes forming large clumps. Roots white, to 1 .5 mm in
diameter. Pseudobulbs 0.5-4 cm apart, fusiform-ellip-
soid, apically 1 -foliate, usually tapering at both ends, 1-
2.5 cm long, 0.5-1.3 cm broad. Leaves linear, 20-35
cm long and 2-3.5 mm wide. Inflorescence 1 per rhi-
zome bract on the rhizome of the most recently com-
pleted shoots; scapes about 1 cm long, nearly concealed
by the rhizome bract and covered by scape bracts; ovary
with pedicel 2-3.5 cm long. Flowers slightly fragrant,
the perianth greenish with strong overlay of red, lip
flushed with red, apically white; callus maroon. Sepals
oblong-elliptic, 1.8-2.7 cm long, 4-9 mm wide, the lat-
eral sepals forming a mentum around the column foot.
Petals somewhat falcate and arcuate, 1.5-2.3 cm long,
3.5-7 mm wide. Lip obovate, apically obtuse to trun-
cate, laterally constricted on the apical Vi, rendering it
somewhat 3-lobate, 1.5-2.0 cm long, 6.5-9.5 mm wide;
callus ligulate on the lower Vi, apically recurved, trun-
cate. Column arcuate, 1.1-1.5 cm long; anther cap pa-
pillose; pollinia 4, supported on a saddle-shaped visci-
dium.
Epiphyte of larger branches in lowland tropical
and premontane rain forests at 10-900 m. Flow-
ering January and February. Costa Rica and west-
ern Panama; erroneously reported from Florida
(USA), apparently on the basis of a specimen
planted in the wild (fide J. Beckner, pers. comm.).
Maxillaria sanguinea is similar to M. tenuifol-
ia, which it replaces geographically. The latter
species, ranging from Mexico to Nicaragua, flow-
ers from March to May, while M. sanguinea flow-
ers in January and February. The flowers of M.
tenuifolia have a pronounced fragrance resem-
bling coconut (or crayons), whereas M. sanguinea
has a slightly fruity fragrance. The lip apex is
broader in M. sanguinea and is usually white, at
least apically, and only minutely spotted (pep-
pered). The lip apex of M. tenuifolia is always
conspicuously spotted with a few large blotches.
Vegetatively, M. sanguinea usually has fusiform
pseudobulbs, while those of M. tenuifolia are usu-
ally cylindric. The leaves of M. sanguinea rarely
exceed 3 mm wide, and the leaf width of M. ten-
uifolia rarely measures less than 3 mm.
Maxillaria schistostele Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:303-304. 1923.
TYPE: Costa Rica, near San Ramon, Acosta
s.n. (photo of type: AMES).
Plant a somewhat spreading, ascending epiphyte; rhi-
zomes 4-5 mm in diameter, the segments about 1 .5 cm
between pseudobulbs. Pseudobulbs ovoid or oblong, 1-
1.3 cm long, subtended by nonfoliaceous sheaths, the
apex 1 -foliate. Leaves linear-ligulate, 6-9 cm long, 6-8
mm wide, the base somewhat narrowed, apex obtuse and
emarginate. Inflorescence slender, borne at the pseudo-
bulb base; scape 6-8 cm long; ovary with pedicel about
1.3 cm long, somewhat exceeded by the floral bract.
Flowers rigid in dried specimens, glabrous except for
the lip; color unknown. Sepals linear or linear lanceo-
late, 2.2 cm long; lateral sepals oblique, forming with
the column foot an obtuse, shortly conic mentum. Petals
linear, acuminate, 1.6 cm long. Lip minutely papillose,
1.25 cm long including the distinct claw (2.5 mm long)
at the base; forming an abruptly lanceolate blade 1 cm
long, 5 mm wide; callus at the base obovate, obtusely
sulcate. Column arcuate, 6 mm long, the apex thick-
ened, the anther bed with fimbriate margin; foot at the
base 2 mm long.
Maxillaria schistostele is known only by a pho-
tograph of the type collection but is clearly related
to the M. acuminata complex in South America,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
75
a group characterized by the rhizomatous habit,
usually 2-foliate pseudobulbs (1 -foliate in this
species), wiry scapes, and entire lips. Nothing is
known of its ecology or phenology. The above
description is translated and condensed from
Schlechter's original description.
Maxillaria schlechteriana J. T. Atwood, Lindley-
ana 9:231. 1994. Basionym: Camaridium im-
bricatum Schltr., Beih. Bot. Centralbl. 36:415-
416. 1918. TYPE: Costa Rica, La Palma, Werc-
kle (isotype: AMES). Ornithidium schlechterian-
um C. Schweinf., nom. illeg., Bot. Mus. Leafl.
4:94. 1937. Maxillaria wrightii (Schltr.) Ames
& Correll var. imbricata (Schltr.) Ames & Cor-
rell, Bot. Mus. Leafl. 11:18. 1943. Not Maxil-
laria imbricata Barb. Rodr. Figure 14 A.
Plant a straggly, erect to pendent, cane-forming epi-
phyte to about 50 cm tall; shoot slender, terminated by
a pseudobulb, to 30 cm long; rhizome wiry, concealed
by scales when young, apparently turning black with
age. Roots white, to 1 mm in diameter. Pseudobulbs
compressed, ovoid to suborbicular, 1-3 cm long, mostly
concealed at the base by foliaceous sheathes, apically 1 -
foliate. Leaves with petiole broadened below the artic-
ulation and about V4— % the blade length; blade elliptic-
lanceolate, carinate beneath, 2.5-18 cm long, 8-40 mm
wide, acute to obtuse. Inflorescence 1 per nonfoliaceous
bract axil, forming on the developing shoot; scapes to
about 2 cm long; ovary with pedicel 3.5-6 mm long,
subtended by a somewhat cucullate floral bract 7-1 1 mm
long. Flowers white or light lavender to rose; column
often darker. Sepals similar, oblong, acute; dorsal 1.2-
1.5 cm long, 4 mm wide; lateral sepals somewhat
oblique, 1.25-1.5 cm long, 4-5 mm wide. Petals ob-
long, 1-1.4 cm long, about 3 mm wide, acute. Lip rigid
with the column foot, lateral view somewhat sigmoid
but base not saccate, 6-8 mm long in natural position;
lateral lobes extending to % the lip; midlobe fleshy; cal-
lus tonguelike between the lateral lobes. Column 2.5-3
mm long without anther; anther about 1 mm long; pol-
linia 4, with somewhat square stipe. Capsule ellipsoid
to globose, about 1 cm long.
Epiphytic and terrestrial in montane cloud for-
ests at (1000)1400-2800 m. Flowering August to
March. Costa Rica and Panama; doubtfully re-
ported from Nicaragua as M. wrightii var. imbri-
cata.
This species is most similar to M. paleata
(Rchb. f.) Ames & Correll, with which it is often
sympatric. However, that species has larger,
darker, red-purple flowers with a lower column-
to-lip-length ratio. Also, the profile of the lip is
somewhat different. Among herbarium speci-
mens, no intermediates suggesting hybrids have
been seen.
Maxillaria scorpioidea Kraenzl., Svensk. Vet.
Akad. Handl. 46:71. 1911. TYPE: Brazil, Matto
Grosso. IMaxillaria rhodosticta Kraenzl., Re-
pert. Spec. Nov. Regni Veg. 24:223. 1928.
TYPE: Mexico. Figure 8D.
Plant a straggly epiphyte, or if cespitose consisting
of several ascending successively borne shoots to 2(3)
cm apart. Roots white, to 1 mm in diameter. Pseudo-
bulbs narrowly ovoid 2-5 cm long, 1-1.7 cm wide,
compressed, conspicuously ribbed when dried, subtend-
ed by 1-2 leaf-bearing rhizome bracts, apically (1)2-3-
foliate. Leaf blades linear to very narrowly elliptic, 2-
24 cm long, 6-15 mm wide. Inflorescence 1 per rhi-
zome bract axil; scapes 1-2 cm long, concealed by 2 or
more acute, somewhat inflated bracts; ovary with pedi-
cel 1.6-2 cm long, subtended by a pair of glumaceous
bracts, the uppermost 16-21 mm long. Flowers green-
ish, yellowish, or buff, lip with maroon spots on the
margins, often tipped with maroon. Sepals similar; dor-
sal narrowly lance-elliptic, acute, apically and dorsally
carinate, 18-25 mm long, to 5 mm wide; lateral sepals
the same size, lanceolate, acute. Petals narrowly elliptic,
falcate, 16-22 mm long, 2.5-4 mm wide. Lip arcuate,
narrowly elliptic, 15-18 mm long, the apex thickened,
oily; callus linear, oily to just below the middle. Column
slender, somewhat arcuate, 12-14 mm long; foot 2 mm
long; pollinia 4, with elongate stipe and saddle-shaped
viscidium. Capsule somewhat down-curved, ellipsoid,
2-3 cm long including the beak.
Epiphytic in moist forests at 500-1800 m.
Flowering mostly November to March. Mexico
(southern Chiapas) to western El Salvador, Costa
Rica and western Panama, and Brazil and Ecua-
dor. In Central America this species is known only
from the Pacific drainage.
The identification of the name is based on the
original description with illustration showing consis-
tent floral dimensions and color. The name M. rho-
dosticta probably applies to this species because the
original description indicates nearly exact floral di-
mensions and the spotting on the lip margin. Max-
illaria scorpioidea has for more than a century
caused confusion with M. friedrichsthalii Rchb. f.
and M. aciantha Rchb. f. because its floral size is
intermediate between the other two. The close spac-
ing of the shoots and spotting pattern on the lip
margin are usually good features for field recogni-
tion. One should look at column length (excluding
the foot), which is shorter in M. friedrichsthalii and
longer in M. aciantha. Floral size and color are usu-
ally closer to those of M. friedrichsthalii than M.
aciantha, but, as with the latter species, M. scor-
pioidea has a much longer bract subtending the
flower. Except perhaps in western Panama, M. scor-
pioidea is sympatric only with M. friedrichsthalii in
its more northern range, and it probably does not
76
FIELDIANA: BOTANY
occur much below 500 m. In dried specimens Af.
scorpioidea often has a golden, waxy appearance
that is lacking in the other two species.
Maxillaria serrulata Ames & Correll, Bot. Mus.
Leafl. 11:16. 1943. Camaridium amparoanum
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
56-57. 1923. TYPE: Costa Rica, San Jeronimo,
Werckle 122 (drawing of type: AMES). Ornithi-
dium amparoanum (Schltr.) Pabst, Bradea 2:87,
1976. Not Maxillaria amparoana Schltr. Figure
17 A.
Plant a somewhat straggly, cane-forming epiphyte, to
about 50 cm high; stems upright or pendent, each ter-
minated by a pseudobulb; rhizome conspicuous, 2-10
cm long. Roots less than 1 mm in diameter. Pseudo-
bulbs ovoid, 1.5-2.5 cm long, protected at the base by
1-3 leaf-bearing, rugose sheaths; apex 1 -foliate. Leaves
articulate, without petiole; blades ligulate, obtuse to
emarginate, abaxially keeled and mucronate, to 13 cm
long, 2.2 cm wide. Inflorescences appearing at the base
of mature pseudobulbs with the flush of the new growth;
scapes to about 3 cm long, concealed by compressed,
subulate bracts; ovary with pedicel to 1 cm long, much
exceeded by the subtending, acute floral bract to 3 cm
long. Flowers dark red with some green within. Sepals
similar, obovate, 2.5 cm long, 1 .5 cm wide, obtuse; dor-
sal somewhat concave; lateral sepals somewhat oblique
at the base. Petals obovate, 1.8-2 cm long, 9-11 mm
wide. Lip complex, hinged to the column foot, 3-lobate
below the middle, about 1 cm long, 8-9 mm across the
midlobe; lateral lobes rounded and erect around the col-
umn; midlobe suborbicular, with a dentate margin; callus
of about 7 keels higher at the base and becoming lower
apically. Column arcuate, 8 mm long excluding the an-
ther; anther 1.5 mm in diameter; foot 3 mm long.
Probably epiphytic but poorly known ecologi-
cally; 1400-2000 m. Flowering July and August.
Endemic to Costa Rica.
It is easily distinguished by the dark red flowers
and serrulate lip but is poorly known despite the
handsome flowers. It seems to be related to other
species, including M. sigmoidea, but little is
known of some aspects of the plant, e.g., whether
the petioles of the foliaceous rhizome bracts are
abruptly narrowed.
Maxillaria sigmoidea (C. Schweinf.) Ames & Cor-
rell, Bot. Mus. Leafl. 11:16. 1943. Ornithidium
sigmoideum C. Schweinf., Bot. Mus. Leafl. 4:
121-122. 1937. TYPE: Costa Rica, Cerro Gallito,
Valeric 72 (holotype: AMES). Figure 13D.
Plant somewhat straggly, epiphytic, or terrestrial,
erect or pendent when large, to about 1 m long; stems
elongate, often branched, each shoot to 30 cm long and
terminated by a pseudobulb. Roots white, to 1 mm in
diameter. Pseudobulbs ovoid to suborbicular, 1-4 cm
long and the uppermost strongly reduced, usually con-
cealed by 1-3 foliaceous sheaths, apically 1 -foliate.
Leaves with petiole up to about V6 the blade length;
blade oblong-elliptic, keeled abaxially, 6-27 cm long,
1.5-5.5 cm wide, much reduced in distal shoots; apex
acute, acuminate, or somewhat attenuate. Inflorescences
a single-flowered scape borne singly within nonfolia-
ceous, scarious, acute sheaths of the developing shoot;
scape to 5 cm long; ovary with pedicel 1-1.5 cm long
subtended by a somewhat membranaceous, cucullate,
acute bract 1.5-2.1 cm long. Flowers light pink to lav-
ender with white lip striped with lavender; column often
yellowish apically. Sepals similar, lanceolate-elliptic, the
lateral somewhat oblique, 1.8-2.4 cm long, 6-8 mm
wide, acute or shortly acuminate. Petals elliptic-oblan-
ceolate, 2-2.2 cm long, 5-5.5 mm wide, acute. Lip com-
plex for the genus, rigid with the column foot, forming
a deeply saccate base, the apical % sharply reflexed and
appearing sigmoid laterally, 9-13 mm long, to 6 mm
wide with the lateral lobes spread; lateral lobes falcate,
diverging near the upper third, embracing the column
apex; midlobe ovate, thickened apically; callus fleshy,
continuous with the lateral lobes, rounded to somewhat
3-dentate in front, exceeding the sinuses between lobes.
Column cylindric, somewhat arcuate, broadened above,
8-12 mm long; with short foot to about 2 mm long at
the base; anther shortly beaked; with protruding visci-
dium; pollinia 4. Capsule 1.5 cm long.
A plant of windy cloud forests at 1400-2100
m. Flowering October to December and February
to April. Costa Rica and Panama.
The unmistakable sigmoid appearance of the lip
is unique among Costa Rican Maxillaria; other-
wise the flowers appear similar to those of a large
M. schlechteriana J. T. Atwood or a small M. va-
ginalis Rchb. f.
Maxillaria strumata (Endres & Rchb. f.) Ames
& Correll, Bot. Mus. Leafl. 11:17. 1943. Orni-
thidium strumatum Endres & Rchb. f., Gard.
Chron. 2:772. 1874. TYPE: Costa Rica, Endres
281 (holotype: w; isotype: AMES). Sepalosaccus
strumatum (Endr. & Rchb. f.) Garay, Caldasia
8:525. 1962. Sepalosaccus humilis Schltr. Re-
pert. Spec. Nov. Regni Veg. Beih. 19:244-246.
1923. TYPE: Costa Rica, San Pedro de San Ra-
mon, Brenes 130 (lectotype: AMES). Figure 19B.
Plant a densely cespitose epiphyte to about 6 cm tall.
Roots filiform, < 0.5 mm in diameter. Pseudobulbs ap-
proximate, narrowly ovoid to cylindric, to about 5 cm
tall, 6 mm wide, subtended when young by 1-2 folia-
ceous bracts, the apex 1 -foliate. Leaf blades thin, nar-
rowly elliptic to oblong, obtuse, to 5.5 cm long, 8 mm
wide, the apical leaf shortly petiolate. Inflorescence 1
per nonfoliaceous bract axil of the rhizome, to about 1 .5
cm long; ovary with pedicel 4.5-6 mm long, exceeded
by the subtending acuminate floral bract. Flowers white,
lip apex yellow, anther dark purple. Sepals dissimilar;
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
77
dorsal oblong, concave at the base, recurved at the ob-
tuse apex, 4-5 mm long, about 1.2 mm wide; lateral
sepals about % connate, each oblong, 7-8 mm long. Pet-
als oblong, 4-4.5 mm long, 1-1.2 mm wide, obtuse. Lip
subsaccate and continuous with the column foot, some-
what 3-lobate on the apical %, about 4.5-5 mm long
when spread; lateral lobes broadly rounded and embrac-
ing the column, connected at the apex by a transverse
tonguelike callus; midlobe ovate, somewhat constricted
below the base, with recurved margins, obtuse. Column
stout, about 2 mm long excluding the anther; foot about
2.5 mm long, anther about 1 mm in diameter.
Epiphytic in wet forests; the type locality is
1100 m. Flowering at least September; in culti-
vation at Selby Gardens, it flowers at various
times of the year.
Maxillaria strumata is related to the M. neglec-
ta group but has a much more cespitose plant hab-
it. It is little collected, probably because of its
small size, but has been collected by R. L. Dress-
ier at "low" elevation near San Joaquin de Dota.
Maxillaria suaveolens Barringer, Brittonia 37(1):
44-46. 1985. TYPE: Costa Rica, Puntarenas,
Esquinas forest, area between the Rio Esquinas
and Palmar, P. H. Allen 5323 (holotype: F; Is-
otypes: MO, NY, SEL). Figure 14C.
Plant known only from the type, epiphytic, somewhat
repent or apparently pendulous; shoots developing 4-8
cm apart. Roots to 1 mm in diameter. Pseudobulbs api-
cally 1 -foliate, somewhat compressed, ovoid, 3.5-5 cm
long, and perhaps 2 cm wide in live material, mostly
hidden when young by 2-4 subtending leaves. Leaves
subcoriaceous, oblong to narrowly obovate, shortly pet-
iolate; blade 10-17 cm long and 3-5 cm wide, retuse.
Inflorescences 1-few per rhizome bract axil of imma-
ture shoots, each 5-7 cm long, covered with about 5
inflated, acute bracts; ovary with pedicel enclosed in the
uppermost bract before pollination, 1.8-2.0 cm long.
Flowers fragrant, white with butter yellow lip. Sepals
elliptic-obovate, 4-5 cm long, 1.8(-2.3) cm wide, acute.
Petals elliptic-obovate, 3.7-4 cm long, 1.5-2 cm wide,
acute to obtuse. Lip hinged to the column foot, about
1.5-2 cm long, 3-lobate above the middle; callus on the
lower half ligulate; midlobe about 6 mm long, ovate.
Column arcuate, 1 cm long; foot about 5 mm long.
"Epiphytic in tops of large trees" at less than
50 m. Flowering September. Costa Rica; known
only from the type collection.
The large, fragrant (Allen compared its fra-
grance to that of narcissus), and attractive flower,
which should be conspicuous to collectors, sug-
gest that this is a rare species, but the plant is
probably a crown epiphyte where orchids are un-
seen except in felled forests or branch falls. Also,
the plant may rarely be in flower and thus over-
looked by collectors if the flowers last only 3
days, as indicated on the herbarium label. More
field collections, including liquid-preserved flow-
ers and observations on growth habit, are badly
needed. Floral dimensions are taken from one re-
hydrated flower and from the original description.
Maxillaria tenuifolia Lindl., Edwards's Bot. Reg.
23: sub. t. 1986. 1837. TYPE: Mexico, Vera-
cruz, Hartweg. ?Maxillaria gracilifolia
Kraenzl., Mitt. Inst. Allg. Bot. Hamburg 6:420.
1927. Figure 10D.
Plant a densely clumping epiphyte to about 75 cm
across, comprised of numerous decumbent to pendent
shoots 1-8 cm apart. Roots white, to 1.5 mm in diameter.
Pseudobulbs cylindric to fusiform, somewhat com-
pressed, 2-5 cm long, 1-2 cm wide, subtended by bla-
deless rhizome bracts, or if with blades these falling at
maturity, the apex 1 -foliate. Leaves linear, 15-50 cm
long, 3-8 mm wide. Inflorescence 1 per rhizome bract
of mature shoots, 1-2 cm long, covered by acute sheaths
and subtending rhizome bract; ovary with pedicel 3-4
cm long, much longer than the subtending bract. Flow-
ers variously blood red, purple-red, or yellowish spotted
with red, the lip yellow or white spotted with red, fra-
grant, similar to that of coconut (crayons?). Sepals lan-
ceolate-ovate, 2-3 cm long, 6-9 mm wide. Petals ellip-
tic-lanceolate, somewhat falcate, apical margins erose,
1.9-2.7 cm long, 5-8 mm wide. Lip simple, elliptic,
constricted on the apical V4, forming a reflexed apex;
callus on the lower V§ or just above, oblong, thickened
and rounded in front, thickened on the sides. Column
arcuate, 1-1.3 cm long; foot 2 mm long. Capsule ellip-
soid, 3-4 cm long.
Large clumping to distinctly rhizomatous epi-
phyte of larger trunks and branches in (semi-)ev-
ergreen forests at 100-1200 m. Flowering March
to May. Mexico (Campeche, Chiapas, San Luis
Potosi, Yucatan, and Veracruz) to Costa Rica.
Maxillaria tenuifolia is replaced geographically
in central Costa Rica by M. sanguinea, which
flowers in January and February, lacks the strong
fragrance of M. tenuifolia, and usually has nar-
rower leaves.
Maxillaria tigrina C. Schweinf., Amer. Orch.
Soc. Bull. 37:409-410. 1968. TYPE: Costa
Rica, Province of Heredia, R. T. Northen s.n.
(holotype: AMES). Chrysocycnis tigrinum (C.
Schweinf.) Atwood, Icon. PL Trop. 14: t. 1306.
1989. Figure 11B.
Plant a straggly, partially pendent epiphyte; shoots 4-
7 cm apart. Roots small, white, to 0.5 mm in diameter.
Pseudobulbs cylindric, subtended by non-blade-bearing
sheaths, apically 1 -foliate, 2-3 cm long. Leaves coria-
78
FIELDIANA: BOTANY
ceous, elliptic, the petiole 1-2.5 cm long; blade 5-10 cm
long, 1-2 cm wide, sulcate along the midvein and keeled
beneath. Inflorescence 1 per rhizome bract axil, 1.5-2
cm long, borne on the rhizome; ovary with pedicel 1.5
cm long, slightly shorter than the subtending, inflated,
acute floral bract. Flowers leathery, translucent brown-
ish, lined with purple. Sepals similar; dorsal lance-ellip-
tic, concave, 2.6 cm long, 8 mm wide, acute; lateral
sepals lance-ovate, reflexed near the base, cuspidate api-
cally, 2.4 cm long, 1 cm wide, acute. Petals elliptic-
ovate, 2.4 cm long, 9 mm wide, acute. Lip 3-lobate,
glabrous, 1.8 cm long; lateral lobes acute, spreading;
midlobe triangular, twice as long as the base; callus
transverse, ridgelike, terminating between the lateral
lobes. Column arcuate, to 8 mm long; pollinia 4, with
squarrose stipe and saddle-shaped viscidium.
Epiphytic in montane cloud forests at 1900-
2000 m. Flowering July. Costa Rica and Panama.
The arcuate column, unique 3-lobate lip, and
plant habit, as well as general floral shape, are
features more consistent with South American
Chrysocycnis. The lip is glabrous, the lateral lobes
more acute, and the column shorter than in the
South American species, but the plant habit as
well as major floral features are those of Chry-
socycnis. Cladistic analyses based on DNA stud-
ies are needed to resolve this question.
Maxillaria tonduzii (Schltr.) Ames & Correll,
Bot. Mus. Leafl. 11:17. 1943. Camaridium cos-
taricense Schltr., Repert. Spec. Nov. Regni Veg.
3:250. 1907. TYPE: Costa Rica, La Palma,
Tonduz 12429 (isotypes: us 577589, 815052,
815053; drawing of type: AMES). Camaridium
tonduzii Schltr., Repert. Spec. Nov. Regni Veg.
8:571. 1910. TYPE: Costa Rica, Tonduz 12429
(same collection as for Camaridium costari-
cense Schltr.). Maxillaria heidelbergensis Sen-
ghas, nom. illeg. Orchidaceen (ed. 3) 29:1751.
1994. Figure 4C.
Plant an erect or decumbent, apparently monopodial
epiphyte to about 2 m tall lacking pseudobulbs; stems
occasionally branched, sometimes fasciculately so, with
each branch arising nearly parallel with the main stem,
concealed by the somewhat rugose leaf sheaths. Roots
white, to about 1 mm in diameter. Leaf blades above
the articulation ligulate, 2-13 cm long, 5-15 mm wide;
apex rounded to unequally 2-lobate. Inflorescence 1-3
per leaf axil, shorter than the subtending leaf; scapes to
about 5 cm long; ovary with pedicel (7)10-1 1 mm long,
subtended by a hyaline, acute floral bract 1.3-1.7 cm
long. Flowers brick red or green stained with reddish
brown; lip white with purple spots. Sepals somewhat
canaliculate, lanceolate, 1.7-2.3(3.7?) cm long, (3.5)4-
5 mm wide, acute; lateral sepals somewhat oblique. Pet-
als linear-lanceolate, 1.4-1.9 cm long, 2.5-3.5 mm wide,
acute. Lip hinged to the column foot, arcuate in natural
position, 3-lobate above the middle, 7-9.5 mm long,
6.5-9 mm wide when spread; lateral lobes somewhat
clasping the column, the apex forming nearly a right
angle; midlobe somewhat flared, rounded in front; callus
ligulate from near the base to just above the middle,
elevated in front, with several sharp teeth at the base
that in some specimens seem to form more or less into
3 rows. Column stout, somewhat arcuate, 5-6 mm long
excluding the anther; foot 3 mm long; anther 1 mm long,
verrucose.
Epiphytic and terrestrial in montane rain forests
at 1300-1650 m. Flowering season incompletely
known; at least August, October, February, and
March. Apparently endemic to Costa Rica, al-
though similar plants have been collected in ad-
jacent Panama with more attenuate sepals and pet-
als.
Maxillaria tonduzii is unique for its tall canes
and brick red flowers with purple-spotted, white
lip.
Maxillaria trilobata Ames & C. Schweinf., Bot.
Mus. Leafl. Harvard Univ. 11:17. 1943. Ca-
maridium brenesii Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:237. 1923. TYPE: Costa
Rica, San Ramon, Brenes 233 (photo of type:
AMES). Figure 3D.
Plant a large epiphyte with apparently monopodial
canes (adults) to 1 m long. Roots white, to about 1 mm
in diameter. Pseudobulbs lacking in adults but possibly
present in unknown juveniles. Leaves supported on
elongate, tapering sheaths to about 15 cm long including
the petiole above the oblique articulation; blade subcor-
iaceous, 10-25 cm long, 4-6 cm wide; apex rounded to
acute. Inflorescences several per leaf axil; scapes about
8 cm long, covered by membranaceous, acute bracts;
ovary with pedicel 7-10 mm long, exceeded by the sub-
tending inflated bract. Flowers small, white with yellow
and red on lip. Sepals oblong-ovate, 1.2-1.8 cm long,
5-7 mm wide. Petals elliptic-oblanceolate, 1-1.5 cm
long, to 6 mm wide, acute. Lip articulated to the column
foot, deeply 3-lobate on the apical VS, 9-10 mm long,
9-10 mm wide; callus ligulate on the lower Vi. Column
cylindric, 3-5 mm long, extended at the base into a foot
1 mm long. Capsule obovoid, about 2-2.5 cm long.
A large epiphyte in cloud forests at 1000-1700
m. Flowering November to May. Costa Rica and
Panama.
Maxillaria trilobata is vegetatively similar to
M. inaudita Rchb. f. in the long, petiolate leaves
but has much smaller flowers with a deeply 3-
lobate lip. The drawing in the Ames herbarium of
the type showing the long, tapering petioles and
small flowers with a deeply 3-lobate lip leaves
little doubt as to the identity.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
79
Maxillaria tubercularis J. T. Atwood, Lindley-
ana 9:229-231. 1994. TYPE: Panama, Chiriqui,
Fortuna Dam region, along trail to Cerro Hor-
nito, McPherson 13567 (holotype: MO). Figure
8A.
Plant a conspicuously rhizomatous epiphyte to about
50 cm tall; stems erect when young, pendent when large,
rooted at the base, concealed by tuberculate, nonfolia-
ceous bracts. Roots to about 1 mm in diameter. Pseudo-
bulbs ovoid, compressed, 2.5-5 cm long, 1-2.25 cm
wide, 3-10 cm apart, concealed when young by 2-3
tuberculate, foliaceous sheaths, the apex 1 -foliate.
Leaves with petiole '/6-% the blade length, the lateral
width 3-4 mm; blade coriaceous, narrowly elliptic, 6-
15 cm long, 1-3 cm wide; apex acute. Inflorescences
apparently restricted to nonfoliaceous bract axils of the
mature or immature rhizome; scapes less than 2 cm long;
ovary with pedicel 1.3-1.6 cm long, subtended by a sub-
equal, inflated, acuminate floral bract. Flowers well ex-
panded, yellow with many red to maroon spots; lip red
or maroon. Sepals dissimilar, ovate, acute; dorsal some-
what concave, 1.9-2.2 cm long, 8.5-14 mm wide; lateral
sepals attached to the column foot, 1.8-2.2 cm long, 1.2
cm wide. Petals elliptic-ovate, somewhat oblique, 1.5-
1.7 cm long, 6-7 mm wide, acute to obtuse. Lip not
firmly attached to the column foot, ovate in general out-
line when spread, 1.2-1.3 cm long, 9-10 mm wide, 3-
lobate at about the middle; lateral lobes variable, acute
to rounded, somewhat exceeded in front by the fleshy,
subquadrate, emarginate callus; midlobe ovate, rounded,
with crenulate margin. Column arcuate, about 9 mm
long excluding foot and anther, foot 5-6 mm long. Cap-
sule ellipsoid, 3-alate.
Epiphytic in evergreen (often oak) forests at
1300-2700 m. Flowering August, December, and
January. Endemic to southern Costa Rica and
Chiriqui, Panama.
Maxillaria tubercularis most closely resembles
M. elatior, found to the north, but is generally
smaller in all vegetative parts and occurs at much
higher elevations.
Maxillaria umbratilis L. O. Williams, Ann. Mis-
souri Bot. Gard. 28:425. 1941. Camaridium nu-
tantiflorum Schltr., Beih. Bot. Centralbl. 36,
Abt. 2:417. 1918. TYPE: Costa Rica, La Palma,
C. Werckle (isotype: CR). Camaridium vinosum
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
240. 1923. TYPE: Costa Rica, San Ramon, A.
M. Brenes 234 (lectotype: CR; isolectotypes:
AMES, NY). Maxillaria vinosa (Schltr.) Senghas,
Orchidaceen (ed. 3) 29:1751. 1994. Not Max-
illaria nutantiflora Schltr. Not Camaridium vi-
nosum Rolfe. Figure 15 A.
Plant a subshrub to about 1 m tall, of 1 or more
canelike shoots, each 10-30 cm long, terminating in a
pseudobulb and forming at the base of the previously
developed pseudobulb. Roots white, to 2 mm in diam-
eter. Pseudobulbs ovoid, biconvex, 3.5-5 cm long, 1(2)-
foliate, with 2-5 blade-forming rhizome bracts at the
base. Leaves coriaceous, petiolate including the rhizome
bracts (early deciduous), the blade narrowly elliptic-lan-
ceolate, to about 20 cm long, 2 cm wide, the apex obtuse
and often retuse. Inflorescence 1 per axil of the half-
lyre-shaped rhizome bract of the developing shoot;
scapes ^4 cm long, covered by acute sheaths; ovary
with pedicel 10-13 mm long, subtended and exceeded
by a cucullate floral bract 15-21 mm long. Flowers var-
iously colored, yellowish green, greenish white to or-
ange, suffused with red or entirely red, the lip often with
red. Sepals similar, lanceolate, concave, 19-21 mm
long, 4.5-5 mm wide, acute. Petals elliptic, attenuate
toward both base and apex, 16-20 mm long, 5-6 mm
wide. Lip hinged to the column foot, deeply 3-lobate,
10 mm long, 10 mm wide; lateral lobes spreading, fal-
cate; midlobe triangular, acute, 5-6 mm long, with a
fleshy V-shaped callus on the lower V4. Column stout,
arcuate, 4-5 mm long; foot at the base 2.5-3 mm long.
Capsule ellipsoid, about 2.5 cm long.
Epiphyte of wet forests or terrestrial on mossy
embankments, especially road cuts at 1000-2800
m. Flowering January to April, July to October,
and perhaps throughout the year. Costa Rica and
Panama.
Maxillaria umbratilis is unique among species
that produce half-lyre-shaped rhizome bracts in
the flush of new growth by the smaller, rather
dull-colored flowers with narrow floral segments.
The basis for the understanding of Camaridium
nutantiflorum is the original description and the
floral diagnosis by Schlechter (Repert. Spec. Nov.
Regni Veg. Beih. 59: t. 68 No. 271. 1931), which
shows the callus rather poorly rehydrated. Actu-
ally, the callus is more V-shaped in front than
shown by his drawing, but when pressed the tips
of the V tend to point forward. A rehydrated flow-
er from Brenes 234 shows the same type of callus.
Schlechter recognized C. vinosum on the basis of
the smaller size and the odd and probably abnor-
mal inflexed lip apex, but there seems to be a
continuum between the types. The dark flower
color indicated by Brenes is probably based on
either aged flowers, a color form, or perhaps
plants in an unusually exposed location.
Maxillaria uncut a Lindl., Ed wards 's Bot. Reg.
23: sub t. 1986. 1837. TYPE: Guiana, Loddiges
s.n. (holotype: K). Camaridium uncatum
(Lindl.) Hoehne, Arq. Bot. Est. Sao Paulo 6:
127. 1952. Maxillaria macleei Bateman ex
Lindl., Edwards's Bot. Reg. 26: misc. 70. 1840.
TYPE: Guatemala. Maxillaria nana J. D.
Hook., Ic. PL t. 315. 1841. TYPE: Guiana, C.
80
FffiLDIANA: BOTANY
S. Parker, Maxillaria squamata Barb. Rodr.,
Gen. & Sp. Orch. Nov. 1:118. 1877. TYPE:
Brazil (illustration in Flora Brasiliensis 3(6): t.
24). Ornithidium squamatum (Barb. Rodr.)
Barb. Rodr., Gen. & Sp. Orch. Nov. 2:209.
1882. Camaridium squamatum (Barb. Rodr.)
Hoehne, Arq. Bot. Est. Sao Paulo 2:72. 1947.
Maxillaria stenostele Schltr., Beih. Bot. Cen-
tralbl. 36, Abt. 2:414. 1918. TYPE: Costa Rica,
Rio Sucio, F. C. Lehmann 1236 (holotype: B,
destroyed). Maxillaria striatella Kraenzl., Re-
pert. Spec. Nov. Regni Veg. 24:359. 1928.
TYPE: Guatemala. Figure IOC.
Plant variable in habit, usually repent with shoots 1
or more cm apart or cespitose with shoots approximate,
or appearing cespitose with many ascending shoots each
spaced 1 or more cm apart. Roots to 1 mm in diameter.
Pseudobulbs cylindric, apically 1 -foliate, about 1 cm
long, concealed by scaly rhizome bracts. Leaves vari-
able, even in the same shoot, always fleshy, flat to terete,
narrowly elliptic to oblong or linear, acute, 1.5-6 (or
more) cm long, 2-6 mm wide. Inflorescence 1 per rhi-
zome bract axil; scapes to 1 cm long, concealed by
bracts; ovary with pedicel 14 mm long, much exceeding
the subtending floral bract. Flowers light lavender to
waxy white, lined with purple. Sepals dissimilar; dorsal
ovate, concave, 9 mm long, 4 mm wide, acute; lateral
sepals triangular, formed around the long column foot,
1 .5 cm long from the tip of the foot to the apex, 5 mm
wide, acute. Petals obliquely ovate, 8-9 mm long, 4 mm
wide. Lip ligulate, canaliculate on the lower %, con-
stricted on the apical 1A, with a ligulate callus thickened
apically and appearing parabolic in front, reaching be-
yond the middle. Column arcuate, 1.4 cm long includ-
ing the 8-mm-long foot, broadened and deepened above
to accommodate the large pollinarium with stipe 2.5-3
mm long appearing hooklike when viewed laterally.
Capsule ellipsoid, about 1 cm long.
Epiphytic on the larger branches of trees in wet
forests, mostly at low elevations but to 1500 m.
Flowering throughout the year. Locally common
in Central America, but in Mexico known only
from Chiapas; probably widespread in South
America.
The fleshy, dark green leaves and waxy, purple-
nerved flowers with a very long column foot and
long stipe are good features for recognition. Flow-
ers of Central American forms are darker and
have a longer callus than do most forms from
South America, and possibly there are two or
more species here accepted as one. Some plants
from northern South America have creamy white
flowers. The type of M. uncata shows the shorter
callus of South American forms.
Maxillaria vaginalis Rchb. f., Beitr. Orchid. -
K.C.Amer. 77. 1866. TYPE: Costa Rica, De-
sengano, Wendland 1247 (holotype: w). Ca-
maridium \vercklei Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:58-59. 1923. TYPE: Costa
Rica, San Cristobal, Werckle 5 (Drawing of
type: AMES). Figure 13F.
Plant a somewhat straggly, erect to decumbent, cane-
forming epiphyte or terrestrial to about 1 m long; stems
anchored at the base, concealed by bracts, formed from
successively borne shoots each terminated by a pseudo-
bulb and diminishing in size from the base. Roots to 1
mm in diameter. Pseudobulbs ovoid, compressed, 1-3.5
cm long, concealed by imbricate bracts and sheaths of
the subtending 1-3 leaves; apex 1 -foliate. Leaves at
base of pseudobulb with conspicuous sheaths constricted
at the articulation, apical leaf conspicuously petiolate;
blades similar, keeled below, narrowly elliptic, 7-30 cm
long, 1.5-6 cm wide, acute. Inflorescence 1 per non-
foliaceous rhizome bract axil of developing shoots, not
within the axils of developing leaves; scapes to about 4
cm long; ovary with pedicel 1-1.2 cm long, exceeded
by the subtending acute floral bract to about 2.5 cm long.
Flowers delicate in texture, white to pink, lip white,
column anther yellowish. Sepals similar, oblong, acute,
3-4 cm long, about 1 cm wide; dorsal somewhat con-
cave; lateral sepals oblique and recurved at about the
middle. Petals falcate-oblanceolate, 3.5 cm long, to 1
cm wide, acute. Lip firmly attached to the column foot,
canaliculate from the base, to about 2.4 cm long, 7 mm
wide in natural position, apical 1A distinctly 3-lobate; lat-
eral lobes somewhat spreading, rounded; midlobe ovate,
obtuse; callus of 5 fleshy ridges within at the base and
with a single fleshy, ligulate callus thickened apically
and extending from below the middle to beyond the si-
nuses between lateral lobes. Column slender, arcuate,
1.4-1.7 cm long excluding the foot and anther; foot
about 5 mm long. Capsule broadly ellipsoid, about 2
cm long with persistent column.
Epiphytic and terrestrial in wet montane forests
at 1500-2600 m. Available data on collections
suggest this may prefer habitats with high precip-
itation but little wind. Flowering July to October
and February. Costa Rica to Panama.
Maxillaria vaginalis is one of the larger-flow-
ered species of the M. sigmoidea complex. It is
easily recognized by the rather large white to pink
flowers on plants with pseudobulbs on the canes
and a straight, not sigmoid, lip.
Maxillaria valenzuelana (A. Rich.) Nash, Bull.
Torrey Bot. Club 34:121. 1907. Pleurothallis
valenzuelana A. Rich., Sagra, Hist. Fis. Cuba
11:234. 1850. TYPE: Cuba, Valenzuela, Wright
3314 (holotype: p?). Marsupiaria valenzuelana
(A. Rich.) Garay, Arq. Jard. Bot. Rio de Janeiro
12:183. 1952. Dicrypta iridifolia Bateman ex
Rchb. f., Bonplandia 2:16. 1854. TYPE: Cuba,
Poeppig (holotype: w). Maxillaria iridifolia
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
81
(Bateman ex Rchb. f.) Rchb. f., Bonplandia 2:
16. 1854. Dicrypta irisphyta Barb. Rodr., Gen.
et Sp. Orch. Nov. 1:126. 1877. TYPE: Brazil,
Minas Gerais (Martius, C. F. P. von, et al. 1896.
Flora brasiliensis 3(6): t. 17, fig. 2). Marsupi-
aria iridifolia Hoehne, Arq. Bot. Est. Sao Paulo
n.s. 2, pt. 4:71. 1947. Figure 18 A.
Plant a pendent, psygmoid, glaucous, essentially
monopodial epiphyte with very abbreviated nonswollen
stems but lacking pseudobulbs. Roots grayish, to 2 mm
in diameter. Mature leaves 5-10, equitant, remaining
unexpanded, linear to lanceolate in profile, somewhat
falcate, acute, 8-60 cm long, 1.4-2.7 cm wide measured
from margin to midvein, articulation oblique. Inflores-
cence a fascicle of successively borne, single-flowered
scapes; each scape to about 2 cm long, hidden by the
subtending rhizome bract and covered by acute scape
bracts; ovary with pedicel 2-2.5 cm long, subtended by
a rudimentary, short, obtuse floral bract. Flowers pre-
sented downward with the pendent shoot, greenish yel-
low, the lip deeper yellow with spots of red or purple.
Sepals narrowly elliptic to broadly lanceolate, 12-18
mm long, 3-5 mm wide, acute. Petals elliptic, 10-16
mm long, about 3 mm wide, acuminate. Lip fleshy,
rhombic to obovate and slightly 3-lobate above the mid-
dle, 10-15 mm long, 5-7 mm wide; midlobe broadly
acute, fleshy, 4-5 mm long. Column about 5 mm long,
forming a short foot at the base. Capsule ellipsoid, 2.5
cm long.
A pendent epiphyte of trunks and larger
branches in premontane wet forests at 1000-1500
m. Flowering throughout the year. Eastern Cuba
(type), Nicaragua, Costa Rica and adjacent west-
ern Panama, Colombia, Venezuela, and Brazil.
Maxillaria valenzuelana varies in plant and
flower size over the range. Specimens seen from
eastern Cuba have leaves that are broad at the
base and somewhat abruptly tapering as in Cos-
ta Rican material. All material from Nicaragua
has noticeably narrower leaves. Floral size of
specimens from Costa Rica and Panama is about
50% larger than in those from Nicaragua and
Honduras, a feature consistent with the larger
leaves.
Maxillaria valerioi Ames & C. Schweinf., Sched.
Orch. 10:96-97. 1930. TYPE: Costa Rica,
Province of Cartago, vicinity of Pejivalle,
Standley & Valeria 46940 (holotype: AMES; iso-
type: us). Figure 2D.
Plant a shrubby epiphyte on large branches; rhi-
zomes forming mostly upright canes sometimes with a
few branches, entirely concealed by sheaths. Roots fi-
brous, white, mostly forming at the base of the plant
and infrequently on the canes, rarely to 1 mm in di-
ameter. Pseudobulbs absent. Leaves densely disti-
chous, the sheath orangish in dried material, with hy-
aline margins; blade coriaceous, short leaves elliptic,
long leaves lanceolate, 1-10 cm long, 5-10 mm wide,
the base clasping, the apex unequally 2-lobate. Inflo-
rescence 1 per leaf axil, mostly concealed by subulate
bracts; scapes 1-3 cm long; ovary and pedicel 3-5 mm
long, exceeded by the subtending floral bract. Flowers
campanulate, white aging yellowish. Sepals 8.5-1 1
mm long, 2-2.5 mm wide; dorsal elliptic-lanceolate,
acute; lateral falcate-lanceolate and acute to rounded.
Petals elliptic, 7.5-9 mm long, 2-3 mm wide, obtuse.
Lip pandurate in outline, with lateral lobes on the basal
Vi, obtuse to acute in front; midlobe rhombic to broadly
oblanceolate; callus fleshy, ligulate on the lower Vi.
Column and foot 5-6 mm long excluding the anther;
foot 1-1.5 mm long.
Epiphytic and locally common, sometimes ter-
restrial in premontane rain forests at 550-1300 m.
Flowering January to July. Costa Rica and west-
ern Panama.
Vegetatively, M. valerioi is extremely variable
in leaf size, shape, and density. It resembles mem-
bers of the M. dendrobioides complex, but the
white flowers and long, narrow, pandurate lip dis-
tinguish it.
Maxillaria variabilis Bateman ex Lindl., Ed-
wards's Bot. Reg. 23: sub. t. 1986. 1837. TYPE:
Mexico. Maxillaria curtipes Hook, f., Ic. PI. 4:
t. 384. 1841. TYPE: Guatemala /. Parkinson
(holotype: K). ?Maxillaria chiriquensis Schltr.
Repert. Spec. Nov. Regni Veg. Beih. 17:68.
1922. TYPE: Panama, Chiriqui, ca. 4000 ft,
Powell 125 (isotypes: AMES, us; drawing of
type: AMES). Figure 11C.
Plant a somewhat straggly epiphyte to about 25 cm
high. Stems elongate; pseudobulbs approximate to
about 4 cm apart. Roots to about 1 mm in diameter.
Pseudobulbs fusiform to cylindric, often glaucous,
subtended by nonfoliaceous sheaths, apically 1 -foliate,
1.5-4 cm long, to 1.25 cm wide. Leaves thin, usually
shortly petiolate, the blade narrowly elliptic to lanceo-
late, 5-14 cm long, 6-20 mm wide, the apex obtuse to
unequally 2-lobate. Inflorescence formed in the flush
of new growth, to about 2 cm long; ovary with pedicel
1.3-2 cm long. Flowers orange to yellow with red
stain, rarely dark red (Mexico), the lip yellow with
dark, shiny red center. Sepals similar, lanceolate to el-
liptic, 1-1.5 cm long, about 5 mm wide, acute to ob-
tuse; dorsal concave and more elliptic, the lateral more
lanceolate to falcate. Petals oblanceolate, 1-1.4 cm
long, 4-5 mm wide; apex acute and strongly recurved.
Lip simple to lightly 3-lobate, more or less straight
with the sides somewhat embracing the column, ellip-
tic-ovate, 1-1.3 cm long, 5-6.5 mm wide; apex trun-
cate to rounded or retuse; callus a fleshy and shiny
ligule on the lower Vi. Column arcuate, 6-9 mm long
excluding the anther; foot 1.5-2 mm long.
82
FIELDIANA: BOTANY
Plants of semideciduous forests at 600-1800 m.
Flowering November to April and August. Mex-
ico to Panama; possibly to Peru.
Maxillaria variabilis is most closely related to
M. caespitifica Rchb. f. and M. costaricensis.
Maxillaria caespitifica is a smaller plant with
smaller, greener flowers, while M. costaricensis
has green to cream red-stained flowers and occurs
in wetter habitats. Populations north of the Central
Valley have closer pseudobulbs than do those
from southern Costa Rica. Maxillaria chiriquen-
sis, here interpreted cautiously as a synonym, has
yellow flowers, but the plant appears more slen-
der, as in M. costaricensis. Other populations in
southern Costa Rica have much stouter shoots
with broader leaves than plants to the north. Var-
ious authors, including this one, have misinter-
preted M. curtipes as an earlier synonym of M.
houtteana Rchb. f., a species that doubtfully oc-
curs in Costa Rica.
Maxillaria vittariifolia L. O. Williams, Ceiba 4:
38. 1953. TYPE: Costa Rica, Puntarenas, El
General Valley C. H. Lankester 1581 (holotype:
us; isotypes: AMES, SEL). Maxillaria linearis L.
O. Williams, name. Not M. linearis C.
Schweinf. Figure 25B.
Plant densely cespitose, very small, to 7 cm tall.
Roots white, to 0.5 mm in diameter. Pseudobulbs or-
bicular to ovoid, 6-10 mm tall, 5-8 mm wide, subtended
by 1-2 leaf-bearing rhizome bracts, apex 1 -foliate.
Leaves linear, 20-55 mm long, 1 .5-2 mm wide, tapering
toward the base. Inflorescence 1 per rhizome bract axil,
slender, erect, equaling or exceeding the leaves, to 6 cm
long, with 2-3 remote and inconspicuous bracts plus 1
subtending the flower; ovary with pedicel shorter than
the subtending bract, about 2 mm long. Flowers white,
the lip with bright orange apex. Sepals elliptic-ovate, 5
mm long, 1.6-2 mm wide, obtuse. Petals elliptic-ob-
long, 4.5 mm long, 1.5 mm wide, obtuse. Lip broadly
obovate when spread, 3.5 mm long, 4 mm wide; apex
divided into 3 equal lobes; midlobe reflexed and side
lobes embracing the column, the central part with a pair
of calli. Column stout, to 2.5 mm long; anther minutely
papillose. Capsule globose.
Poorly known ecologically; epiphytic in pre-
montane and lower montane primary forests at
900 m. Flowering probably throughout the year.
Costa Rica, but to be expected in adjacent Pana-
ma.
This species is one of the smallest-flowered
species of Maxillaria in Costa Rica with odd fi-
liform peduncles and might better be classified in
its own genus.
Maxillaria wercklei (Schltr.) L. O. Williams,
Ann. Missouri Bot. Card. 27:284. 1940. Orni-
thidium wercklei Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:60. 1923. TYPE: Costa
Rica, San Pedro de San Ramon, Werckle 61
(not found), Brenes 112 (neotype selected here:
AMES 28776). Figure 13 A.
Plant a somewhat straggly epiphyte to 15 cm tall;
rhizome erect or decumbent, rooted mostly at the base.
Roots to 0.5 mm in diameter. Pseudobulbs about 4 cm
apart, cylindric to somewhat tapered, 1-1.5 cm long,
about 5 mm wide at the base; subtended when young
by 1-2 foliate sheaths; apex 1 -foliate. Leaves coriaceous
to fleshy, sessile, elliptic, 7-25 mm long, 5-10 mm wide,
with unequally 2-lobate apex. Inflorescence 1 per rhi-
zome bract axil of the developing shoot; scape less than
1 cm long; ovary with pedicel 3.5-4.5 mm long, ex-
ceeded by the subtending floral bract. Flowers translu-
cent tan with red veins or entirely red. Sepals similar,
elliptic-lanceolate, 6.5-9 mm long, 2-3 mm wide, acute;
laterals inserted on the column foot. Petals elliptic-lan-
ceolate, about 6-8 mm long, 2-3 mm wide, acute. Lip
hinged to the column foot, ovate in general outline, 4.5-
5 mm long, about 3 mm wide when spread, 3-lobate at
the base; lateral lobes short, not exceeding V4 the lip;
midlobe suborbicular, obtuse, the side margins much re-
curved; callus tonguelike exceeding and connecting the
lateral lobes. Column stout, 1.5-2 mm long excluding
the anther; foot about 1 mm long; anther about 1 mm in
diameter.
Maxillaria wercklei is a premontane rain forest
species at (300)500-1350 m. Flowering Novem-
ber to January and August. Apparently endemic
to Costa Rica, but to be expected in adjacent Pan-
ama. Plants identified as M. wercklei from Pana-
ma appear to be M. lankesteri Ames.
The understanding of M. wercklei is based on
the Brenes collection cited by Schlechter (Repert.
Spec. Nov. Regni Veg. Beih. 19:244. 1923). Nei-
ther the holotype nor any isotypes of M. wercklei
based on Werckle 61 have been found. Maxillaria
lankesteri is usually considered as a later syn-
onym, but that species is easily distinguished by
the much larger flowers.
Excluded Taxa
Maxillaria cobanensis Schltr., Repert. Spec. Nov.
Regni Veg. 10:295. 1912. This was reported by
L. O. Williams (1956) as occurring in Costa
Rica, but specimens have been seen only from
Mexico, possibly to Nicaragua. Despite the lack
of specimens, R. L. Dressier believes he may
have seen it in Costa Rica. Plants from Hon-
duras appear similar to a small, cespitose, rather
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
83
broad-leaved variety of M. variabilis with
brown-purple flowers.
Maxillaria houtteana Rchb. f., Hamb. Gartenz.
14:212. 1858. This species occurs from Mexico
possibly to Nicaragua. It was treated (by J. T.
A.) as a synonym of M. curtipes Hook. f. (Icon.
PI. Trop. 14: t. 1342) based on a collection from
Mexico in the belief that it probably occurs in
Costa Rica based on the checklist of L. O. Wil-
liams ( 1 956), but no specimens south of Nica-
ragua have been seen. Subsequent examination
of the type of M. curtipes, rather than photo-
graphs, revealed that M. curtipes is a synonym
of M. variabilis Batem. ex Lindl., as suggested
by M. Soto A. (pers. comm.).
Mormolyca Fenzl
(J. T. Atwood)
REFERENCES — L. A. Garay & M. Wirth, On the
genera Mormolyca Fenzl and Cyrtoglottis Schltr.
Canad. J. Bot. 37:479-490. 1959. L. van der Fiji
& C. H. Dodson, Orchid Flowers. University of
Miami Press, Coral Gables, Florida. 214 pp. 1966.
Epiphytic cespitose or rhizomatous and creeping
herb. Pseudobulbs ovoid or elongate, variously com-
pressed. Leaves 1-4 at the pseudobulb apex, none at the
base, conduplicate, chartaceous to coriaceous, linear to
elliptic-oblong. Scapes 1 -flowered, laterally produced at
the base of the pseudobulb, elongate, usually with the
internodes exposed between bracts; ovary distinctly ped-
icellate, at least in the Central American species. Flow-
ers ringent, usually with rather lurid colors. Sepals sim-
ilar or dissimilar, never connate. Petals similar or dis-
similar to the sepals. Lip simple or 3-lobate, often with
appearance of an insect. Column stout or slender; pol-
linia 4, usually without a stipe (stipe present in at least
1 South American species) but supported on a viscidium.
Fruit a capsule.
A Neotropical genus of about six species. The
primary distinguishing feature of Mormolyca is
the footless column, although one could argue that
a foot is actually present that does not extend be-
yond the receptacle. The flowers appear adapted
for pseudocopulation, although the author is un-
aware of any observations of natural pollination.
Only a single species is known in Central Amer-
ica (see note under Maxillaria piestopus Schltr.).
Garay and Wirth (1959) suggested that Mor-
molyca and related Trigonidium may prove insep-
arable from Maxillaria, a view also held by one
of us (J. T. A.). The heavily veined leaves, ces-
pitose pseudobulbs lacking foliaceous sheaths at
the base, obsolete column foot, usually sharp lat-
eral lobes of the lip, and distinctly pedicellate
ovaries strongly suggest a relationship with the
Maxillaria rufescens complex.
Mormolyca ringens (Lindl.) Schltr., Die Orchi-
deen 436. 1914. Trigonidium ringens Lindl.,
Edwards's Bot. Reg. 26: misc. 57. 1840. TYPE:
Mexico, Hartweg (K, photo seen). Mormolyca
lineolata Fenzl, Denkschr. Kaiserl. Akad. Wiss.,
Math.-Naturwiss. Kl. Figure 26A.
Plant a cespitose epiphyte to about 40 cm tall. Rhi-
zome short, roots to 2.5 mm in diameter. Pseudobulbs
ovoid, congested, smooth when turgid, rugose if at all
dehydrated, somewhat compressed, 2-4 cm long, 1.5—4
cm wide, subtended by fugaceous, nonfoliaceous bracts,
apex 1 -foliate. Leaves coriaceous and strongly veined
above, shortly petiolate, elliptic, acute, 9.5-44 cm long,
1.3-5.5 cm wide. Inflorescence 1 or more per shoot,
lateral, wiry, with 5-7 appressed bracts including the
floral bract, internodes conspicuously exposed; ovary
with pedicel 3-4.5 cm long, the pedicel 2.25-3.6 cm
long, subtended by a floral bract 7-12 mm long. Flow-
ers ringent, not very variable, with petals pointing up-
ward and parallel with the dorsal sepal, lateral sepals
pointing downward and parallel to each other, greenish
aging tan, veined with red; lip with maroon callus at
base. Sepals dissimilar; dorsal concave, elliptic acute to
shortly acuminate, 1.4-1.6 cm long, 7-8 mm wide; lat-
eral convex, elliptic-falcate, 1.5-1.7 cm long, 5.5-7 mm
wide, with apex bluntly acute and somewhat recurved.
Petals similar to the lateral sepals but smaller, somewhat
convex, elliptic-falcate, somewhat twisted, 1.3-1.5 cm
long, 4-5 mm wide, apex obtuse and acuminate. Lip
porrect and parallel with the column, somewhat velvety,
3-lobate below the middle, elliptic-rhombic in general
outline, 7-8 mm long when spread, 4-6 mm across the
lateral lobes; lateral lobes short, triangular, and acute:
midlobe recurved apically; calli 2, the first at the base
triangular, velvety, a second callus at the middle, some-
what elevated and horseshoe-shaped in front. Column
arcuate, clavate, essentially lacking a foot, 8 mm long
without anther, anther ca. 1 mm long; pollinia 4, with
horseshoe-shaped viscidium.
Epiphytic on larger branches of evergreen to
semideciduous forests; usually common locally in
the northern parts of its range. Flowering probably
throughout the year. Widespread from Mexico to
at least northern Costa Rica.
Mormolyca ringens is easily recognized
throughout its range by the unifoliate, clustered
pseudobulbs and wiry scapes supporting rather
gaping, greenish to tan flowers marked with red
and maroon. Even sterile specimens should be
easily distinguished if old scapes are present.
84
FIELDIANA: BOTANY
Trigonidium Lindl.
(J. T. Atwood)
Epiphytic or lithophytic, cespitose or rhizomatous and
creeping herb. Pseudobulbs ovoid or elongate, variously
compressed laterally. Leaves 1-5 at pseudobulb apex, none
at the base, conduplicate, chartaceous to coriaceous, linear
to elliptic-oblong. Scapes elongate, bracteate, 1 -flowered,
produced at the base of the mature pseudobulb or immature
shoot. Flowers campanulate, appearing triangular from the
front, usually more or less brown with petal apices metallic
blue to white. Sepals similar when spread. Petals much
smaller than sepals. Lip 3-lobate, shorter than lateral petals,
and with a simple callus. Column nearly straight, wingless,
forming at most a very short foot at the base; pollinia 4,
waxy. Fruit a capsule.
A perplexing Neotropical genus of perhaps 12
species differing mainly in floral size and growth
habit. Three species are known in Central Amer-
ica, all in Costa Rica.
The relationships of Trigonidium with the
rest of the Maxillariinae are unclear, but there
are vegetative and floral similarities with some
of the Brazilian species of Maxillaria (M. picta
group). Trigonidium itself is probably mono-
phyletic, held together by the similar and rather
odd floral apomorphies such as the campanulate
presentation of the perianth and shiny spot on
the petal apex.
Key to the Species of Trigonidium
la. Plants conspicuously rhizomatous, pseudobulbs bearing >2 leaves at the apex T. lankesteri
Ib. Plants cespitose, pseudobulbs apically 2-foliate 2
2a. Flowering mostly November to June; flowers arising from the axils of bracts of mature shoots;
ovary with pedicel 4.7-7 cm long; narrowest leaves at least 14 mm wide . . T. egertonianum
2b. Flowering mostly August to October; flowers arising from the axils of bracts of immature
shoots; ovary with pedicel 4-4.5 cm long; widest leaves 13 mm wide or less
T. riopalenquense
Trigonidium egertonianum Bateman ex Lindl.,
Edwards's Bot. Reg. 24: misc. p. 73. 1838.
TYPE: Honduras, G. U. Skinner (holotype: K,
photo seen). Trigonidium seemannii Rchb. f.,
Seem. Bot. Voy. Herald. 214. 1854. TYPE:
Panama, Seemann s.n. (holotype: w). Figure
26B.
Plant a cespitose epiphyte often forming massive
clumps. Roots velamentous, to 2.5 mm in diameter.
Pseudobulbs approximate, ovoid in outline, somewhat
laterally flattened, apically 2-foliate, 4.5-7 cm long and
1.7-3 cm wide, with several longitudinal grooves be-
coming more apparent on drying, invested basally by
several nonfoliaceous bracts. Leaves 2 at the pseudobulb
apex, narrowly oblanceolate, arcuate near the apex in
live material, acute, 30-48 cm long and 1.2-2.5 cm
wide. Inflorescence 1 -flowered, arising laterally at the
base of the most recently completed shoot on peduncles
25-45 cm long, covered by 8-10 closely appressed tu-
bular sheaths; ovaries with pedicels 4.7-7 cm long.
Flowers green to orange, faintly reticulated with purple
thus appearing brown, the petals provided with a metal-
lic blue spot near the apex. Sepals closely appressed at
the base but broadly expanded apically, quadrate to ob-
ovate, acuminate, 3-4 cm long and 1-2 cm wide. Petals
lanceolate, acute, shortly clawed, 1.5-2 cm long and 4-
6 mm wide. Lip 8-10 mm long with 3 lobes diverging
above the middle; lateral lobes turned upward around
the column and provided centrally with a ligulate callus;
apical lobe orbicular, verrucose, and reflexed. Column
terete, 4-6 mm long.
Epiphytic on larger branches of trees of ever-
green or less often semideciduous forests at 0-
1000 m; rarely lithophytic; usually common.
Flowering in November to June. Widespread from
Mexico into South America.
Trigonidium egertonianum resembles closely T.
riopalenquense Dodson, with which it occasion-
ally occurs on the same host. They are best dis-
tinguished by the length of the ovary with the
pedicel and by the habit of flowering either on
completed growth (T. egertonianum) or on im-
mature growth (T. riopalenquense). Leaf width is
helpful in determining sterile specimens, but a de-
pauperate plant of T. egertonianum could easily
be misdetermined as T. riopalenquense on this
feature alone.
Trigonidium seemannii Rchb. f. is here inter-
preted as a synonym of T. egertonianum. The ho-
lotype of T. seemannii (w) consists only of a
drawing of a scape and flower, but only T. eger-
tonianum is known from the type locality, low-
land central Panama. Furthermore, Reichenbach
intended the name T. seemannii to replace T.
egertonianum, a species with a description he be-
lieved to be defective. Described from Honduran
material, T. egertonianum is the only species of
Trigonidium known north of Nicaragua.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
85
Triton id in m lankesteri Ames, Sched. Orch. 5:
32-34. June 1923. TYPE: Costa Rica, Lankes-
ter 367 (holotype: AMES). Trigonidium ampa-
roanum Schltr, Repert Spec. Nov. Regni Veg.
Beih. 19:61. Nov. 1923. TYPE: Costa Rica,
Carillo, C. Werckle 26 (not found). Trigonidium
insigne of authors. Not Rchb. f. ex Benth. &
Hook. Figure 26D.
Plant conspicuously rhizomatous, the ascending
shoots more than 15 cm apart; roots to about 1.5 mm in
diameter. Pseudobulbs conical to cylindrical, to about 8
cm long, sheathed basally with obtuse sheaths about 2
cm long, apically 3-5 foliate. Leaves imbricate basally,
thin, coriaceous, elliptic-oblong to elliptic-lanceolate,
18-28 cm long and 2.5-4(5) cm wide, shortly petiolate.
Inflorescence a solitary flower borne on a scape con-
cealed by imbricating bracts and rising from the base of
the most recently formed shoot. Flowers large for the
genus, brown. Sepals 4.5-6 cm long and 1 .5-2 cm wide,
connivent basally, sharply reflexed above the middle,
acute apically. Petals elliptic-oblanceolate, about 2.5 cm
long, thickened apically. Lip 1.4 cm long and about 5
mm wide, 3-lobate apically, the lateral lobes erect, the
midlobe minutely glandular; callus bilobed. Column
about 1 cm long.
An uncommon and striking epiphyte of tropical
and premontane rain forests from Costa Rica to
South America to about 1000 m. Flowering Au-
gust and September.
The large flowers, ascending rhizomatous habit,
and apically 3-5 -foliate pseudobulbs readily dis-
tinguish this species from others. Trigonidium
lankesteri also occurs at higher elevations. The
name T. insigne Rchb. f. ex Benth. & Hook. f.
has been misapplied to South American plants
that appear to be T. lankesteri. The holotype of T.
insigne based on a Colombian specimen collected
by Purdie (K) is a large plant related to and prob-
ably conspecific with T. egertonianum Batem. ex
Lindl.
Trigonidium riopalenquense Dodson, Selbyana
2:56. 1977. TYPE: Ecuador, Los Rios, Dodson
5251 (holotype: SEL). Figure 26C.
Plant a cespitose epiphyte, sometimes shortly creep-
ing, rarely forming large clumps. Roots velamentous, to
2 mm in diameter. Pseudobulbs approximate, even
when exhibiting a creeping habit, ovoid, compressed,
rarely conical, with furrows formed rather irregularly on
drying, 2-3.5 cm long and 1.7-2.3 cm wide, 2-foliate,
sheathed basally by 2 or more bracts disintegrating with
age. Leaves linear to very narrowly oblanceolate, acute,
30-40 cm long and 0.6-1.2 cm wide. Inflorescences
lateral, 1 -flowered, arising from the axil of a bract pro-
tecting the immature shoot, supported on a peduncle 17-
25 cm long covered by 7-9 closely appressed bracts;
ovary with pedicel 4-4.5 cm long. Flowers greenish to
pinkish yellow to brown. Sepals rhombic, obovate, acu-
minate, 2.8-3.4 cm long and 1-1.5 cm wide. Petals with
a metallic blue "eye" apically, lanceolate, acute, 1 .5 cm
long and 5-6 mm wide. Lip 6-8 mm long, with 3 lobes
diverging above the middle, the lateral lobes turned up-
ward around the column, the apical lobe orbicular, ver-
rucose, and reflexed, with a ligulate callus on the lower
half. Column terete, 4-5 mm long. Capsule ellipsoid,
about 3 cm long.
Uncommon epiphytes of lowland humid tropics
from Nicaragua to Ecuador but easily confused
with T. egertonianum (see notes under that spe-
cies). Flowering in Costa Rica from August to Oc-
tober.
Subtribe Oncidiinae Benth.
REFERENCES — M. W. Chase, A reappraisal of
the Oncidioid orchids. Syst. Bot. 11:477-491.
1986. M. W. Chase & J. D. Palmer, Floral mor-
phology and chromosome number in subtribe On-
cidiinae (Orchidaceae): Evolutionary insights
from a phylogenetic analysis of chloroplast DNA
restriction site variation. In P. S. Soltis, et al., Mo-
lecular Systematics of Plants. Chapman and Hall,
New York, New York, pp. 324-337. 1992. R. L.
Dressier, The Orchids. Harvard University Press,
Cambridge, Massachusetts, and London, England,
332 pp. 1981. R. L. Dressier, Phylogeny and Clas-
sification of the Orchid Family. Dioscorides Press,
Portland, Oregon, 314 pp. 1993. L. O. Williams,
An enumeration of the Orchidaceae of Central
America, British Honduras and Panama. Ceiba 5:
1-256. 1956.
Plant usually an epiphytic herb, rarely terrestrial or
more commonly opportunistically terrestrial. Rhizomes
short or variously elongate; roots with white velamen,
variously thickened. Pseudobulbs usually present, of a
single internode, sheathed at the base with articulate fo-
liaceous bracts or by nonfoliaceous and nonarticulate
bracts; apex of 1 -several leaves. Leaves thin to fleshy,
conduplicate and flat, less often equitant or terete, artic-
ulate or less often nonarticulate, linear to elliptic, peti-
olate to nonpetiolate. Inflorescence a raceme, panicle
with various patterns of branching, or less often a scape
with solitary flower; ovary usually with long pedicel.
Flowers usually complex in form and color pattern, col-
or various but perianth segments often marked with ma-
roon, red, or brown. Sepals similar or dissimilar, often
concave, often unguiculate, the laterals often variously
connate. Petals usually large and showy, sometimes
larger than the sepals, sometimes unguiculate. Lip sim-
ple to complexly lobed, with complex calli consisting of
various plates and tubercles. Column usually complex,
86
FIELDIANA: BOTANY
often with a plate ventrally near the base (tabula infra-
stigmatica), often complexly winged above; anther with
2 pollinia with stipe of varying length and a viscidium.
Fruit a capsule.
An exclusively Neotropical subtribe of about
77 genera and more than 1,200 species. Thirty-
five genera are known in Costa Rica with a total
of 112 species.
The Oncidiinae is best developed in South
America and grows in wet to nearly desert habi-
tats from sea level to more than 3500 m. Although
rather well defined as a subtribe, the genera within
it are particularly problematic, especially the ge-
nus Oncidium, which is defined on the basis of
plesiomorphic character states, including the ob-
tuse angle formed by the column and lip.
Key to Subtribe Oncidiinae
la. Leaves laterally flattened (equitant) 2
2a. Stems elongate; plant never fanlike 3
3a. Plants creeping monopodials 4
4a. Flowers white to green, segments < 5 mm Pachyphyllum
4b. Flowers red to purple, segments > 5 mm long Fernandezia (tica)
3b. Plants never creeping Lockhartia
2b. Stems not elongate, plant fanlike 5
5a. Sepals and petals narrowly lanceolate to attenuate, "spidery" Macroclinium
5b. Sepals and petals elliptic to ovate, never "spidery" 6
6a. Inflorescence a panicle of subumbels; floral segments < 3.5 mm long
Trizeuxis (falcata)
6b. Inflorescence never subumbellate; floral segments > 3.5 mm long . . Psygmor-
chis
Ib. Leaves not laterally flattened, either dorso- ventrally flattened or terete 7
7a. Plant monopodial, usually creeping 8
8a. Flowers red to purple, segments > 5 mm long Fernandezia (tica)
8b. Flowers white to green, segments < 5 mm Pachyphyllum
7b. Plant sympodial 9
9a. Spur, chin, or column foot present 10
lOa. Only 1 leaf per shoot 11
lla. Basal side margins of lip as well as petals embracing the column
Plectrophora (alata)
lib. Neither lip base nor petals embracing the column Trichocentrum
lOb. Leaves 2 or more per shoot 12
12a. Appendage present at base of lip; may be enclosed in sepaline spur ... 13
13a. Appendage of lip single, undivided Rodriguezia (compacta)
13b. Appendages of lip either 2 or 1 and furcate at apex 14
14a. Flower red; appendages of lip 2, about as long as blade of lip . .
Comparettia (falcata)
14b. Flower yellow; appendage of lip divided near apex, much shorter
than blade Scelochilus (aureus)
12b. Appendage absent at base of lip 15
15a. Column foot conspicuous, broad; flowers green; blade of lip much
shorter than sepals Systeloglossum
15b. Column foot inconspicuous; flowers either white or lavender, never
green; lip much longer than sepals lonopsis
9b. Spur, chin, or column foot absent 16
16a. Lip partially adnate to, or the basal margins embracing the column, sometimes
in-rolled and forming a tubular base 17
17a. Lip clearly 3-lobate with sharp sinuses between the lobes, the midlobe much
narrower than the lateral lobes 18
18a. Flowers white; lip and petals denticulate . . . Warmingia (margaritacea)
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
87
18b. Flowers colored, lip and petals entire . . . Macradenia [unknown but
expected in Costa Rica; reported from Nicaragua and Panama]
17b. Lip simple or 3-lobate but with blunt sinuses, or if sharp, then the midlobe
much broader than the lateral lobes 19
19a. Shoots with 1 apical leaf 20
20a. Sepals less than 2 cm long; leaf fleshy . . Leucohyle (subulata)
[unknown but expected in Costa Rica; reported from Nicaragua
and Panama]
20b. Sepals > 2 cm long; leaf thin Trichopilia
19b. Shoots with leaves at base of pseudobulb in addition to the 1-2 apical
leaves 21
2 la. Side lobes of lip not clasping nor in-rolled around the column;
pseudobulb apex with 2(1) leaves Aspasia
21b. Side lobes of lip either clasping or in-rolled around the column;
pseudobulb apex with 1 leaf Cischweinfia
16b. Lip entirely free from, although sometimes parallel with, the column when viewed
laterally 22
22a. Leaf 1 restricted to pseudobulb apex 23
23a. Sepals > 4 cm long 24
24a. Dorsal sepal linear; lateral sepals much broader, falcate; petals
similar to the dorsal sepal, antenna-like
Psychopsis (krameriana)
24b. Sepals all similar Rossioglosswn (schlieperianum)
23b. Sepals < 4 cm long 25
25 a. < 4 flowers per inflorescence Ticoglossum
25b. > 4 flowers per inflorescence Oncidium
22b. Leaves > 1 per shoot 26
26a. Petals broad, at least 1.3 cm broad, never yellow or olivaceous . . 27
27a. Flowers mostly chestnut brown .... Otoglossum (chiriquense)
27b. Flowers white to pink Miltoniopsis (warscewiczii)
26b. Petals narrower, < 1 .25 cm broad, of various colors 28
28a. Lateral sepals > 7.5 cm long, lanceolate Brassia
28b. Lateral sepals much shorter than 7.5 cm long 29
29a. Lip > 2.3 cm long 30
30a. Lip more or less acute Rhynchostele
30b. Lip obtuse, rounded, or emarginate Oncidium
29b. Lip < 2.2 cm long 31
3 la. Plant with combination of pseudobulb hidden by sev-
eral subtending leaves, starlike flowers with narrow
sepals, the lateral sepals > 1.9 cm long, lip less than
% the lateral sepals Ada (chlorops)
31b. Combination of features otherwise 32
32a. Flowers nonresupinate, white or tinged with lavender; the lip with yellow or red on the callus . .
Osmoglossum
32b. Rowers otherwise 32
33a. Angle formed by the column and base of lip ^ 30° when viewed laterally 34
34a. Lip simple, biblike or flabellate and emarginate, the blade flat, neither with recurved
nor incurved margins 35
35a. Lip deeply emarginate or with slightly concave side margins, white to deep lav-
ender lonopsis
35b. Lip if emarginate, then with convex side margins, greenish white to yellowish,
sometimes spotted Leochilus
34b. Lip 3-lobate, pandurate, or with recurved or incurved margins 36
36a. Lip 3-lobate, with subtriangular lateral lobes near the middle
88 FIELDIANA: BOTANY
Goniochilus (leochilinus)
36b. Lip otherwise 37
37a. Lip shorter than lateral sepals Mesospinidium
37b. Lip longer than lateral sepals Hybochilus (inconspicuus)
33b. Angle formed by column and lip base > 30° (often much greater than 60°) 38
38a. Column stout, less than twice as long as wide 39
39a. Lip simple, biblike Leochilus
39b. Lip 3-lobate Oncidium
38b. Column more than twice as long as wide 40
40a. Inflorescence a pendent, densely flowered raceme; lip simple, arrow-shaped ....
Notylia
40b. Inflorescence and lip otherwise 41
4 la. Column slender, arcuate, at least half as long as and usually more than %
the lip length 42
42a. Sepals < 1 cm long Sigmatostalix
42b. Sepals > 1 cm long Amparoa (costaricensis)
41b. Column not slender and arcuate, usually less than Vi the lip length .... 43
43a. Sepals > 7 mm long Oncidium
43b. Sepals < 6 mm long Sigmatostalix
Ada Lindl.
(D. E. Mora de Retana)
REFERENCES — L. A. Garay, Orquideas colom-
bianas nuevas o criticas. Orquideologia 4:14-21.
1969. N. H. Williams, A reconsideration of Ada
and the glumaceous brassias (Orchidaceae). Brit-
tonia 24:93-1 10. 1972.
Epiphytic, cespitose or rhizomatous and creeping
herb. Pseudobulbs present to obsolete, variously com-
pressed. Leaves several per shoot, 1-2 at pseudobulb
apex, chartaceous, conduplicate, with usually several
conspicuous veins. Inflorescence a lateral raceme of
several flowers, often 2 or more per shoot. Flowers stel-
late to campanulate with lanceolate to attenuate seg-
ments, the lip nearly parallel to the column; pollinia 2,
waxy, supported on an obdeltoid stipe and broadly ovate
viscidium. Fruit a capsule.
A genus of about 14 currently recognized,
mostly South American species; two species in
southern Central America, but only one in Costa
Rica. The genus is easily recognized by the re-
duced or absent pseudobulbs, the thin leaves with
several prominent veins, and the starry to cam-
panulate flowers borne on lateral racemes.
The DNA evidence of Chase and Palmer sup-
ports Williams' separation of Ada from closely
related Brassia. Unfortunately, Chase and Palmer
did not sample Mesospinidium, which is vegeta-
tively similar to Ada but usually has smaller and
fewer leaves.
Ada chlorops (Endres & Rchb. f.) N. Williams,
Brittonia 24:105. 1972. Brassia chlorops En-
dres & Rchb. f., Card. Chron. 542. 1873.
TYPE: Costa Rica, Endres (w). Brassia parvi-
flora Ames & C. Schweinf., Sched. Orch. 8:74-
75. 1925. TYPE: Costa Rica, Cascajal, Alfaro
s.n. (holotype: AMES) Figure 27A.
Plant epiphytic, cespitose, forming small clumps to
ca. 40 cm tall. Rhizome short; roots 2-3 mm in diameter.
Pseudobulbs reduced, ovate-lanceolate, compressed, 3-
5 cm long, 1.5 cm wide, covered at the base by the
conduplicate sheaths of several leaves, 1 -foliate apically.
Leaf blades articulate, carinate, subcoriaceous, elliptic-
lanceolate to obovate, acute, to 34 cm long, 1.3-2.5 cm
wide. Inflorescence a lateral, 5-10 flowered raceme,
20-35 cm long including the scape, the latter with ap-
pressed, acute, membranaceous bracts; ovary with ped-
icel 1.5-2.5 cm long, subtended by shorter to longer,
triangular, apiculate bracts. Flowers somewhat fleshy,
greenish yellow with brown spots. Sepals subequal, free,
ovate-lanceolate, acuminate; dorsal erect, 1.5-2.2 cm
long, 3-5 mm wide; lateral sepals subfalcate and spread-
ing, 1.9-2.5 cm long, 3-4 mm wide. Petals similar to
the sepals, 1.2-1.5 cm long, 2-3 mm wide. Lip oblong-
lanceolate, acute to acuminate, reflexed above the mid-
dle, the base adnate to the column, 1.2-1.4 cm long, 5-
6 mm wide; callus fleshy, sulcate and pubescent basally,
and with a pair of elevated teeth apically. Column 5-6
mm long, stout, foot lacking; anther terminal, rounded,
papillose, with 2 imperfect cells; pollinia 2, pyriform,
with subcordate stipe and viscidium. Fruit an ellipsoid
capsule ca. 5 cm long.
Epiphytic in premontane and lower montane
rain forests mostly at (200)1000-1600(2000) m,
where it grows on larger trunks of the host in
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
89
FIG. 27. A, Ada chlorops (Icon. PI. Trop. 16: t. 1501. Illustrator: F. Pupulin). B, Amparoa costaricensis (Icon. PI.
Trop. 7: t. 601. Illustrator: O. Achf). C, Aspasia epidendroides (Icon. PI. Trop. 16: t. 1502. Illustrator: C. Marfn). D,
Aspasia principissa (Icon. PI. Trop. 16: t. 1503. Illustrator: C. Marin).
90
FIELDIANA: BOTANY
FIG. 28. A, Brassia arcuigera (Icon. PI. Trop. 16: t. 1507. Illustrator: L. Alfaro). B, B. caudata (Icon. PI. Trop. 16:
t. 1508. Illustrator: L. Alfaro). C, B. gireoudiana (Icon. PI. Trop. 16: t. 1509. Illustrator: C. Marin). D, B. verrucosa
(Icon. PI. Trop. 16: t. 1510. Illustrator: C. Marin).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
91
3 mm
1 cm
2cm
FIG. 29. A, Cischweinfia dasyandra (Icon. PI. Trop. 15: t. 1413. Illustrator: L. Alfaro). B, Cischweinfia pusilla
(illustrator: O. Achf). C, Comparettia falcata (Icon. PI. Trop. 15: t. 1416. Illustrator: O. Achi). D, Fernandezia tica
(Brenesia 39-40:165. 1993. Illustrator: C. Marin).
92
FIELDIANA: BOTANY
1 cm
1 cm
2mml
5 mm
2 mm
FIG. 30. A, Goniochilus leochilinus (Icon. PI. Trop. 15: t. 1436. Illustrator: L. Alfaro). B, lonopsis utricularioides
(illustrator: O. Achf). C, Leochilus labiatus (illustrator: C. Marin). D, L. tricuspidatus (Icon. PI. Trop. 15: t. 1443.
Illustrator: L. Alfaro).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
93
2 mm
B
5 cm
2 mm
10 cm
5 mm
1 cm
FIG. 31. A, Lockhartia amoena (Icon. PI. Trop. 15: t. 1446. Illustrator: L. Alfaro). B, L. hercodonta (Icon. PI. Trop.
15: t. 1448. Illustrator: L. Alfaro). C, L. micrantha (Icon. PI. Trop. 15: t. 1449. Illustrator: L. Alfaro). D, L. oerstedii
(Icon. PI. Trop. 15: 1450. Illustrator: L. Alfaro).
94
FIELDIANA: BOTANY
1 mm
1 mm
FIG. 32. A, Lockhartia acuta (Icon. PI. Trop. 15: t. 1445. Illustrator: L. Alfaro). B, L. pittieri (illustrator: O. Achf).
C, Macroclinium confertum (Lindleyana 11:139. Illustrator: F. Pupulin). D, M. generalense (Lindleyana 11:137.
Illustrator: F Pupulin).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
95
5 mm
FIG. 33. A, Macroclinium cordesii (illustrator: E Pupulin). B, Macroclinium ramonense (illustrator: F. Pupulin). C,
Mesospinidium horichii (Icon. PI. Trop. 16: t. 1564, incorrectly as Mesospinidium endresii. Illustrator: L. Alfaro). D,
Mesospinidium warscewiczii (Icon. PI. Trop. 16: 1565. Illustrator: C. Marin).
96
FIELDIANA: BOTANY
FIG. 34. A, Miltoniopsis warscewiczii (illustrator: W. Zuchowski). B, Notylia trisepala (illustrator: O. Achi). C, N.
pittieri (illustrator: O. Achi). D, N. pittieri (illustrator: O. Achi).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
97
B
FIG. 35. A, Oncidium ascendens (illustrator: O. Achf). B, O. teres (illustrator: O. Achf). C, O. cebolleta (Icon. PI.
Trop. 15: t. 1467. Illustrator: L. Alfaro). Note: Plant habit is similar to that in Figures 35A, B. D, O. carthagenense
(Icon. PI. Trop. 16: t. 1570. Illustrator: L. Alfaro). E, O. schroederianum (Icon. PI. Trop. 15: t. 1472. Illustrator: L.
Alfaro).
98
FIELDIANA: BOTANY
1 cm
FIG. 36. A, Oncidium cariniferum (Icon. PI. Trop. 16: t. 1569. Illustrator: C. Marin). B, O. stenoglossum (illustrator:
O. Achf). C, O. panduriforme (Icon. PI. Trop. 15: t. 1471. Illustrator: L. Alfaro). D, O. storkii (Icon. PL Trop. 15: t.
1474. Illustrator: L. Alfaro).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
99
5 mm
FIG. 37. A, Oncidium globuliferum (illustrator: C. Marin). B, O. cristagalli (Icon. PI. Trap. 16: t. 1571. Illustrator:
F. Pupulin). C, O. warscewiczii (illustrator: O. Achi). D, O. bryolophotum (illustrator: S. Dalstrom).
100
FIELDIANA: BOTANY
6 mm 2 mm
1 cm
FIG. 38. A, Oncidium bracteatum (illustrator: O. Achf). B, O. luteum (Icon. PI. Trop. 16: t. 1572. Illustrator: S.
Dalstrom). C, O. cheirophorum (Icon. PI. Trop. 15: t. 1468. Illustrator: L. Alfaro). D, O. ampliatum (Icon. PI. Trop.
15: t. 1464. Illustrator: L. Alfaro).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
101
FIG. 39. A, Oncidium obryzatoides (illustrator: O. Achf). B, O. klotzschianum (Icon. PL Trap. 15: t. 1470 as synonym
O. obryzatum. Illustrator: L. Alfaro). C, O. dichromaticum (Illustrator: O. Achf). D, O. parviflorum (illustrator: L.
Alfaro).
102
FIELDIANA: BOTANY
1 cm
FIG. 40. A, Oncidium ansiferum (Icon. PI. Trop. 15: t. 1465. Illustrator: L. Alfaro). B, O. stenobulbon (illustrator: S.
Dalstrom). C, O. ensatum (illustrator: O. Achf). D, O. stenotis (Icon. PI. Trop. 15: t. 1473. Illustrator: L. Alfaro).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
103
FIG. 41. A, Oncidium polycladium (illustrator: O. Achf). B, Oncidium isthmi (illustrator: O. Achi). Note: Plant habit
is similar to that in Figure 41A. C, Osmoglossum convallarioides (Icon. PI. Trop. 15: t. 1475. Illustrator: L. Alfaro).
D, Osmoglossum egertonii (illustrator: C. Marin). E, Otoglossum chiriquense (Icon. PI. Trop. 16: t. 1573. Illustrator:
L. Alfaro).
104
FIELDIANA: BOTANY
FIG. 42. A, Pachyphyllum crystallinum (Icon. PI. Trap. 14: t. 1375 as P. pastii. Note: Floral diagnosis is based on a
forcefully opened bud. Illustrator: B. N. Culbertson). B, Pachyphyllum hispidulum (illustrator: S. Dalstrom). C,
Plectrophora alata (Icon. PI. Trop. 15: t. 1479. Illustrator: L. Alfaro). D, Psychopsis krameriana (Icon. PI. Trop. 15:
t. 1486. Illustrator: L. Alfaro).
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
105
2cm
B
FIG. 43. A, Psygmorchis pumilio (Icon. PI. Trop. 15: t. 1487. Illustrator: L. Alfaro). B, Rhvnchostele bictoniensis
(Icon. PI. Trop. 15: t. 1442. Illustrator: L. Alfaro). C, R. hortensiae (Icon. PI. Trop. 16: t. 1 54 1 .' Illustrator: F. Pupulin).
D, R. stellata (illustrator: C. Marin).
106
FIELDIANA: BOTANY
FIG. 44. A, Rodriguezia compacta (Icon. PI. Trop. 16: t. 1578. Illustrator: F. Pupulin). B, Rossioglossum schlieper-
ianum (Icon. PI. Trop. 16: t. 1579. Illustrator: F. Pupulin). C, Scelochilus aureus (Icon. PI. Trop. 16: t. 1585. Illustrator:
C. Marin). D, Sigmatostalix unguiculata (illustrator: O. Achi).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
107
4 mm
1 mm
FIG. 45. A, Sigmatostalix picta (illustrator: O. Achi). B, 5. macrobulbon (Icon. PI. Trop. 16: t. 1588. Illustrator: S.
Dalstrom). C, 5. brownii (illustrator: O. Achi). D, 5. hymenantha (illustrator: O. Achi).
108
FIELDIANA: BOTANY
1 mm
10cm
5 mm
1 cm
FIG. 46. A, Sigmatostallx adamsii (illustrator: O. Achf). B, Systeloglossum acuminatum (illustrator: S. Dalstrom). C,
Systeloglossum costaricense (Icon. PI. Trop. 15: t. 1494. Illustrator: L. Alfaro). D, Ticoglossum krameri (Icon. PI.
Trop. 16: t. 1596. Illustrator: F. Pupulin).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
109
FIG. 47. A, Ticoglossum oerstedii (Icon. PI. Trap. 14: t. 1398. Illustrator: B. N. Culbertson). B, Trichocentrum
dianthum (illustrator: F. Pupulin). C, Trichocentrum pfavii (illustrator: F. Pupulin). D, Trichocentrum estrellense
(illustrator: F. Pupulin).
110
FIELDIANA: BOTANY
B
FIG. 48. A, Trichocentrum caloceras (Icon. PI. Trop. 16: t. 1597. Illustrator: F. Pupulin). B, T. costaricense (illustrator:
F. Pupulin). C, T. brenesii (Selbyana 15:97. 1994. Illustrator: F. Pupulin, redrawn from type). D, T. capistratum (Icon.
PI. Trop. 16: t. 1598. Illustrator: F. Pupulin).
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
111
FIG. 49. A, Trichocentrum cymbiglossum (Selbyana 15: 100. 1994. Illustrator: L. Alfaro). B, Trichopilia maculata
(illustrator: O. Achf). C, Trichopilia suavis (Icon. PI. Trap. 15: t. 1498. Illustrator: L. Alfaro). D, Trichopilia turialbae
(Icon. PI. Trop. 16: t. 1600. Illustrator: C. Marin).
112
FIELDIANA: BOTANY
1 cm
FIG. 50. A, Trichopilia tortilis (Icon. PI. Trop. 15: t. 1499. Illustrator: L. Alfaro). B, Trichopilia marginata (Icon. PL
Trop. 15: t. 1497. Illustrator: L. Alfaro). C, Trichopilia galeottiana (Icon. PL Trop. 16: t. 1599. Illustrator: L. Alfaro).
D, Trizeuxis falcata (illustrator: O. Achi).
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
113
substantial shade. Flowering August to November.
Nicaragua to Panama.
Ada chlorops is easy to recognize in the field
by the stellate, green flowers spotted with tan on
fan-shaped plants with thin, almost plicate leaves.
Sterile plants appear similar to Miltoniopsis \var-
scewiczii (Rchb. f.) Garay & Dunsterv., with
which it often grows, but the latter has a much
more conspicuous pseudobulb, and its leaves are
more gray in color.
Amparoa Schltr.
(D. E. Mora de Retana)
Epiphytic, somewhat rhizomatous herb with rather
thick roots. Pseudobulbs ovate, 1 -foliate, compressed,
subtended by 2 or more foliaceous bracts. Leaf oblong-
lanceolate to somewhat elliptic, acute, thin in texture.
Inflorescence a lateral, scapose raceme borne at the base
of the pseudobulb, with 5-10 flowers; ovary with slen-
der pedicel. Flowers spreading, green or yellowish
green, the lip with yellow or orange. Sepals soft in tex-
ture, oblong-lanceolate, acute. Petals similar in shape
but notably smaller than the sepals, linear-oblanceolate,
porrect, acute. Lip somewhat fleshy, simple, straight,
subpandurate, with simple callus. Column slender, cy-
lindric to subclavate, without auricles; pollinia 2, with
stipe and viscidium. Fruit a capsule.
A genus of two species ranging from Mexico
to Costa Rica. One species occurs in Costa Rica.
Schlechter (Repert. Spec. Nov. Regni Veg.
Beih. 19:64. 1923) separated Amparoa from
Odontoglossum on the basis of the relatively small
petals, rather porrect and straight lip with simple
callus, and the slender column. Little is known of
its relationships, but the long column suggests a
relationship with Sigtnatostalix.
Amparoa costaricensis Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:65. 1923. TYPE:
Costa Rica, La Palma, C. Werckle 129 (AMES,
photo seen). Figure 27B.
Plant a cespitose to shortly creeping epiphyte 20-25
cm high; rhizome thick, to about 2 cm long between
pseudobulbs; roots to 2.5 mm thick. Pseudobulbs ellip-
tic-ovate, strongly compressed laterally and with slender
margins, 4-4.5 cm long, 2.5-2.7 cm wide, the apex 1-
foliate, subtended by ca. 4-6 soon deciduous, foliaceous
bracts. Leaves coriaceous, shortly petiolate; blades ellip-
tic-ovate, carinate, conduplicate at the base; apex acute
and apiculate. Inflorescence scapose, 20-35 cm long (to
40 cm long in Nicaragua), with 5-10 flowers; ovary with
pedicel 2-3 cm long subtended by much shorter acute
bracts. Flowers 2-4 open at a time, spreading, yellowish
green, the lip yellow with orange-yellow callus. Sepals
subequal, oblong-lanceolate, acute, carinate; dorsal 1.3-
1.4 cm long, 3-3.4 mm wide; laterals somewhat subfal-
cate, 1.4-1.6 cm long, 3-3.4 mm wide. Petals much
smaller than sepals, linear-oblanceolate, porrect, acute,
1-1.1 cm long, 2-3 mm wide. Lip simple, subpandurate,
1.3-1.4 cm long, 5-6 mm wide; with a short, semiterete
claw at the base; apical portion flat, truncate, and retuse;
margins entire; callus on the basal 2/5 trapezoid, smooth,
the base forming "shoulders," the apex emarginate.
Column slender, broadened above, without auricles, 1-
1.1 mm long; anther with 2 pollinia, obovate, sulcate,
with ovate stipe and viscidium. Capsules ca. 3.5 cm
long; pedicel 2-3 cm long.
Middle-canopy epiphytes growing in premon-
tane cloud forests on larger branches exposed to
strong winds at 1200-1400 m. Rare from Hon-
duras to Costa Rica. Flowering in August and
September.
Vegetatively A. costaricensis is typical of the
Oncidiinae, but the inflorescence has few flowers
that have a simple lip and a similarly simple cal-
lus. The petals are notably smaller than the sepals,
and the flower color is unique among Costa Rican
Oncidiinae.
Alphonse Heller wrote an unpublished note that
he found plants in Nicaragua at 2150 ft with a
pubescent callus among other differing features.
These plants may represent an undescribed species.
Aspasia Lindl.
(D. E. Mora de Retana)
REFERENCES — R. L. Dressier and N. H. Wil-
liams, An overlooked genus in the Oncidiinae.
Amer. Orchid Soc. Bull. 39:988-994. 1970. N. H.
Williams, Taxonomy of the genus Aspasia Lind-
ley (Orchidaceae: Oncidieae). Brittonia 26:333-
346. 1974.
Epiphytic, cespitose or shortly rhizomatous and creep-
ing herb. Pseudobulbs stalked, elliptic to oblong, var-
iously compressed laterally, subtended by 2-6 sheathing
and often foliaceous bracts. Leaves 1-2 at the pseudo-
bulb apex, conduplicate, usually thin. Inflorescence 1-
2, lateral at the base of the pseudobulb, usually not ex-
ceeding the shoot, of 1-10 flowers. Flowers usually rin-
gent, the base of the lip parallel to and partially united
to the column by the lateral margins. Sepals and petals
similar, the petals somewhat smaller and usually par-
tially adnate to the column. Lip larger than sepals and
petals (at least in Central American species), recurved
or reflexed sharply above the claw, the callus of keels,
or teeth, usually 3-lobate, the midlobe often emarginate.
Column erect, cylindric; anther with 2 pollinia, waxy,
with angular-ovate stipe with viscidium.
114
FIELDIANA: BOTANY
A genus of about six species from lower ele-
vations differing mainly in flower size and in de-
tails of the column and lip. Two species are
known in Central America, both in Costa Rica.
Williams compared Aspasia to several genera,
including Brassia, and Odontoglossum. Dressier
and Williams removed Aspasia pusilla C.
Schweinf. to the genus Cischweinfia. Chase and
Palmer's work suggests a relationship with a clus-
ter of genera including Ada, Brassia, Cischwein-
fia, Odontoglossum, Otoglossum, and Symphy-
glossum.
Key to the Species of Aspasia
la. Sepals < 2.5 cm, brown spotted . .
Ib. Sepals > 2.6 cm, lined with brown
A. epidendroides
. . A. principissa
Aspasia epidendroides Lindl., J. Bot. (Hooker)
1:6. 1834. TYPE: Panama and western Colom-
bia, Cuming s.n. (K, photo). Odontoglossum as-
pasia Rchb. f., Ann. Bot. Syst. 6:851. 1864. As-
pasia fragrans Klotzsch, Ind. Sem. Hort. Berol.
12. 1853. Aspasia barclayi Rolfe, Bull. Misc.
Inform. 210. 1892. Figure 27C.
Plant epiphytic, cespitose to shortly repent, forming
small clumps. Rhizome stout; roots 1-1.5 mm in diam-
eter. Pseudobulbs oblong-elliptic, strongly compressed
laterally, 4-16 cm long, 4-6 cm wide, somewhat stipi-
tate and hidden basally by 2-4 foliate sheaths, apically
2-foliate. Leaves articulate, the blade elliptic to lanceo-
late, conduplicate at the base and apically acute to acu-
minate, 8-30 cm long, 1.5-4.5 cm wide. Inflorescences
generally 2 borne laterally at the pseudobulb base, with
4-8 successive flowers, shorter than the subtending leaf;
ovary with pedicel 2-3 cm long. Flowers fragrant, the
sepals green spotted with maroon, petals rose, and lip
white, turning yellow with purple stain centrally. Sepals
subequal, subcoriaceous, elliptic-ovate to obovate,
somewhat concave, acuminate apically, 2.1-2.3 cm long,
8-10 mm wide; dorsal erect and inserted on the column;
lateral-spreading and reflexed. Petals spreading, elliptic-
ovate, somewhat spatulate, apiculate, adnate basally to
the column, 2.1-2.3 cm long, 9-10 mm wide. Lip ad-
nate to less than :/3 the column, lightly 3-lobate, reflexed
below the middle, 2-2.5 cm long, 1.5-2.5 cm wide, the
margin lightly undulate, the apex emarginate and some-
what fimbriate; callus of 2 fleshy, longitudinal, elevated
projections centrally, usually flanked by 2-4 shorter,
more or less parallel calli. Column stout, somewhat ar-
cuate, (1.7-)2.3 cm long; anther terminal, the anther pa-
pillose; pollinia 2, obpyriform, waxy, with prominent
stipe and viscidium. Capsule 6-8 cm long, cylindric,
pedicel 1-1.5 cm long.
A fairly common epiphyte of humid forests on
larger branches and trunks at 0-750(1000) m
along the Pacific slope. Flowering January to
April. Reported from Guatemala and El Salvador
to Colombia.
This species is easily distinguished from the
similar A. principissa Rchb. f. by the smaller
flowers, the spotted sepals and petals, and the lip,
which has a darker purple color. The synonymy
is based on Williams's treatment.
Aspasia principissa Rchb. f., Bot. Zeitung (Ber-
lin) 10:637. 1852. TYPE: Panama, Veraguas(?),
Warscewicz (w 45457, photo). Odontoglossum
principissa (Rchb. f.) Rchb. f., Ann. Bot. Syst.
6:852. 1864. Aspasia epidendroides Lindl. var.
principissa (Rchb. f.) P. H. Allen, Ann. Missou-
ri Bot. Card. 36:165. 1949. Aspasia bibriana
Rchb. f., Bonplandia 2:90. 1854. TYPE: Local-
ity unknown (w). Odontoglossum biberianum
[sic] (Rchb. f.) Rchb. f., Ann. Bot. Syst. 6:849.
1864. Aspasia rousseauae Schltr. Gartenflora
72:76. 1922. (lectotype: fig. 10 in Gartenflora
72:73). Figure 27D.
Plant epiphytic, to 40 cm high. Rhizome short; roots
about 1 .5 mm in diameter. Pseudobulbs oblong elliptic,
compressed, stalked, 7.8-15 cm long, 3.1-3.6 cm wide,
concealed at the base by 2-3-foliaceous sheaths, apically
2-foliate. Leaves subcoriaceous, deciduous, elliptic-lan-
ceolate to oblanceolate, conduplicate at the base, acute,
the blade 10-41 cm long, 1.3-3.8 cm wide. Inflores-
cence a lateral 4-6 successive-flowered raceme; ovary
with pedicel about 3 cm long subtended by subulate
bracts to 1.4 cm long. Flowers about 4.5 cm high and
2.5 cm wide in natural position, with greenish yellow
sepals and petals; sepals with diffused bars of reddish
brown, the petals longitudinally marked with the same
color; lip white turning yellow, stained centrally or lined
with magenta and with a yellow callus. Sepals subequal,
elliptic-lanceolate, lightly concave, apiculate and thick-
ened at the apex; dorsal erect, inserted on the back of
the column, 2.8-3 cm long, 7-9 mm wide. Petals ellip-
tic-lanceolate, carinate, enlarged and involute apically,
united to the base of the column, the apex emarginate,
and with a fleshy apicule; lateral sepals subfalcate, with
lightly undulate margins, 2.6 cm long, 7 mm wide. Lip
pandurate when spread, united to nearly Vi the column,
2.5-3.5 cm long, 2-2.8 cm wide; callus of a pair of
fleshy lamellate keels from the base to about the middle.
Column stout lacking wings apically, 2-2.5 cm long;
anther terminal, the anther lightly papillose; pollinia 2,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
115
obovoid with well-developed stipe and viscidium. Cap-
sule cylindric, about 8 cm long, with pedicel about 1.5
cm.
Epiphytic in lowland tropical rain forests be-
tween 10 and 300(500) m on the Atlantic slope,
where it grows on the larger, shaded branches of
the host. Flowering December to April. Reported
from Nicaragua(?), Costa Rica, Panama, and Co-
lombia.
This species closely resembles A. epidendro-
ides but differs in the larger flower, in the sepals
and petals with brown lines rather than spots, in
the much more constricted lip, and in the much
deeper anther. The synonymy given above follows
that of N. Williams.
Brassia R. Br.
(D. E. Mora de Retana)
REFERENCE — N. H. Williams, A reconsideration
of Ada and the glumaceous brassias (Orchida-
ceae). Brittonia 24:93-110. 1972.
Cespitose or repent epiphytic herb, usually with well-
developed pseudobulbs subtended by sometimes folia-
ceous bracts, the apex with 1-3 leaves. Leaves condu-
plicate, lanceolate, elliptic to oblanceolate. Inflores-
cence an arcuate, several- to many-flowered raceme
borne laterally at the base of the pseudobulb. Flowers
large and conspicuous, subtended by small elongate spa-
thaceous bracts. Sepals free, narrowly attenuate to cau-
date, sometimes of equal length but usually the dorsal
much shorter. Petals similar to the dorsal sepal but usu-
ally shorter. Lip simple or obscurely lobed, forming
more than a right angle with the column. Column short,
stout, and erect, lacking wings; anther terminal, pollinia
2, waxy. Fruit a capsule.
A Neotropical genus of about 38 species dif-
fering from Oncidium mainly by its attenuate flo-
ral segments. Four species are known in Costa
Rica. Norris Williams separated plants once as-
signed to the glumaceous section of Brassia into
the genus Ada on the basis of the smaller pseudo-
bulbs, leaves with several major veins, the often
several inflorescences per shoot, the large sheath-
ing leaves, and different pollinarium. According
to Chase and Palmer, DNA evidence suggests re-
lationships with Aspasia, Cischweinfia, and pos-
sibly some species of Odontoglossum and Sym-
phyglossum.
Key to the Species of Brassia
la. Pseudobulb apex with a single leaf B. arcuigera
Ib. Pseudobulb apex with >2 leaves
2a. Lip with conspicuous raised warts B. verrucosa
2b. Lip lacking warts
3a. Lip rhombic, at least 2 cm wide; petals with solid dark pigment on the lower third; pseudo-
bulbs tapering apically B. gireoudiana
3b. Lip lanceolate to ovate, < 2 cm wide; petals at most spotted on lower Va; pseudobulbs
fusiform to oblong B. caudata
Brassia arcuigera Rchb. f., Card. Chron. (1869)
388. TYPE: Peru, Veitch (w, photo). Brassia
antherotes Rchb. f., Card. Chron. 12:782. 1879.
TYPE: Colombia, Dagua, E. Klaboch (w, pho-
to). Brassia lawrenceana Lindl. var. longissima
Rchb. f., Card. Chron. 1313. 1868. TYPE: Bull-
er s.n. (w, photo). Brassia longissima (Rchb. f.)
Nash, Bail. Stand. Cyclop. Hort. 1:541. 1914.
Brassia antherotes Rchb. f. var. longissima
(Rchb. f.) Teuscher, Baileya 9:121-124. 1961.
Figure 28A.
Plant epiphytic, rarely terrestrial, cespitose, forming
small clumps to ca. 40 cm. Rhizome short; roots to 3 mm
in diameter. Pseudobulbs ovoid to elliptic-oblong, strongly
compressed laterally and sharp-edged, 6-14 cm long, 2.5-
4.5 cm wide, apically 1 -foliate, protected at the base by 0-
1 foliaceous sheaths. Leaves articulate, subcoriaceous, el-
liptic, conduplicate at the base, acute, prominently keeled
abaxially, 20-40 cm long, 4-6 cm wide. Inflorescence an
arcuate raceme with numerous flowers borne at the base of
the pseudobulb, 25-45 cm tall including the scape, the
nodes with triangular acute bracts; ovary with pedicel 1.5-
3 cm long. Flowers "spidery" in shape, very variable in
size, pleasantly fragrant in Costa Rican populations, disti-
chous, greenish white turning orange-yellow, with reddish
brown spots at the base of the sepals, to 30 cm tall, 7 cm
wide. Sepals similar, free, linear-lanceolate, attenuate, the
dorsal erect, to about 10 cm long, 4-5 mm wide; lateral
8-21 cm long and 5-6 mm wide. Petals much shorter than
sepals, free, linear-lanceolate, lightly falcate, attenuate, 4-
6 cm long, 4-6 mm wide. Lip simple, ovate-lanceolate,
acuminate, the margins smooth, 5-6 cm long, 1.2-1.5 cm
116
FIELDIANA: BOTANY
wide; callus at the base consisting of a pair of white, ele-
vated keels between which is a median, yellow-orange, pu-
bescent furrow. Column stout, 7-9 mm long; anther with
2 pollinia, waxy, obpyriform, with viscidium and broad
stipe.
Epiphytic, rarely terrestrial on embankments in
premontane rain forests at 500-1600 m. Flower-
ing in Costa Rica mostly March to October. Re-
ported from Costa Rica to Peru and Venezuela.
This species is the largest-flowered brassia in
Costa Rica. It is most similar to B. caudata, which
occurs at much lower elevations, and the pseudo-
bulbs of that species are apically 2-foliate with
rounded edges. We are following authors in rec-
ognizing the above synonymy.
Brassia caudata (L.) Lindl., Bot. Reg. 10: t. 832.
1824. Epidendrum caudatum L., Sp. PL (ed. 2)
1349. 1763. Malaxis caudata Willd. Sp. PL (ed.
4) 4:93. 1805. Oncidium caudatum Rchb. f.,
Ann. Syst. Bot. 6:766. 1863. Brassia caudata
var. hieroglyphica Rchb. f., Dl. Hort. 28:20.
1881. TYPE: "La Perle des Antilles" (w, pho-
to). Figure 28B.
Plant epiphytic, cespitose to 35 cm high. Rhizome short;
roots to 2.5 mm in diameter. Pseudobulbs fusiform to ob-
long, somewhat compressed, 6-10 cm long, 1.5-2.5 cm
wide, subtended at the base by foliaceous bracts when
young, apically 2-foliate. Leaves elliptic oblong to some-
what oblanceolate, coriaceous, carinate abaxially, 10-20
cm long, 2.5-4.5 cm wide, apex acute. Inflorescence a
lateral arcuate raceme of 6-12 distichous flowers, to about
30 cm long including the scape; ovary with pedicel 1-2
cm long. Flowers "spidery" with spreading segments very
variable in attenuation, greenish yellow turning orangish
with age, marked with maroon; lip white turning yellow
with wine red spots. Sepals dissimilar, free, linear-lanceo-
late, attenuate; dorsal erect and with incurved apex, to 7(12)
cm long, 5-8 mm wide; lateral sepals 12-20(33) cm long,
5-7 mm wide. Petals lanceolate, falcate, attenuate, 2.5-5.3
cm long, 3-5 mm wide. Lip simple, oblong lanceolate,
acuminate, 3—4.5(5.5) cm long, 1.2-1.5 cm wide; callus of
2 longitudinal, pubescent keels, each terminated by a re-
curved tooth and between which is a pubescent furrow.
Column stout, 3-5 mm long; anther cucullate; pollinia 2,
obpyriform, with stipe and viscidium. Capsules 4-5 cm
long including the beak, pedicel about 1.5 cm long.
This species grows in tropical lowland rain for-
ests at 0-200 m, where it is usually uncommon
though widespread. Flowering in Costa Rica
mostly March to July. Reported from Florida,
Cuba, and Jamaica and from Mexico and Central
America to Bolivia and northern South America.
Brassia caudata is most similar to B. arcuigera
Rchb. f., a species that has 1 -foliate, sharp-edged
pseudobulbs and occurs at higher elevations.
Also, the flowers of that species average larger in
Costa Rican populations.
Available specimens from the Atlantic drainage
have flowers that match very well Linnaeus's con-
cept based on the plate by Plumier. Two herbari-
um sheets (AMES) based on a plant collected at
Pigres on the Pacific Coast by C. Lankester have
much larger flowers (lateral sepals 33 cm long)
and may represent natural hybrids or a different,
perhaps unnamed species.
Brassia gireoudiana Rchb. f. & Warsc., Allg.
Gartenzeitung 22:273-274. 1854. TYPE: War-
scewicz (not seen). Figure 28C.
Plant a somewhat scandent epiphyte. Rhizome elon-
gate, 3-4 cm between shoots, covered by dark bracts;
roots fleshy, to 2.5 mm in diameter. Pseudobulbs ellip-
tic-ovoid, tapering, little compressed laterally, 6-14 cm
long, 2.5-4.5 cm wide, subtended by 1 or more folia-
ceous bracts when young, becoming deciduous with age,
apically 2(rarely 1 )-foliate, the second leaf borne higher
than the first. Leaves petiolate, subcoriaceous, carinate
beneath, elliptic-ovate, 20-35 cm long, 2.2-5 cm wide,
acute and apiculate apically. Inflorescence a lateral, ar-
cuate raceme of 1-15 flowers neatly arranged on the
axis, 75-100 cm long including the scape, borne at the
base of the immature to mature shoot; ovary with ped-
icel 1.8-3 cm long. Flowers large and spectacular, 20-
25 cm high, greenish white with brown dots at the base
of the sepals, the base of the petals solid dark brown;
lip greenish white with brown spots. Sepals similar, free,
linear-lanceolate, attenuate to the apex; dorsal erect, 10-
13 cm long, 4-5 mm wide; lateral sepals 1 1-20 cm long,
4-5 mm wide. Petals similar to sepals but shorter, 5-10
cm long, 4-5 mm wide, linear-lanceolate, subfalcate.
Lip simple rhombic, narrowed at the base forming a
narrow claw united to the base of the column, the apex
acuminate, the margins lightly crisped, 3.5-7 cm long,
2-3 cm wide; callus at the base fleshy, a pair of keels,
somewhat pubescent at the base, rounded and elevated
apically. Column stout, 5-6 mm long, somewhat broad-
ened apically; anther with 2 waxy, ovoid, furrowed, pol-
linia supported on a stipe and viscidium.
Brassia gireoudiana grows in semideciduous to
evergreen premontane forests at 500-1000 m at
least on the Pacific slope of southern Costa Rica.
Flowering February to May. Southern Costa Rica
and adjacent Chiriqui, Panama.
This species can be confused with B. verrucosa
Lindl. but lacks the warts on the lip of that spe-
cies, and the inflorescence often forms on imma-
ture shoots.
Brassia gireoudiana is apparently closely relat-
ed to Brazilian B. arachnoidea Barb.-Rodr. Both
species share nearly identical color patterns in the
flowers and a similar rhombic lip. Also, the pseu-
dobulbs are little compressed laterally.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
117
Brassia verrucosa Lindl., Bot. Reg. 26: misc. 36.
1840. Figure 28D.
Plant epiphytic, cespitose, forming small clumps, to
about 65 cm tall. Rhizome short; roots to 3 mm in di-
ameter. Pseudobulbs ovoid-conic, furrowed, little com-
pressed laterally, 6-11 cm long, 3.1-4 cm wide, sub-
tended by 1-2 foliaceous bracts, apically 2-foliate.
Leaves coriaceous, conduplicate at the base, the apical
leaves shortly petiolate, the blade elliptic-lanceolate,
acute, 16-30 cm long, 2.6-4.5 cm wide. Inflorescence
a lateral, erect, or arcuate raceme of 6-10 flowers in
Costa Rican populations, 50-65 cm long including the
scape; ovary with pedicel 2.5-3 cm long, subtended by
much shorter subulate bracts. Flowers attractive, pleas-
antly fragrant in Costa Rican populations, to 25 cm high,
distichous, yellowish white with bars and spots of red-
dish brown on the sepals and petals, olive green warts
on the lip. Sepals similar, free, linear-lanceolate, atten-
uate and somewhat canaliculate; dorsal erect, 6.5-13.5
cm long, 6-7 mm wide; lateral sepals 7.5-17.5 cm long,
6-7 mm wide. Petals linear-lanceolate, falcate, attenu-
ate, 4-9 cm long, 6-7 mm wide. Lip obscurely 3-lobate,
trullate-subpandurate, broadest just below the middle,
acuminate at the apex, verrucose below, 3.5-5.2 cm
long, 2.4-3.2 cm wide; callus a pair of keels on the
lower 1A, each keel terminated by a tooth. Column erect,
7 mm long; anther with 2 pyriform, sulcate pollinia with
stipe and viscidium.
Brassia verrucosa is a species of evergreen to
semideciduous cloud forests at 1300-1800 m,
where it grows on trunks and larger branches of
its host. Flowering December to May. Reported
from Mexico to Costa Rica.
Plants observed from northern Nicaragua have
smaller and usually more numerous flowers with a
rather unpleasant odor compared to those from Cos-
ta Rica. Also, Costa Rican plants have fewer warts
on the Up than those farther north. We hesitate to
repeat the synonymies given by authors because
some of them may prove to be different. Although
the type of B. verrucosa was not determined, a spec-
imen on microfiche, identified by Lindley as the
species, was seen.
Cischweinfia Dressier & N. Williams
(D. E. Mora de Retana)
REFERENCE — R. L. Dressier and N. H. Williams,
An overlooked genus in the Oncidiinae. Amer.
Orchid Soc. Bull. 39:988-994. 1970.
Cespitose, usually small epiphytic herbs. Pseudo-
bulbs usually ellipsoid to ovoid, 1 -foliate, compressed,
subtended by 1 or more foliaceous sheaths. Leaves con-
duplicate, carinate, elliptic to lanceolate, acute, thin in
Costa Rican species. Inflorescence a lateral raceme,
shorter than the subtending foliaceous bract in Costa Ri-
can plants, with 1-5 flowers. Flowers spreading, usually
attractive. Sepals and petals free, similar. Lip simple,
with basal margins usually clasping the column. Col-
umn with a pair of auricles beneath the stigma, with
distinct hood-shaped and often petaloid anther bed; an-
ther often papillose; pollinia 2 supported on an elongate
stipe with viscidium. Fruit a capsule.
A Neotropical genus of about nine species
ranging from Costa Rica to South America. There
are two species in Costa Rica.
Cischweinfia was described by Dressier and
Williams to accommodate species previously clas-
sified in the genera Aspasia, Miltonia, and Tri-
chopilia. Chase and Palmer published two clado-
grams suggesting relationships with Ada, Aspasia,
Brassia, and possibly several other genera as well.
Key to the Species of Cischweinfia
la. Lip surrounding column at base, tepals green suffused with pink; lip white with pink spots
C. dasyandra
Ib. Lip cuneate, not surrounding the column, tepals brown with yellowish green apex; lip white with
yellowish orange at the base C. pusilla
Cischweinfia dasyandra (Rchb. f.) Dressier &
N.Wms., Amer. Orchid Soc. Bull. 39:991. 1970.
Trichopilia dasyandra Rchb. f., Xen. Orch. 3:
64, t. 230. 1883. TYPE: Costa Rica, Endres
(w). Figure 29A.
Plant epiphytic, cespitose. Rhizome short, the roots
slender, 1-1.5 mm in diameter. Pseudobulbs oblong,
compressed, 3-8 cm long, 6-11 mm wide, apically 1-
foliate, subtended by 2-3 foliaceous bracts. Leaves lin-
ear-lanceolate, conduplicate at the base, acute, keeled
abaxially, 9-20 cm long, 7-12 mm wide. Inflorescence
a short, somewhat pendulous 1-4-flowered raceme
borne from the base of the pseudobulb, 4-7 cm long,
ovary and pedicel 13-20 mm long. Flowers with green
sepals and petals suffused with pink, the lip white with
pink spots and yellow in the center, 2.4-2.7 cm long and
1.8-2.2 cm wide in natural position. Sepals subequal,
118
FIELDIANA: BOTANY
free, elliptic-lanceolate, acute, 1.2-1.4 cm long, 2-3 mm
wide; lateral subfalcate and somewhat larger than the
dorsal. Petals elliptic-oblong, acute, subfalcate, 1-1.2
cm long and 2-3 mm wide. Lip with basal margins fold-
ed around the column, obovate to suborbicular when
spread, emarginate, 1-1.2 cm long and 9-10 mm wide,
the margins crenulate; callus basal, glabrous, yellow,
with 2 small keels. Column 5-7 mm long, erect, semi-
terete, white, with a prominently dentate to fimbriate an-
ther bed; pollinia 2, orbicular translucent, with narrow
stipe and elongate viscidium; stigma bilobed.
Uncommon epiphyte of wet premontane forests
at 700-1100(1650) m. Flowering mostly June to
September. Costa Rica and Panama; reported also
from Colombia and Ecuador.
This species is distinguished from similar C.
pusilla (C. Schweinf.) Dressier & N. H. Williams
by the smaller plants, larger flowers of different
color, and overlapping sides of the lip base folded
around the column.
Cischweinfia pusilla (C. Schweinf.) Dressier &
N. H. Williams, Amer. Orchid Soc. Bull. 39:
992. 1970. Aspasia pusilla C. Schweinf., Bot.
Mus. Leafl. 10:21-23, t. 1. 1941. TYPE: Pan-
ama, Darien, M. E. & R. A. Terry 1502 (holo-
type: AMES, photo; isotype: F). Trichopilia pus-
illa (C. Schweinf.) Garay, Orquideologia 5:20.
1970. Figure 29B.
Plant a cespitose epiphyte sometimes forming large
clumps, 20-25 cm high; rhizome short; roots to 1.5 mm
in diameter. Pseudobulbs elliptic-ovate, compressed, 1-
foliate, 3-3.5 cm long, 1-2.1 cm wide, concealed at the
base by 2-3 foliaceous bracts. Leaves coriaceous, cari-
nate, with prominent conduplicate petiole; blade elliptic-
lanceolate, 10-20 cm long, 1-1.5 cm wide, acute. Inflo-
rescence 1-2, lateral at the base of the pseudobulb,
about Vi as tall as the plant, 8-10 cm long; lax raceme
of 4-5 flowers; ovary with pedicel 2.5-3 cm long, sub-
tended by a triangular, acute bract to 8 mm long. Flow-
ers with brown tepals with lemon yellow apices; lip
white with yellow-orange lines at the base. Sepals sim-
ilar, oblong-elliptic, dorsally carinate, 1.1-1.3 cm long,
3-4 mm wide, the apex acute, apiculate; dorsal erect;
lateral sepals subfalcate. Petals similar to the sepals, ob-
lanceolate-elliptic, 9-11 mm long, 3-3.5 mm wide,
oblique at the base, the apex acute. Lip cuneate, simple,
suborbicular to obovate when spread, 1.1-1.3 cm long,
1-1.2 cm wide, adnate basally to the middle of the col-
umn, rounded. Column short, 5 mm long, with a pair
of auricles below the stigma; anther bed petaloid, irreg-
ularly dentate, forming a hood; anther papillose, trian-
gular in profile; pollinia 2, obovate with elongate stipe
and viscidium.
A poorly known and rare epiphyte of tropical
humid forests at 600-700 m. Flowering June to
August. Costa Rica and Panama; probably also in
Colombia.
Cischweinfia pusilla closely resembles C. das-
yandra but differs notably in the cuneate lip,
which does not enclose the column, and in flower
color.
Comparettia Poeppig & Endl.
(D. E. Mora de Retana)
REFERENCE — I. Bock, Revision der Gattung
Comparettia Poepp. & Endl. (part 1). Orchidee
(Hamburg) 37:193-196. 1986.
Epiphytic, cespitose herb often forming small clumps.
Pseudobulbs cylindric, small, often hidden by subtending
foliaceous bracts, apex 1 -foliate. Leaves subcoriaceous,
conduplicate, usually ovate to elliptic, obtuse to acute,
petiole essentially none. Inflorescence a laterally borne
raceme or panicle, usually with long scape; ovary dis-
tinctly pedicellate, subtended by short, triangular bracts.
Flowers brightly colored. Sepals dissimilar, the dorsal
free, the lateral sepals connate and forming a conspicuous
single spur. Petals ovate, usually acute. Lip much longer
than the tepals, with a bifid appendage at the base inserted
within the sepaline spur, the lamina conspicuously emar-
ginate to 2-lobate. Column stout, usually with broadened
stigma; anther with 2 pollinia supported on a slender stipe
with viscidium. Fruit a capsule.
A genus of 10-12 vegetatively similar species
distributed throughout the neotropics. Only a sin-
gle species occurs in Costa Rica. The genus is
easily recognized by the brilliantly colored, prob-
ably bird-pollinated flowers with large, emargin-
ate to 2-lobate lip and conspicuous sepaline spur.
According to Chase and Palmer, Comparettia is
closely related to Rodriguezia and Scelochilus but
surprisingly not to lonopsis.
Comparettia falcata Poeppig & Endl., Nov. Gen.
Sp. 1:42, t. 73. 1836. TYPE: Peru, between
Cassapi and Pampayacu, Poeppig 1646 (not
seen). Comparettia rosea Lindl., Edwards's
Bot. Reg. 26: misc. 78. 1840. TYPE: "Spanish
Main," drawing based on Loddiges 752 (K).
Figure 29C.
Plant epiphytic, cespitose, forming small clumps.
Rhizome short, the roots stout, much branched, 1-1.5
mm in diameter. Pseudobulbs cylindric, 1 -foliate, 1.5-
2.5 cm long, 7-10 mm broad, green suffused with pur-
ple, subtended by papery bracts. Leaves variable, ellip-
tic, coriaceous to somewhat fleshy, the apex rounded and
apiculate or bluntly acute, 5.5-19 cm long, 1.5-5.5 cm
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
119
wide. Inflorescence lateral, 15-55 cm long including the
scape and panicle, rarely subtended by a foliaceous
bract; ovary and pedicel about 1.5-2 cm long. Flowers
purplish red or pink with white centrally, 2-2.3 cm high
and 1 .5-0.7 cm wide in natural position. Sepals dissim-
ilar; the dorsal free, elliptic-lanceolate, acuminate, 10-
12 mm long and 4-6 mm wide; lateral connate for their
entire length, forming a spur at the base, 2.2-2.4 cm
long including the spur, 5-6 mm wide. Petals free, ob-
ovate, the apex rounded, apiculate, 11-12 mm long, 5-
7 mm wide. Lip obscurely 3-lobulate, 1.3-1.5 cm long
and 1.4-1.6 cm wide, the lateral lobes small, subtrian-
gular; midlobe reniform, deeply emarginate, contracted
at the base, forming a claw united by its margins and
central keel to the column and extending at the base into
2 filiform appendages; disk glabrous and somewhat
fleshy with a white callus. Column stout, 4-5 mm long,
broadened above, with a pair of wings near the apex;
anther with 1 cell; pollinia 2, waxy, with conspicuous
stipe and viscidium; stigma entire, somewhat bilobed.
Capsules variable depending on plant size, 2-5 cm long
including the conspicuous beak, with pedicel 0.8-1.7 cm.
Epiphytic on small branches and trunks in open
canopies of premontane and lower montane for-
ests at 800-2000 m. Comparettia falcata is one
of few orchids that colonize twigs of guava trees.
Flowering mostly May to August, September to
November, and probably sporadically .throughout
the year. Widespread throughout the neotropics.
Comparettia falcata is easily identified by the
pink to red or purple flower with proportionately
large, 2-lobate lip and conspicuous spur.
Fernandezia Ruiz & Pav.
(D. E. Mora de Retana)
Plant a monopodial epiphyte. Stems erect, creeping
or pendulous, concealed by leaf sheaths; pseudobulbs
lacking. Leaves conduplicate, distichous, articulate,
chartaceous, coriaceous, to fleshy, usually elliptic to ob-
long. Inflorescence lateral, short, a few-flowered ra-
ceme. Flowers showy, red to yellow, orange, or purple,
never green, spreading to campanulate. Sepals similar,
the laterals sometimes partially connate. Petals usually
larger than the sepals. Lip usually simple, often flabel-
late, sometimes the base parallel with the column and
the apex somewhat reflexed, callus present and usually
very different between species. Column simple, often
with broad, dentate wings apically; anther with 2 pollinia
with elongate, simple, or furcate stipe, or each with sep-
arate stipe; viscidium elongate. Fruit a capsule.
A high-elevation genus of perhaps 12 species.
Only a single species is known from Central
America.
Fernandezia is probably most closely related to
Pachyphyllum, which has an identical growth
habit but much smaller, greenish to whitish flow-
ers. Dressier (1993) notes the similarity of growth
habits and pollinia to Lockhartia and Raycadenco
and places both Fernandezia and Pachyphyllum
in the Oncidiinae.
Fernandezia tica Mora-Retana & J. B. Garcia,
Brenesia 39-40:163-165. 1993. TYPE: Costa
Rica, Heredia, road to Vara Blanca, 2000 m, 27
Oct. 1992, J. T. Atwood & O. Rodriguez 4181
(holotype: usj). Figure 29D.
Plant epiphytic, monopodial, small, to about 8 cm
long. Stems somewhat creeping; roots to 2 mm in di-
ameter. Leaves fleshy, green, purplish and keeled be-
neath, elliptic, ca. 20 mm long, 8 mm wide; forming a
prominent sheath at the base ca. 5 mm long; apex acute.
Inflorescence lateral, short, of 2-3 flowers; ovary with
pedicel prominently winged, ca. 13 mm long. Flowers
fleshy campanulate, red-purple, ca. 8 mm long. Sepals
subequal, shortly connate at the base, ovate -elliptic,
prominently keeled abaxially, 10 mm long, 6 mm wide,
acute; dorsal erect; lateral sepals subfalcate. Petals el-
liptic-ovate, 8 mm long, 6 mm wide, the apex recurved.
Lip simple, elliptic-ovate to somewhat trulliform, 8 mm
long, 4 mm wide, somewhat recurved near the middle,
forming a cavity at the base; callus on the lower % of 2
slender prominent keels ending in a fleshy transverse
ridge and bifurcating above. Column robust, 7 mm long,
with prominent wings above; anther about 1.5 mm in
diameter; pollinia 2, elliptic-ovate supported on separate
stipes and connected to a prominent elongate viscidium.
Capsule unknown.
Very poorly known epiphyte at 2000 m, where
the type was found growing in strong winds and
frequent cloud cover but occasionally exposed to
nearly full sun. Flowering in late October but to
be expected at other times of the year. Apparently
endemic to Costa Rica.
The existence of a fernandezia in Costa Rica
has been known for some time, but few specimens
had been collected until recently. No other plants
were observed growing near the holotype, and the
plant is apparently rare.
Goniochilus M. W. Chase
(D. E. Mora de Retana)
REFERENCE — M. W. Chase, Revisions of Hy-
bochilus and Goniochilus (Orchidaceae). Contr.
Univ. Michigan Herb. 16:109-127. 1987.
Plant a cespitose herb epiphytic on twigs. Pseudo-
bulbs ovoid to suborbicular, ancipitous; base concealed
by 2-3 foliaceous bracts; apex 1 -foliate. Leaves coria-
ceous, elliptic-lanceolate, shortly petiolate at the base,
120
FIELDIANA: BOTANY
the apex acute. Inflorescence lateral at the base of the
pseudobulb, a raceme in small plants, paniculate when
large. Flowers small, colors weakly attractive. Sepals
dissimilar, carinate, acuminate, the laterals united to %
their length. Petals simple, oblong-elliptic, obtuse. Lip
3-lobate, the midlobe emarginate; callus simple. Col-
umn stout, the base forming a short nectary with the lip;
apex with a pair of auricles; anther more or less dorsal;
pollinia 2. Fruit a capsule.
A monotypic genus differing from closely re-
lated Leochilus by the conspicuously 3-lobate lip
and the column, which is somewhat bent upward.
The close apparent relationship with Leochilus is
supported by evidence from DNA by Chase and
Palmer.
Goniochilus leochilinus (Rchb. f.) M. W. Chase,
Contr. Univ. Mich. Herb. 16:125. 1987. Rodri-
guezia leochilina Rchb. f., Gard. Chron. 1:970.
1871. TYPE: Costa Rica. Mesospinidium leo-
chilinum (Rchb. f.) Schltr., Repert. Spec. Nov.
Regni Veg. 19:253. 1923. Figure 30A.
Plant cespitose, epiphytic, somewhat reddish 6-15
cm tall excluding the inflorescence. Rhizome short, roots
slender, to 1(1.5) mm in diameter. Pseudobulbs 1 -foli-
ate, ellipsoid-ovoid to suborbicular, ancipitous, (0.6) 1 .5-
2 cm long, 1.3-1.7 cm wide, covered when young by
foliaceous bracts deciduous with age. Leaves condupli-
cate, petiolate, the blade elliptic-lanceolate, carinate
abaxially, acute, (2.5)6-14 cm long, (0.6)1.3-2.4 cm
wide, the apex asymmetric. Inflorescences 1-2, lateral,
pendulous, 10-32 cm long including the scape, with few
to many flowers; ovary and pedicel 3-6 mm long, sub-
tended by a triangular, acute bract 2-4 mm long. Flow-
ers 1 cm high and 5 mm wide; sepals and petals greenish
yellow with red-brown spots, the lip white with reddish
spots. Sepals subequal, strongly concave, carinate, the
dorsal ovate, 6-9 mm long, 3-5 mm wide; lateral sepals
fused for about % their length, each 6-9 mm long, 2-3
mm wide. Petals ovate-elliptic, 5-7 mm long, about 3
mm wide. Lip 3-lobate, 9-11 mm long, 4 mm wide;
lateral lobes semicircular to triangular; midlobe su-
bquadrate, convex, emarginate, the apical Vi forming an
obtuse angle with the base; callus on the lower % a sim-
ple sulcate pad, higher toward the base. Column 4-7
mm long; anther globose; pollinia 2, ovoid, with broad
triangular stipe and prominent elliptic viscidium. Cap-
sules ellipsoid, with 3 conspicuous ribs, 1.5-2.5(4.5) cm
long including the beak; pedicel 4-5 mm long.
Goniochilus leochilinus grows on twigs and
slender branches of trees, often in abandoned cof-
fee, citrus, and guava plantations in premontane
and lower montane rain forests at 600-1600 m.
Plants are common but little collected. Flowering
September to January. Reported from Nicaragua
to Panama.
This species can be confused with species of
Leochilus, but the lip is clearly 3-lobate and the
column somewhat bent upward, putting the anther
in a dorsal position.
Hybochilus Schltr.
(D. E. Mora de Retana)
REFERENCE — M. W. Chase, Revisions of Hy-
bochilus and Goniochilus (Orchidaceae). Contr.
Univ. Michigan Herb. 16:109-127. 1987.
Cespitose epiphytic herb of twigs and small branches.
Pseudobulbs ovoid to suborbicular, subtended by 2-3
foliaceous sheaths; apex l(2)-foliate. Leaves coriaceous,
conduplicate, elliptic-lanceolate, the apex unequally 2-
lobate. Inflorescences 1-2 per shoot, lateral, pendent pa-
niculate. Flowers minute, campanulate with rather dull
colors. Sepals dissimilar, the lateral to % united; petals
free, connivent with the sepals; lip simple or obscurely
3-lobate, forming a nectary at the base with the column,
subquadrate, retuse; column stout, anther terminal.
A monotypic genus endemic to Costa Rica and
possibly western Panama.
According to Chase, Hybochilus is closely re-
lated to both Leochilus and to Goniochilus. It is
easily distinguished from both by the much small-
er, campanulate flowers borne in dense panicles.
Hybochilus inconspicuus (Kraenzl.) Schltr., Re-
pert. Spec. Nov. Regni Veg. 16:430. 1920. Rod-
riguezia inconspicua Kraenzl. Bull. Herb. Bois-
sier 3:630. 1895. Rodriguezia candelariae
Kraenzl. Bot. Jahrb. Syst. 54, Beibl. 117:32.
1916. Leochilus parviflorus Standley & L. O.
Williams, Ceiba 1:235. 1951.
Plant a cespitose epiphyte 2-11 cm tall. Pseudobulbs
suborbicular to elliptic-ovoid, rugose when old, 1-2.5
cm long, 0.5-2 cm wide, subtended by 1-2 foliaceous
bracts, apex 1 -foliate. Leaves coriaceous, carinate, ellip-
tic to lanceolate, 1.5-7.5 cm long, 0.6-2.5 cm wide,
sometimes shortly petiolate, apex unequally 2-lobate.
Inflorescences 1-2 per shoot, paniculate, lateral, pen-
dent, many-flowered, 6-20 cm tall; ovary with pedicel
2-4 mm long, subtended by shorter, triangular, acute
bracts. Flowers minute, inconspicuous, rather closed, se-
pals greenish with purplish margins, petals whitish with
purplish margins, lip white with pale purple spots. Se-
pals dissimilar, the dorsal free, laterals to % connate,
ovate, 2.5-3 mm long, 1.5-2 mm wide. Petals ovate,
2.5-3 mm long, 2 mm wide, obtuse. Lip simple or ob-
scurely 3-lobate, 3.5-5 mm long, 1.5-2 mm wide, the
apex emarginate; callus an elongate, 2-lobate pad. Col-
umn stout, 0.5-1 mm long, about 1 mm in diameter, the
apex with a pair of extrorse stigmatic arms, forming a
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
121
nectary at the base with the lip. Capsules 0.8-1(1.5) mm
long including pedicel and beak.
Rather common twig epiphytes in evergreen
and often disturbed cloud forests at 950-1500 m;
often seen in coffee plantations. Flowering (Jan-
uary) February to May (June). Costa Rica, es-
pecially in the Central Valley; also reported from
Panama.
This species is easily recognized by the minute
flowers on plants resembling Leochilus. The
above synonomy and floral measurements are
taken from Chase's revision.
lonopsis H. B. K.
(D. E. Mora de Retana)
Epiphytic herb usually on twigs; rhizomes short to
much elongate, with at most rudimentary pseudobulbs.
Leaves lateral, rarely at the pseudobulb apex, terete or
flat, conduplicate and somewhat fleshy. Inflorescence
lateral, slender, erect or arcuate, a simple raceme or pan-
icle, with elongate scape. Flowers usually small. Sepals
and petals subequal, the lateral sepals connate at the
base producing a short sac; lip clawed and adnate to the
column base, much exceeding the tepals, 2-lobate. Col-
umn footless, lacking wings; pollinia 2, waxy, supported
on viscidium and elongate stipe. Fruit a capsule.
A mostly lowland genus of three species rang-
ing throughout the neotropics.
Key to the Species of lonopsis
la. Leaves terete
Ib. Leaves flat .
. . /. satyrioides
I. utricularioid.es
lonopsis satyrioides (Sw.) Rchb. f., Ann. Bot.
Syst. 6:683. 1863. Epidendrum satyrioides Sw.,
Prod. 123. 1788. TYPE: Hispaniola. lonopsis
costaricensis Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 19:62. 1923. TYPE: Costa Rica,
Werckle 25 (not seen).
Plant a cespitose epiphyte of twigs and smaller
branches; roots fibrous, to 0.5 mm in diameter. Pseudo-
bulb nearly obsolete and often hidden by subtending
leaves. Leaves 3-6, terete, acute, 2.5-13 cm long, to 3-
4 mm in diameter. Inflorescence a lateral successively
flowered raceme 6-19 cm long including the scape; ova-
ry with pedicel 8-10 mm long, subtended by much
shorter subulate bracts. Flowers white or with fine pur-
ple lines, with yellow callus. Sepals somewhat campan-
ulate, shortly connate at the base and not widely spread-
ing, lanceolate-elliptic, acute, 5-6 mm long, 1.5-2 mm
wide, the lateral forming a short mentum. Petals oblan-
ceolate to elliptic, obtuse, recurved, 5.5-6.5 mm long,
1.5-2 mm wide. Lip cuneate, lightly emarginate, 7.5-8
mm long, 2.5-3, with a pair of mammillate calli at the
base. Column stout, 1.5-2 mm long. Capsule obovoid,
about 1 cm long.
Epiphytic and common in disturbed moist for-
est at 0-750 m. Flowering August to October. Re-
ported throughout Mesoamerica, West Indies, and
South America. lonopsis satyrioides is easily rec-
ognized by the terete leaves and small, whitish
flowers with large, emarginate lip. There is some
suggestion that there may be more than one spe-
cies here accepted as one.
lonopsis utricularioides (Sw.) Lindl., Coll. Bot.
t. 39A. 1821. Epidendrum utricularioides Sw.,
Prodr. 122. 1788. Figure 30B.
Plant an epiphyte, usually of twigs and smaller
branches, usually with only 1-2 shoots, to 15 cm high;
roots fibrous, 0.5-1 mm in diameter. Pseudobulbs short,
6-10 mm tall, 4-5 mm wide, concealed by the subtend-
ing 2-4 foliaceous sheaths. Leaves distichous, some-
what fleshy, with conspicuous raised veins adaxially,
variable in shape from elliptic to oblong or oblanceolate,
acute, 4-15 cm long, 0.6-2 cm wide. Inflorescence a
somewhat successively flowered panicle borne laterally
near the pseudobulb base, to 50 cm high including the
peduncle; ovary with pedicel 5-10 mm long. Flowers
white or pink, veined with magenta. Sepals dissimilar;
dorsal ovate, acute to obtuse, 3.5-4.5 mm long, 1.8-2.3
mm wide; lateral sepals shortly connate, acute, 4.5-5.5
mm long, 1.4-2 mm wide, forming a short mentum at
the base. Petals ovate, 3.5-5 mm long, 1.8-2.5 mm
wide. Lip shortly clawed at the base, flabellate-obovate
and deeply emarginate, 10-12 mm long, 7-10 mm wide,
with 3 purple lines at the base, and a pair of white calli.
Column cylindric, stout, 2 mm long; anther with cucul-
late, somewhat beaked anther, pollinia 2 supported on
an elongate stipe and elongate viscidium. Capsules el-
lipsoid, without beak, about 2.5 cm long; pedicel 7-8
mm long.
Epiphytic on twigs and smaller branches of
usually disturbed evergreen forests, often seen in
coffee and guava at 0-880 m. Flowering mostly
January to May (August). Widespread and locally
common throughout the wetter lowland neotrop-
ics.
lonopsis utricularioides is easily recognized by
122
FIELDIANA: BOTANY
the flat, somewhat fleshy leaves, the pseudobulb,
which usually lacks an apical leaf, and the pani-
cles of the pinkish flowers with a large, emargin-
ate lip.
Leochilus Knowles & Westcott
(D. E. Mora de Retana)
REFERENCES — M. W. Chase, A Monograph of
Leochilus (Orchidaceae). Systematic Botany
Monographs 14. 97 pp. 1986. M. W. Chase, Pol-
lination ecology of two sympatric, synchronously
flowering species of Leochilus in Costa Rica. Lin-
dleyana 1:141-147. 1986.
Small cespitose herb, epiphytic, often on twigs and
smaller branches. Pseudobulbs elliptic-ovoid to subor-
bicular, compressed, 1-2-foliate, subtended by 2-3
leaves, the apex 1-2-foliate. Leaves coriaceous, condu-
plicate, elliptic to lanceolate, acute to obtuse. Inflores-
cence an erect or pendent lateral raceme or panicle with
slender peduncles and 1-many successively borne flow-
ers. Flowers small, colors generally dull. Sepals spread-
ing, the laterals free or variously connate, adnate to lip
base. Petals similar to the sepals, free. Lip adnate to
column base forming a nectary, longer than the sepals
and petals, simple or lightly 2-lobate, with a fleshy callus
near the base. Column footless, short, with arms on each
side at about the middle; pollinia 2, waxy. Fruit a cap-
sule.
A Neotropical genus of about nine species cen-
tered mainly in Mesoamerica and the Caribbean;
also in northern and western South America.
Three species are known in Costa Rica.
The relationships of Leochilus are clearly with
Goniochilus and Hybochilus, and possibly with
other genera of mostly twig epiphytes. For syn-
onymies see the revision of Leochilus by Chase.
Key to the species of Leochilus (including Hybochilus and Goniochilus)
la. Sepals > 3.5 mm long 2
2a. Lip simple, whitish to yellow often with dull spots 3
3a. Plants clearly reddish Leochilus labiatus
3b. Plants clearly green 4
4a. Pseudobulbs prominent, ovoid, usually > 2 cm long, low-elevation plants
Leochilus scriptus
4b. Pseudobulbs mostly hidden by foliaceous bracts, < 1.9 cm tall, plants mid to high
elevation Leochilus tricuspidatus
2b. Lip 3-lobate, white with dark spots Goniochilus leochilinus
Ib. Sepals < 3.25 mm long Hybochilus inconspicuus
Leochilus labiatus (Sw.) Kuntze, Revis. gen. pi.
2:656. 1891. Epidendrum labiatum Sw., Nov.
gen. & sp. pi., Prodr. 124. 1788. Figure 30C.
Plant a small cespitose twig epiphyte 2-5 cm tall ex-
cluding inflorescence; roots long, to about 1 mm in di-
ameter. Pseudobulbs compressed, orbicular to ovate, 1-
1.8 cm tall, with 1-2 leaves at the base, apex 1-foliate.
Leaves coriaceous, elliptic to lanceolate, acute, 2-4.5
cm long and 1-1.5 cm wide. Inflorescence a raceme or
panicle with 1-2 branches, lateral at the pseudobulb
base, successively flowered, 2-12 cm long including the
peduncle; ovary with pedicel about 6-9 mm long. Flow-
ers with sepals and petals yellowish green marked with
red-brown and with a yellow lip marked with red-brown.
Sepals dissimilar, the dorsal sepal ovate, acute, cucul-
late, 5-6.5 mm long, 2.5 mm wide; the lateral sepals
free to l/z united, about 5-6 mm long. Petals ovate-el-
liptic, about 4-5 mm long. Lip obovate, emarginate,
about 7-8 mm long and 3-4 mm wide, provided with a
shallow basal nectary and a central fleshy callus most
prominent below the middle. Column about 2.5 mm
long, with a pair of oblong arms at about the middle;
anther with 2 pollinia supported on an elongate stipe and
viscidium. Capsules ellipsoid, weakly 3-carinate, 2.5 cm
long including the prominent beak; pedicel 8-10 mm
long.
Common twig epiphytes, of evergreen forests
at 50-2000 m. It is often weedy in coffee plan-
tations, citrus groves, and guava. Flowering most-
ly October to April. For distribution Chase in-
cludes the West Indies, Mesoamerica, and north-
ern South America.
Leochilus labiatus is easily distinguished from
the other species by the reddish plants, suborbi-
cular pseudobulbs, and greenish yellow flowers
with red-brown markings. Chase did not encoun-
ter specimens below 500 m in Costa Rica, but a
specimen at SEL collected at less than 100 m (At-
wood 85-72) appears to be this species.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
123
Leochilus scriptus (Scheidw.) Rchb. f., Xenia
Orchid. 1:15. 1854. Cryptosanus scriptus
Scheidweiler, Allg. Gartenzeitung 11:101.
1843.
Plant a cespitose epiphyte 8-15 cm high; rhizomes
short, roots to 1 .5 mm in diameter. Pseudobulbs ovoid,
not strongly compressed, grooved, 2-5 cm long, 1-2.5
cm wide, subtended by 1-2 foliaceous sheaths, apex 1-
foliate. Leaves somewhat petiolate, the blade elliptic-
lanceolate, 4-12 cm long, 1-3 cm wide, apex acute. In-
florescences 1-2, 4-15(29) cm long, each a raceme or
panicle; ovary with pedicel about 1 cm long, subtended
by shorter, triangular bracts. Flowers fragrant, 1-6, te-
pals greenish yellow with reddish spots, lip yellow to
yellowish green with red to rose spots. Sepals oblong-
lanceolate, dissimilar, acute, 7-12 mm long, 3-5 mm
wide; dorsal free; lateral sepals to l/i connate. Petals
ovate, acute, 7-10(15) mm long, 5-7 mm wide. Lip ob-
ovate, emarginate, 6-12(14) mm long, 5-9 mm wide,
forming a rounded nectary at the base with the column.
Column 6-8 mm long, pollinia 2 on an elongate stipe
with viscidium. Capsules 3-4 cm long including the
long beak; pedicel 1-1.2 cm long.
Leochilus scriptus grows in moist to semideci-
duous forests at 0-600(1100) m. Flowering in
Costa Rica mostly December to April. Reported
from Mesoamerica to coastal tropical South
America, Cuba, and the Dominican Republic.
This species is easily distinguished from others
principally by its lower-elevation distribution and
in its large pseudobulbs. The flowers otherwise
resemble those of the other species.
The above measurements incorporate those
published by Chase in his revision of Leochilus.
Leochilus tricuspidatus (Rchb. f.) Kraenzl.,
Pflanzenr. IV. 50(Heft 80):297. 1922. Oncidium
tricuspidatum Rchb. f., Beitr. Orch.-K.C.Amer.
72. 1866. Figure 30D.
Plant epiphytic, cespitose, 8-12 cm high excluding
inflorescence. Rhizome short; roots 1-1.5 mm in diam-
eter. Pseudobulbs ovate to lanceolate, 1 (2)-foliate, com-
pressed, 1-1.8 cm long, 4-6 mm wide. Leaves subcor-
iaceous, elliptic-lanceolate, apically asymmetric, 5-9.5
cm long, 0.9-2 cm wide. Inflorescences paniculate, 1-
2 from the base of the pseudobulb, 8-35 cm high, with
5-15 flowers, lightly fractiflex, the scapes covered by
triangular, papery bracts; ovary with pedicel 5-1 1 mm
long. Flowers opening successively, greenish yellow
with brown spots, the lateral sepals with the spots coa-
lescing into a central line, 1.4-1.6 cm long and 9-11
mm broad. Sepals free, subequal; dorsal sepal elliptic-
lanceolate, concave, obtuse; lateral sepals elliptic-lan-
ceolate, subfalcate, acute, carinate, 5-8 mm long, 3-4
mm wide. Petals free, elliptic, obtuse, concave, 4-7 mm
long, 3-4 mm wide. Lip simple, oblanceolate, obtuse,
pubescent, with a small nectary at the base, 5-12 mm
long, 2.5-5 mm wide; blade with 2 erect projections at
the base and an erect callus with numerous small hairs,
the apex emarginate. Column semiterete, 4-8 mm long,
the apex denticulate, with 2 fleshy auricles beside the
stigma; pollinia 2, with elongate stipe and viscidium.
Capsule ellipsoid, strongly 3-carinate, 2-3 cm long,
pedicel 1-1.2 cm long.
This species is a twig epiphyte of evergreen
forests at 1200-2000 m. Flowering mostly Feb-
ruary to June and sporadically to October. Endem-
ic to Costa Rica and adjacent Chiriqui, Panama.
Leochilus tricuspidatus is easily distinguished
by the small pseudobulbs hidden by subtending
foliaceous bracts.
Lockhartia Hook.
(D. E. Mora de Retana)
REFERENCE — M. W. Chase, A reappraisal of the
Oncidioid orchids. Syst. Bot. 11:477-491. 1986.
Epiphytic, cespitose, psygmoid herb, stems un-
branched above. Pseudobulbs none. Leaves congested
and sheathing the stem, imbricate, remaining condupli-
cate, inarticulate, usually triangular in profile. Inflores-
cences 1-several racemes, lateral, of 1-several flowers.
Flowers usually attractive, usually yellow with red
marking or white, complex, especially the callus of the
lip. Sepals simple, ovate, free, concave, reflexed. Petals
simple, ovate. Lip simple to complexly lobed, the callus
often complex. Column short, usually winged, lacking
a tabula infrastigmatica; pollinia 2, usually obovate to
clavate, with stipe and small viscidium. Fruit a capsule.
An odd genus of 20-25 species native from
Mexico to South America but noticeably absent
from the Antilles. Six species are known from
Costa Rica.
The relationships of Lockhartia are unclear, but
they probably belong within the Oncidiinae. The
flowers superficially resemble those of Oncidium
but lack the tabula infrastigmatica. Plant habit is
similar to that of Psygmorchis Dodson & Dressier
but has a longer stem.
124
FIELDIANA: BOTANY
Key to the Species of Lockhartia
la. Lateral lobes present at the base of the lip 2
2a. Flowers < 1 cm tall L. micrantha
2b. Flowers > 1 cm tall 3
3a. Lip apex less than twice as broad as base of midlobe; floral bracts 6-10 mm wide
L. amoena
3b. Lip apex at least twice as broad as base of midlobe; floral bracts < 6 mm wide
L. oerstedii
Ib. Lateral lobes absent at the base of the lip, lip entire, 2-lobate or with lateral lobes near the middle
4
4a. Flowers white, lip suborbicular, callus orange, leaves slender and flexible . . . . L. hercodonta
4b. Flowers yellow, lip longer than wide; leaves stiff 5
5a. Flowers at least 1.2 cm tall, inflorescence compact with inconspicuous branches
L. pittieri
5b. Flowers ^ 1 cm tall; inflorescence spreading and diffuse, the branches conspicuous ....
. L. acuta
Lockhartia acuta (Lindl.) Rchb. f., Bot. Zeitung
(Berlin) 10:767. 1852. Fernandezia acuta
Lindl., Edwards's Bot. Reg. 21: t. 1806. 1836.
TYPE: Trinidad. Lockhartia pallida Rchb. f.,
Bonplandia 2:14. 1854. TYPE: Venezuela, Car-
abobo, Wagener (w, photo). Figure 32A.
Plant epiphytic, cespitose, to 40 cm; roots slender,
0.5-1 mm in diameter. Stems arcuate or pendulous, lat-
erally flattened, leafy, 25-40 cm long. Leaves disti-
chous, densely imbricate, somewhat fleshy, triangular in
profile, acute, 2-3.5 cm long, 3-9 mm deep. Inflores-
cence terminal or lateral near the apex, diffusely
branched, with few flowers opening in succession; ovary
and pedicel ca. 9 mm long, subtended by cordate bracts
1-2 mm long and 1-2 mm wide. Flowers cream-white,
small for the genus, 5-9 mm tall, 7-8 mm wide. Sepals
similar, rounded or obtuse, free, concave, 2-3 mm long,
2-3 mm wide. Petals similar to the sepals in form and
size. Lip simple to obscurely 3-lobate, subquadrate, with
reddish brown marks on the basal half, 5-6 mm long,
3-4 mm wide, the apex retuse and with somewhat ir-
regular lateral margins; lower l/i with a pale, papillose
callus, the apex with 2 rounded projections. Column
white with reddish brown spots at the base, with rounded
wings, the margins smooth, 1-2 mm high, 2-3 mm
wide; anther papillose; pollinia 2, pyriform with rela-
tively large stipe and viscidium.
Epiphytic in seasonally moist forests of the Pa-
cific drainage at 50-600 m. Flowering December
to March. Known from Costa Rica to Colombia,
Venezuela, Trinidad, and Tobago.
Lockhartia acuta is similar to L. micrantha
Rchb. f. but may be distinguished by the much
larger, usually pendulous stems and by the more
diffuse, branching inflorescence. Costa Rican ex-
amples lack the extreme acute lateral lobes of the
lip found in the type from Trinidad.
Lockhartia amoena Endres & Rchb. f., Gard.
Chron. 666. 1872. TYPE: Costa Rica, Endres
(w, photo). Lockhartia costaricensis Schltr., Re-
pert. Spec. Nov. Regni Veg. 3:81-82. 1906.
TYPE: Costa Rica, Los Palmares, Pittier 10592
(AMES). Lockhartia dipleura Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:69. 1923.
TYPE: Costa Rica, Werckle 102. Lockhartia
grandibractea Kraenzl., Pflanzenr. IV. 50 (Heft
83): 15. 1923. TYPE: Costa Rica, Endres (w,
photo). Lockhartia triangulabia Ames & C.
Schweinf., Sched. Orch. 8:80-81. 1924. TYPE:
Panama, Province of Chiriqui, Powell 362a
(AMES). Lockhartia amoena var. triangulabia
(Ames & C. Schweinf.) C. Schweinf. & P. H.
Allen, Bot. Mus. Leafl. 13:150. 1948. Figure
31A.
Plant epiphytic, cespitose, to 40 cm tall. Roots slen-
der, 0.5-1 mm in diameter. Stems erect, pendulous when
large, flattened. Leaves distichous, somewhat fleshy, tri-
angular in profile, acute, 1.3-3.5 cm long, 5-10 mm
deep. Inflorescence branched, terminal or lateral near
the apex, few- to many-flowered; ovary with pedicel 15
mm long, subtended by cordate, suborbicular bracts 4-
10 mm long and 6-10 mm wide. Flowers deep yellow
with reddish brown spots, of medium size for the genus,
1.3-1.9 cm long, 1.6-1.7 cm wide. Sepals similar, free,
ovate, apiculate, 5-7 mm long, 4-6 mm wide, the dorsal
lightly reflexed, the lateral strongly reflexed. Petals sim-
ilar, free, somewhat larger than the sepals, oblong to
elliptic, with undulate margins, 7-8 mm long, 4-5 mm
wide, obtuse. Lip 3-lobate, 5-9 mm long, 6-8 mm wide;
the lateral lobes linear, obtuse, incurved, 5-7 mm long,
1-3 mm wide; midlobe subquadrate, emarginate, with
the base more than Vi the width of the apex; disk with
callus of papillae from the base to ca. % the lip, emar-
ginate in front, column broadly winged, 2-3 mm high,
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
125
5-6 mm wide, the margins denticulate; anther papillose;
pollinia 2, pyriform; stipe and viscidium small.
Lockhartia amoena grows on larger branches
and tree trunks in premontane forests at 800-1400
m. Flowering throughout the year. Reported from
Mexico to Colombia.
Lockhartia amoena, as here interpreted, is a
most variable species, especially in the size of the
inflorescences, number of flowers, and midlobe of
the lip. Lockhartia dipleura Schltr. is based on a
plant with a nearly entire, not emarginate, mid-
lobe, but we find intermediates between plants
with entire and plants with deeply emarginate lips,
and both extreme forms have the same linear lat-
eral lobes and similar calli.
Lockhartia hercodonta Rchb. f. ex Kraenzl.,
Pflanzenr. IV. 50(Heft 83):8, t. 2A. 1923.
TYPE: Costa Rica, Endres (w, photo). Figure
31B.
Plant cespitose, epiphytic, 10-45 cm high. Rhizome
short, with slender roots < 1 mm in diameter. Stems
erect or pendulous, compressed, foliaceous. Leaves tri-
angular in profile, thin and flexible, imbricate, 2.8-4.3
cm long, lateral width 5-10 mm deep; apex usually in-
curved, acute to somewhat acuminate. Inflorescence ap-
parently terminal or subterminal, usually branched, suc-
cessively flowered; ovary and pedicel 5-6 mm long,
subtended by variable, ovate-lanceolate, acuminate
bracts 5-7 mm long. Flowers small, 8-10 mm in di-
ameter, white, the lip with orange callus. Sepals con-
cave, ovate, acute, 3-4 mm long, 2-3 mm wide. Petals
lightly rounded, concave, 4-5 mm long, 3-4 mm wide.
Lip simple, suborbicular, 4-5 mm long, 6-7 mm wide;
with an elevated, cuplike, papillose, apically bidentate
callus. Column short, broad, 2-3 mm long, with cren-
ulate wings on the upper l/r, pollinia 2 with stipe and
minute viscidium. Capsule globose to ellipsoid, 8 mm
long, with pedicel ca. 5 cm long.
This species grows in premontane and lower
montane rain forests at 900-2000 m, usually in
deep shade. Flowering August to March and spo-
radically in June and July. Reported from Guate-
mala to Panama and Colombia.
Lockhartia hercodonta is easily distinguished
from others, especially the cream-white-flowered
L. acuta (Lindl.) Rchb. f., by the soft, thin leaves,
suborbicular lip, and orange callus. Vegetatively
it is similar to L. pittieri Schltr., a low-elevation
species with yellow flowers.
Lockhartia micrantha Rchb. f., Bot. Zeitung
(Berlin) 10:768. 1852. TYPE: Panama(?), Ver-
agua, Hinds (w, photo). Lockhartia chiriquensis
Schltr., Repert. Spec. Nov. Regni Veg. 12:215.
1913. TYPE: Panama, Chiriqui near San Felix,
Pittier (AMES, drawing). Lockhartia lankesteri
Ames, Sched. Orch. 5:36. 1923. TYPE: Costa
Rica, San Carlos, Lankester 448 (holotype:
AMES). Figure 3 1C.
Plant epiphytic, cespitose, compact, 12-30 cm high.
Roots slender, 0.5-1 mm in diameter. Stems erect or
pendulous, laterally flattened, densely foliaceous.
Leaves distichous, densely imbricate, triangular in pro-
file, somewhat fleshy, acute, 0.8-2 cm long, 5-8 mm
wide. Inflorescence short, terminal or lateral near the
apex, of few flowers; ovary with pedicel 8-9 mm long,
subtended by cordate, acuminate bracts, ca. 6 mm long.
Flowers small for the genus, 7-9 mm long and about as
wide, clear or pale yellow, column pale yellow with
small reddish brown spots at base, callus with small,
brown spots. Sepals concave, similar, ovate, apiculate,
free, 5-6 mm long, 4-5 mm wide. Petals elliptic, 5-6
mm long, 2-3 mm wide; margins lightly undulate. Lip
3-lobate, 5-6 mm long, 6-7 mm wide; lateral lobes lin-
ear, ascending, obtuse; midlobe, cuneate, emarginate
with a small notch on each side; callus suborbicular, ver-
rucose, broad at the base, narrow at the apex. Column
2 mm long with broad wings; anther cucullate, papillose;
pollinia 2, pyriform, with relatively large stipe. Cap-
sules obovoid to ellipsoid, 6-8 mm long, pedicel ca. 1-
1.2 cm long.
This species generally grows in humid open
forests at 10-900 m in bright light. Flowering No-
vember to April. Broadly distributed from Nica-
ragua to Panama; also reported from northwestern
South America.
Lockhartia micrantha is characterized by the
small, pale yellow flowers and linear lobes at the
base of the lip.
Lockhartia oerstedii Rchb. f., Bot. Zeitung (Ber-
lin) 10:767-768. 1852. TYPE: Costa Rica, Bar-
ba, Oersted s.n. (K, photo seen). L. verrucosa
Lindl. ex Rchb. f., Hamburger Garten-Blumen-
zeitung 15:53-54. 1859. TYPE: Stange? (K,
photo). Lockhartia lamellosa Rchb. f. Ham-
burger Garten-Blumenzeitung 21:300. 1865.
TYPE: Mexico, Stange. Fernandezia robusta
Bateman, Bot. Mag. t. 5592. 1866. TYPE: Gua-
temala, Skinner s.n. Lockhartia robusta (Bate-
man) Schltr., Repert. Spec. Nov. Regni Veg. 3:
82. 1906. Figure 31D.
Plant epiphytic, cespitose, sometimes densely so.
Roots slender, 0.5-1 mm in diameter. Stems erect or
arcuate, (12)25-40 cm long. Leaves distichous, imbri-
cate, triangular in profile, somewhat fleshy, acute, about
1-3 cm long, 4-8 mm deep. Inflorescence of few suc-
cessively borne flowers, terminal or lateral near the stem
apex; ovary with pedicel 1-1.5 cm long, subtended by
126
FIELDIANA: BOTANY
cordate bracts about 5 mm long, 4 mm wide. Flowers
bright yellow with reddish brown spots, rather large for
the genus, 2-2.5 cm high and 1-1.5 cm wide. Sepals
similar, free, elliptic, obtuse, reflexed, 5-9 mm long, 6-
7 mm wide. Petals ovate, larger than the sepals, the
margins undulate, 8-1 1 mm long, 5-7 mm wide. Lip 3-
lobate, the margins undulate, 1.3-1.6 cm long, 1.2-1.5
cm wide; basal lateral lobes spatulate, obtuse, incurved,
7-9 mm long, 2-4 mm wide; midlobe 2-lobate, with
narrow base less than Vi the width of the apex, the disk
with a complex callus centrally of 7 rows of teeth, the
central row with smaller teeth. Column 2-3 mm long,
4-5 mm wide, with very broad wings with dentate mar-
gins; anther papillose; pollinia 2, pyriform, with minute
stipe and viscidium. Capsules obovoid, ca. 1.2-1.5 cm
long, pedicel ca. 1.4 cm long.
Lockhartia oerstedii is a midcanopy epiphyte of
evergreen premontane and lower montane forests
at 1100-1800 m. Flowering December to May,
August to September, and perhaps throughout the
year. Reported from Mexico to Panama and Co-
lombia.
This species has among the largest and most
attractive flowers of the genus. It is distinguished
from the similar L. amoena Endr. & Rchb. f. by
the more slender shoots, the more compact inflo-
rescences, and the more complex callus with 7
rows of teeth.
Lockhartia pittieri Schltr., Repert. Spec. Nov.
Regni Veg. 12:216. 1913. TYPE: Panama, Pit-
tier (drawing of type: AMES). Lockhartia var-
iabilis Ames & C. Schweinf., Sched. Orch. 8:
81-82. 1925. TYPE: Panama, near Frijoles,
Powell 355 (holotype: AMES). Lockhartia Inte-
gra Ames & C. Schweinf., Sched. Orch. 10:
108. 1930. TYPE: Costa Rica, Guanacaste, El
Silencio, near Tilaran, Standley & Valeria
44675 (holotype: AMES). Figure 32B.
Plant an erect or pendent epiphyte 15-20 cm tall;
rhizome short, roots < 1 mm in diameter. Leaves dense-
ly imbricate, triangular in profile, 2-3.5 cm long, 2.5-6
cm wide, acute. Inflorescence short, 1 -apparently few
flowers, lateral on the upper Vi of the stem; ovary and
pedicel 9-10 mm long, subtended by an ovate, cordate,
acuminate bract 3.5 mm long and 2.5 mm wide. Flowers
yellow with orange-brown callus, ca. 1 .4 cm high, 1 cm
wide. Sepals concave, elliptic-ovate, reflexed at the
base, 5-6 mm long, 4-5 mm wide; apex dorsally cari-
nate, acute. Petals ovate-lanceolate, dorsally carinate, 6-
8 mm long, 5-6 mm wide, acute. Lip convex, generally
obovate to subquadrate, 8-9 mm long, 5-6 mm wide;
apex 2-lobate; callus ovate, concave, margin lightly pi-
lose, with 3 fleshy keels at the base, the central larger
than the lateral. Column stout with broad, denticulate
wings for the entire length, 3 mm long; pollinia 2 with
caudicles, obovoid, with stipe and small viscidium. Cap-
sule suborbicular to ellipsoid, 8-10 mm long, pedicel
ca. 1 cm long.
Epiphytic in midcanopies in humid tropical for-
ests at 30-800 m. Flowering November to May.
Belize to Panama; also reported from Ecuador.
Plant is similar to that of L. hercodonta Rchb.
f. ex Kraenzl. but lives at lower elevations, and
the flowers are larger and yellow.
Lockhartia pittieri Schltr. is based on a type
from Panama with a 4-lobate lip, a feature that
may be an artifact. For the moment we are fol-
lowing others in accepting this as the oldest avail-
able synonym for L. Integra Ames & C.
Schweinf., which is based on Costa Rican mate-
rial with a 2-lobate lip.
Macroclinium Barb. Rodr.
(F. Pupulin and D. E. Mora de Retana)
REFERENCE — F. Pupulin, New and critical Ma-
croclinium (Orchidaceae) from Central America.
Lindleyana 11:135-140. 1996.
Plant perennial, epiphytic, cespitose, sympodial with
pseudobulbs or submonopodial without. Pseudobulbs
when present more or less inconspicuous, ovoid or
rounded, compressed, apically 1 -foliate, concealed by
leaf sheaths. Leaves usually 3-7 (34) per shoot, con-
duplicate, laterally flattened, the basal ones articulate
with imbricate sheaths often provided with hyaline mar-
gins; blades linear-lanceolate to subfalcate-lanceolate.
Inflorescence lateral, subumbellate or elongate with
flowers well separated, often branched, 2-many-flow-
ered, often successive, arising from the axils of the upper
leaf sheaths. Flowers small, hyaline. Sepals free or
shortly connate, petals similar to the sepals, spreading.
Lip free or adnate to the column base, with or without
calli or lateral lobules near the base. Column slender,
linear-clavate, sometimes abruptly and dorsally reflexed
at the apex, sometimes slightly adnate to the lip, foot
lacking; pollinia 2, strongly compressed, supported on a
slender, triangular stipe. Fruit a capsule.
A genus of about 38 species distributed from
southern Mexico to Brazil, generally as twig ep-
iphytes on the upper or the outer portion of the
canopy. Fifteen species are reported from Central
America, 10 of which occur in Costa Rica. Phy-
logenetically, Macroclinium has several derived
character states within the Rodriguezia clade
(Chase & Palmer, 1992), a group of orchids spe-
cialized to occupy the more xeric habitat of twigs
in Neotropical canopies, including lonopsis, Tri-
zeuxis, Notylia, Warmingia, and Macradenia.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
127
Key to the Species of Macroclinium
la. Inflorescence a raceme with flowers well separated 2
2a. Callus of the lip papillose-hirsute M robustum
2b. Callus of the lip glabrous 3
3a. Lateral sepals free; lamina of the lip cordate-sagittate M. ramonense
3b. Lateral sepals connate; lamina of the lip ovate M. confertum
Ib. Inflorescence subumbellate, the flowers close together 4
4a. Lip without lateral, retrorse auricles M. lineare
4b. Lip with two lateral, retrorse auricles at base 5
5a. Lip ecallose M. doderoi
5b. Lip with a basal callus between auricles 6
6a. Column with abruptly and dorsally reflexed apex 7
7a. Lip free from the column; callus of the lip with transverse hairs
M. glicensteinii
7b. Lip adnate to the base of the column; callus glabrous 8
8a. Lamina of lip cordate, with a single callus between auricles M. allenorum
8b. Lamina of the lip hastate, with 2 membranous calli between auricles
M. generalense
6b. Column without such a reflexed apex 9
9a. Pseudobulbs > 1 cm long M. cordesii
9b. Pseudobulbs < 1 cm long M. paniculatum
Macroclinium allenorum Dressier & Pupulin,
Lindleyana 11:34-36. 1996. TYPE: Costa Rica,
Puntarenas, vicinity of Palmar Norte, Rio Ter-
raba, Allen 5244 (holotype: us; isotypes: F, SEL).
Plant epiphytic, cespitose, small. Rhizome short,
roots filiform. Pseudobulbs inconspicuous, compressed-
ellipsoid, 4-8 mm long, closely invested by 3-5 con-
duplicate, leaf -bearing sheaths, apex 1 -foliate. Leaves
conduplicate, laterally flattened, lanceolate to narrowly
elliptic-lanceolate, 12-25 mm long, 2.8-5.8 mm wide;
the basal leaves articulate to the broad sheaths with hy-
aline margins. Inflorescence a simple subumbellate ra-
ceme, becoming paniculate, 1.5-11 cm long, provided
with 2-3 conspicuous, concave, acute bracts 8-11 mm
long, 5 mm wide; ovary with pedicel subclavate, sub-
tended by a lanceolate floral bract, shorter than the ped-
icellate ovary. Flowers small; sepals white, hyaline; pet-
als white, spotted with rose-red; lip lavender. Sepals dis-
similar; dorsal lanceolate-elliptic, acuminate, cucullate,
5.5-7 mm long, 0.8-1.3 mm wide; lateral sepals oblong-
lanceolate, acuminate, free, dorsally carinate, 6-8 mm
long, 0.8-1 mm wide. Petals obliquely lanceolate, acu-
minate, 4.7-6 mm long, 0.8 mm wide. Lip with a linear
claw ca. 2 mm long, adnate to base of column for more
than l/3 of claw; blade 5-5.5 mm long, triangular-cordate,
with serrulate margins, acuminate to mucronate, the
midlobe unguiculate, the isthmus subequal to basal claw
with 2 short retrorse auricles near the base, with thick-
ened, glabrous callus between auricles. Column slender,
ca. 5 mm long, abruptly dorsally reflexed at apex; anther
triangular, cucullate; pollinia 2, compressed, on a long,
triangular stipe; viscidium elliptic.
Epiphytic in moist premontane forests at 0-400
m. Flowering December to May. Costa Rica; sup-
posedly endemic in the region of Palmar Norte to
Golfito.
Macroclinium allenorum may be distinguished
from M. glicensteinii J. T. Atwood and M. gener-
alense Pupulin by the lip with long claw partially
adnate to the base of the column and by the cor-
date midlobe with serrulate margins. In addition,
both the transverse hairs on the callus of M. gli-
censteinii and the two membranous calli of M.
generalense are absent in M. allenorum.
Macroclinium confertum Pupulin, Lindleyana
11:138-140. 1996. TYPE: Costa Rica, San
Jose, Dota, San Marcos, road to San Joaquin,
Pupulin 204 (holotype: us:). Figure 32C.
Plant epiphytic, small, cespitose, sympodial. Rhi-
zome short, roots filiform. Pseudobulbs inconspicuous,
ovate, somewhat compressed, 6-7 mm long, 4.5-5 mm
wide, covered at the base by 3-7 leaf-bearing sheaths,
apex 1 -foliate. Leaves conduplicate, laterally flattened,
linear-lanceolate to subfalcate-lanceolate, to 5.5 cm long,
6 mm wide, acute; the basal leaves articulate to sheaths
with hyaline margins. Inflorescence a pendent raceme
with many flowers (11-13), to 8 cm long, provided at
the base with 2-3 lanceolate, subulate bracts ca. 4 mm
long; ovary with pedicel subclavate, to 7 mm long, sub-
tended by a lanceolate, acuminate, floral bract ca. 3 mm
long, 1.5 mm wide. Flowers rather large for the plant;
sepals white, hyaline; petals pale rose to lavender spot-
ted with purple; lip lavender. Sepals dissimilar; dorsal
concave, linear-lanceolate, acuminate, to 16 mm long,
128
FIELDIANA: BOTANY
2.2 mm wide; lateral sepals shortly connate for ca. 1
mm, somewhat concave toward the base, narrowly lin-
ear-lanceolate, ca. 1.6 cm long, 1.3 mm wide, acuminate
to setaceous. Petals obliquely lanceolate, to 1.2 cm long,
1.9 mm wide, acuminate to setaceous. Lip with linear
claw 1 mm long free from the column; blade 9.7-10.1
mm long, 2 mm wide at the middle, the base narrowly
cuneate and with a pair of triangular, deflexed, and
somewhat twisted auricles; apex setaceous; margins
crisped; callus between auricles glabrous, obtrullate.
Column slender, clavate, ca. 4 mm long, with cuneate,
acute apex; anther cucullate; pollinia 2, strongly com-
pressed; stipe hyaline, elongate, triangular; viscidium el-
liptic, brown.
Epiphytic on lower twigs in wet premontane
forests at 1300-1500 m. Flowering at least May
to June. Endemic to valleys of the Dota region in
central Costa Rica.
The shortly connate lateral sepals, the blade of
the lip with ovate central portion, and the obtrul-
late, glabrous callus easily distinguish M. confer-
tum from its closest allied species, M. ramonense
(Schltr.) Dodson and M. robustum Pupulin &
Mora-Retana.
Macroclinium cordesii (L. O. Williams) Dodson,
Icon PL Trop. 10: t. 938. 1984. Notylia cordesii
L. O. Williams, Ann. Missouri Bot. Gard. 26:
286. 1939. TYPE: Panama, Bocas del Toro,
Mosquito Hill, Cordes ex Woodson, Allen, &
Seibert 1932 (MO, not seen). Figure 33A.
Plant epiphytic, cespitose, small. Rhizomes short,
roots filiform, < 1 mm in diameter. Pseudobulbs com-
pressed, obovoid-ellipsoid, 1-1.5 cm long, 4 mm wide,
concealed at the base by ca. 4 leaf sheaths, apex 1 -fo-
liate. Leaves equitant, sessile, linear-lanceolate to ovate-
lanceolate, 4-7 cm long, 3-5 mm wide, acute to acu-
minate. Inflorescence a simple subumbellate raceme or
panicle with subumbellate branches, each subumbel with
5-6 flowers; peduncle slender, 4.5-6 cm long, with sev-
eral triangular, funnelform bracts 1.5-2 mm long; ovary
with pedicel to 9 mm long, subtended by a scarious,
lanceolate, spreading, acute to acuminate floral bract
1.5-2 mm long. Flowers small; sepals white, hyaline;
petals lavender tinged with rose-purple near the base; lip
and column purple. Sepals similar, linear-lanceolate,
acuminate; dorsal ca. 10 mm long, 1.25 mm wide; lateral
sepals slightly oblique, 1.2-1.3 cm long, 1 mm wide.
Petals similar to the dorsal sepal but slightly narrower,
ovate-lanceolate, abruptly attenuate toward the apex, ca.
1 cm long, 1 mm wide. Lip free from the column, with
claw 4 mm long; blade with a basal isthmus, provided
at the base with an obscurely papillose thickening and
2 retrorse auricles, then abruptly hastate, acuminate, to
5 mm long, 2 mm wide; lateral lobes somewhat re-
curved, serrate; apex strongly acuminate. Column slen-
der, ca. 4 mm long with prominent foot; anther subglo-
bose; pollinia 2, pyriform, strongly compressed, long tri-
angular stipe.
Macroclinium cordesii is a rare species from
the coastal plains of the Province of Limon. Flow-
ering late August to October. Costa Rica and Pan-
ama.
Among Costa Rican Macroclinium species (es-
pecially M. paniculatum (Ames & C. Schweinf.)
Dodson) with subcapitate inflorescences, M. cor-
desii may be distinguished by the straight, not
abruptly reflexed, column and the conspicuous
pseudobulbs.
Macroclinium doderoi Mora & Pupulin, Selby-
ana 18:7. 1997. TYPE: Costa Rica, Province of
Cartago, Turrialba, Cano Seco Torito, Dressier
et al. s.n. (holotype: usj; isotype: usi).
Plant epiphytic, pendent. Rhizome short; roots fili-
form, ca. 1 mm in diameter. Pseudobulbs inconspicu-
ous, elliptic-obovate, ca. 3.5 mm long, 2 mm wide, con-
cealed by 3-5 leaf sheaths, apex 1 -foliate. Leaves con-
duplicate, compressed, lanceolate to narrowly elliptic-
lanceolate, 1.2-1.8 cm long, 2 mm wide, acute,
articulate to the imbricate sheaths with hyaline, scarious
margins. Inflorescence a simple subumbellate raceme or
panicle with many subumbellate branches produced suc-
cessively from the last bud below the rachis of the ter-
minal flower cluster, lax, to ca. 4 cm long, with 5-6
flowers per subumbel; ovary with pedicel subclavate, ca.
7 mm long, subtended by a shorter, triangular, acumi-
nate, floral bract. Flowers large for the plant, tepals
white, hyaline, petals with a purple blotch near the base;
column and lip lavender to purple. Sepals similar; dorsal
lanceolate, ca. 9 mm long, 1.6 mm wide, attenuate; lat-
eral sepals narrowly lanceolate, ca. 1.1 cm long, 1 . 1 mm
wide, long-attenuate. Petals ovate-lanceolate, falcate, ca.
9 mm long, 1.3 mm wide, attenuate. Lip free, inserted
at a very narrow angle with the column, with short claw
0.9 mm long; blade slightly incurved toward the column,
abruptly sagittate, ca. 7.5 mm long, 2 mm wide at mid-
point, with 2 short, twisted lobes near the base; margins
erose-dentate; callus none. Column terete, slender, 5.5
mm long, with abruptly reflexed apex; anther cucullate,
narrowly ovate-triangular; pollinia 2, ovate-pyriform,
strongly compressed, with long triangular stipe with lat-
eral projections apically; viscidium elliptic, brown.
Epiphytic and endemic in moist tropical forests
of the Central Valley of Costa Rica. Flowering
late June through August.
Macroclinium doderoi may be easily distin-
guished from its Costa Rican relatives by a unique
combination of characters including the subcapi-
tate inflorescence, the lip entirely free from the
column, the very short claw, and the absence of
a callus between the lateral lobes of the lip.
Macroclinium generalense Pupulin, Lindleyana
11:136-138. 1996. TYPE: Costa Rica, San
ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS
129
Jose, Perez Zeledon, Alto de San Juan, Pupulin
24 (holotype: us;). Figure 32D.
Plant epiphytic, small, cespitose. Rhizome short;
roots filiform, glabrous. Pseudobulbs inconspicuous,
compressed, ellipsoid, to 5 mm long, 4 mm wide, con-
cealed by 3-5 leaf sheaths, apex 1 -foliate. Leaves con-
duplicate, laterally flattened, lanceolate to narrowly el-
liptic-lanceolate, to 4 cm long, 1 cm wide, acute, the
basal leaves articulate to imbricate sheaths provided with
broad, hyaline, scarious margins. Inflorescence subum-
bellate, with lateral, subumbellate branches produced
consecutively from the last bract of the inflorescence
below the rachis, few-flowered, to 1 1 cm long, provided
at the base with 3-5 conspicuous, ovate, concave, acute
bracts 8-1 1 mm long, 5 mm wide; ovary with pedicel
subclavate, to 6 mm long, subtended by a lanceolate,
acuminate floral bract to 4 mm long. Flowers small,
hyaline, sepals white, petals pale rose spotted with pur-
ple, lip lavender. Sepals similar; dorsal lanceolate -ellip-
tic, cucullate, to 1.2 cm long, 3.5-4 mm wide, acumi-
nate; lateral sepals free, dorsally carinate, oblong-lan-
ceolate, ca. 1.1 cm long, 3-3.5 mm wide, acuminate.
Petals obliquely lanceolate, acuminate, to 1.2 cm long,
3.5 mm wide. Lip shortly clawed, sagittate, to 1.1 cm
long, acuminate; claw linear, ca. 2 mm long, with more
than i/3 adnate to column base; blade with 2 short, re-
trorse, twisted auricles at the base and 2 membranous,
slightly thickened, glabrous calli between the auricles;
basal portion of the blade narrowly linear cuneate, grad-
ually expanded into the distal, triangular-rhombic por-
tion, with irregular margins, apex acuminate. Column
slender, abruptly dorsally reflexed at the apex; anther
triangular, cucullate; pollinia 2, pyriform, strongly com-
pressed laterally on a long triangular, hyaline stipe; vis-
cidium elliptic, brown.
Epiphytic and endemic in the northern part of
the Valle del General in Costa Rica. Flowering
January through April.
The triangular-rhombic blade of the lip with the
short claw adnate to the column and the presence
of two membranous, glabrous calli easily distin-
guish M. generalense from its close allies, es-
pecially M. glicensteinii J. T. Atwood.
Macroclinium glicensteinii J. T. Atwood, Sel-
byana 10:60. 1987. TYPE: Costa Rica, Vara
Blanca, QIC 7473 (holotype: SEL). Holotype
originally collected by L. Glicenstein.
Plant epiphytic, cespitose, small. Rhizome short;
roots filiform, to 1 mm in diameter. Pseudobulbs ovate,
compressed, inconspicuous, to 4 mm long, concealed by
the base of leaf sheaths, apex 1 -foliate. Leaves sessile,
conduplicate, laterally flattened, narrowly elliptic, 2-4
cm long, 3 mm wide; sheaths with conspicuous hyaline
margins. Inflorescence subumbellate, sometimes with
more than a subumbel developing from the same pedun-
cle; peduncle slender, with lanceolate subulate bracts
each sometimes developing a subumbel basipetally; ova-
ry with pedicel 1 cm long, subtended by a lanceolate
floral bract 3-4 mm long. Flowers small, pale lavender
with spotting mostly on the petals. Sepals similar, sub-
conduplicate, narrowly elliptic-attenuate, 1.1-1.4 cm
long, 1.5 mm wide. Petals lanceolate-attenuate, 8-12
mm long, to 1.5 mm wide. Lip free from the column,
with a short claw, 3.5 mm long, presenting 2 auricles
and 2 membranous calli apically, below which are nu-
merous papillae and between which are transverse hairs;
blade of the lip hastate, 5-7 mm long, 2-2.5 mm wide,
acuminate, with irregularly crenulate margins, the apex
strongly acuminate. Column slender, the apex abruptly
dorsally reflexed, 7-8 mm long; anther cap ovate-subu-
late; pollinia 2, pyriform, strongly compressed, with
long cuneate stipe.
Originally described from a cultivated speci-
men, M. glicensteinii is endemic to Costa Rica,
where it probably lives in moist montane forests
in the region of Dota. Flowering in cultivation
(Connecticut) in December.
The presence of a few long, transverse hairs on
the callus at the base of the blade of the lip and
the short claw completely free from the column
are distinct features for field recognition. This
species is known only by the type collection.
Macroclinium lineare (Ames & C. Schweinf.)
Dodson, Icon. PI. Trop. 10: t. 938. 1984. No-
tylia linearis Ames & C. Schweinf., Sched.
Orch. 8:72. 1925. TYPE: Costa Rica, La Fuen-
te, East Turrialba, Alfaro s.n. (holotype: us).
Plant epiphytic, cespitose, small. Roots filiform.
Pseudobulbs elliptic, to 7 mm long, 3 mm wide. Leaves
conduplicate, spreading, laterally flattened, semielliptic
to linear-oblong, sharply acute, sessile, 1-3 cm long,
2.5-4.5 mm wide; imbricate sheaths with broad, scari-
ous, hyaline margins. Inflorescence subumbellate, se-
miglobose; peduncle slender, flexuous, suberect, to 4 cm
long, with 3 funnelform, acuminate bracts; ovary with
pedicel subclavate, 7 mm long, subtended by a narrowly
lanceolate, acuminate, concave, spreading, floral bract
1.8-2 mm long. Flowers large for the plants; sepals
white, hyaline; petals tinged with rose-purple; lip purple.
Sepals similar; dorsal strongly concave, narrowly lan-
ceolate, dorsally conspicuously carinate, ca. 1 cm long,
1.8-2 mm wide, apex caudate-acuminate; lateral sepals
free or very shortly connate forming a saccate base, lan-
ceolate-linear, 1.2-1.3 cm long, 1 mm wide; apex long
acuminate and conduplicate. Petals similar to the dorsal
sepal, 9-10 mm long, ca. 1.5 mm wide, caudate. Lip
free from the column, linear, contracted near the base,
tapering to the carinate tip, 5-9 mm long, ca. 1 mm wide
above the middle, shortly acuminate, with a clump of
low, approximate papillae near the base. Column slen-
der, ca. 3 mm long; pollinia 2, pyriform, strongly com-
pressed, with long cuneate stipe.
Epiphytic and rare in wet premontane forests at
the typical locality near Turrialba, along the At-
130
FIELDIANA: BOTANY
lantic slopes of Parque Nacional Braulio Carrillo,
and in northern Valle del General at 1100-1200
m. Flowering at least August and September. Cos-
ta Rica, Panama, Colombia, and Ecuador.
Macroclinium lineare is unmistakable for its
linear lip lacking auricles and for the clump of
slender papillae at the base.
Macroclinium paniculatum (Ames & C.
Schweinf.) Dodson, Icon. PI. Trop. 10: t. 938.
1984. Notylia paniculata Ames & C. Schweinf.,
Sched. Orch. 10:102. 1930. TYPE: Costa Rica,
Cartage, vicinity of Pejivalle, Standley & Val-
eria 46858 (holotype: AMES).
Plant epiphytic, cespitose, small. Rhizome short;
roots filiform, < 1 mm in diameter. Pseudobulbs incon-
spicuous, suborbicular, compressed, sulcate, 7 mm long,
5 mm wide; concealed by 4 scarious, leaf sheaths; apex
1 -foliate. Leaves densely rugose, conduplicate, laterally
flattened, narrowly lanceolate to narrowly elliptic-lan-
ceolate, 2.5-3.5 cm long, 3.5-5 mm wide, acute. Inflo-
rescence subumbellate, subequaling or surpassing the
leaves, commonly with successive short branches, few-
flowered in each branch; floral bracts scarious, lanceo-
late, spreading, 3-3.5 mm long, much shorter than the
ovary with pedicel 6 mm long. Flowers large for the
plant; sepals white, hyaline; petals pinkish white tinged
with rose-purple; lip purple. Sepals dissimilar; dorsal
ovate-lanceolate, cucullate, 6-7 mm long, 1.2-1.6 mm
wide, shortly acuminate; lateral sepals nearly Vi connate,
each oblong-lanceolate, dorsally carinate near the apex,
ca. 9 mm long, 1.2-1.3 mm wide, mucronate. Petals
ovate-lanceolate, 6-7 mm long, 1.8-2 mm wide, acu-
minate. Lip free from the column, with a long claw
slightly dilated and thickened near the blade; the blade
sagittate at the base, 3 mm long, 2 mm wide, the upper
Vi triangular-cordate, apex sharply mucronate; margins
irregularly crenate; callus consisting of a central, longi-
tudinal fleshy keel and a pair of fleshy auricles at the
base of the blade. Column slender below, dilated above,
2.3 mm long; anther subglobose; pollinia 2, pyriform,
strongly compressed, with long stipe.
Epiphytic and rare in premontane wet forests at
600-800 m. Flowering at least February (type)
and April. Nicaragua to Costa Rica.
The sagittate lip with callus consisting of a cen-
tral keel and fleshy auricles at the apex of the claw
and the cordate, mucronate anterior portion of the
blade are useful characters for field recognition.
Macroclinium ramonense (Schltr.) Dodson,
Icon. PL Trop. 10: t. 939. 1984. Notylia ramo-
nensis Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 19: 250-251. 1923. TYPE: Costa Rica,
Santiago de San Ramon, Brenes 91 (isotype:
CR). Figure 33B.
Plant epiphytic, small, cespitose, pendent. Rhi-
zome short; roots filiform, < 1 mm in diameter. Pseu-
dobulbs obovate-ellipsoid, compressed, ca. 1.5 cm
long, 5-6 mm wide, protected at the base by 2-5 con-
duplicate leaf-bearing sheaths; apex 1 -foliate. Leaves
rugulose, conduplicate, laterally flattened, ligulate-
lanceolate, subfalcate, acute, to 4.5 cm long, 4-8 mm
wide; basal leaves articulate to sheaths provided with
conspicuously hyaline, scarious margins. Inflores-
cence a pendent raceme, with many flowers (to 30),
to 13 cm long, provided with triangular, acute bracts
ca. 1.5 cm long; ovary with pedicel linear-subclavate,
ca. 8 mm long, subtended by a lanceolate, acuminate,
spreading floral bract 3 mm long. Flowers large for
the genus, sepals white, petals pale lavender with pur-
ple blotches, lip lavender. Sepals subequal; dorsal lin-
ear-lanceolate, concave, acuminate, 1.7 cm long, 2.1-
2.5 mm wide; lateral sepals obliquely linear-lanceo-
late, slightly concave, acute, 1.6-1.7 cm long, ca. 2
mm wide. Petals obliquely linear-lanceolate, long-at-
tenuate, ca. 1.3 cm long, 1.9 mm wide. Lip with a
linear, terete, slender claw 1.8 mm long, shortly ad-
nate to the column; blade 6.5 mm long, 3.7 mm wide
at the middle, the base with a pair of short, rounded,
twisted auricles, then abruptly anchor-shaped; apex
sharply attenuate; margins erose; callus between the
auricles glabrous, narrowly elliptic. Column slender,
terete, ca. 3 mm long; anther ovate-cucullate, trun-
cate; pollinia 2, subtriangular, strongly compressed on
a long, triangular, attenuate stipe; viscidium elliptic,
brown.
Native to moist evergreen forests at 950(type)-
1500 m. Flowering at least August and September.
Costa Rica and western Panama.
Among Costa Rican species of Macrocli-
nium, M. ramonense has the widest distribution,
ranging from Monteverde southward along the
Cordillera de Tilaran to southern and western
drainage of Cordillera Central just to the west-
ern slopes of Cerro Vueltas of the Dota region.
Among the species with elongate, rather than
condensed, racemes, it is easily distinguished
by the anchor-shaped blade of the lip and the
narrowly elliptic callus. The long claw of the
lip and prominent, central callus readily distin-
guish this from M. bicolor, with which it has
been confused.
Macroclinium robustum Pupulin & Mora, Sel-
byana 18:7-10. 1997. TYPE: Costa Rica, Prov-
ince of San Jose, Alajuelita, Pupulin & Flores
315 (holotype: usj; isotype: usj).
Plant epiphytic, cespitose, pendent. Rhizome short;
roots filiform, ca. 1 mm in diameter. Pseudobulbs
ovate, to 8.5 mm long, 5 mm wide, concealed at the
base by 2-5 leaf sheaths with hyaline, scarious mar-
gins. Leaves rugose, compressed, 5.3-6.2 cm long,
5-7 mm wide, acute, the basal leaves articulate with
their sheaths. Inflorescence a lax raceme with 11-17
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
131
flowers, to 8.6 cm long, the base provided with 2 tri-
angular, acute bracts 8 mm long; ovary with pedicel
linear-subclavate, ca. 8 mm long, subtended by a tri-
angular, acuminate, floral bract ca. 4.5 mm long.
Flowers rather large for the genus; sepals greenish
white, petals lavender with purple blotches, lip lav-
ender. Sepals similar; dorsal concave, lanceolate, acu-
minate, dorsally carinate, 1.2-1.3 cm long, 3-3.5 mm
wide; lateral sepals slightly concave, linear-lanceo-
late, 1.4-1.5 cm long, 1.8 mm wide, acute. Petals
sigmoid, ovate-lanceolate, ca. 1.2 cm long, 1.5 mm
wide, long-attenuate. Lip shortly adnate to the col-
umn; claw linear, terete, 3.2 mm long; blade inflexed
toward the column 6 mm long, 2.6 mm wide, with 2
short, subquadrate, twisted lobes near the base,
abruptly sagittate-cordate at the middle, margins
erose; disk verruculose, with a papillose-hirsute cal-
lus. Column short, terete, slender at the base, 3 mm
long, thickened abruptly at the reflexed apex; anther
ovate-cucullate, apically mucronate; pollinia 2 ovate,
strongly compressed; stipe triangular-elongate; visci-
dium elliptic, yellow.
Epiphytic and endemic in montane cloud for-
ests of Costa Rica at 1850 m. Flowering late July
and August.
In the type locality, this species was found
growing exclusively on the introduced cypress
Cupressus lusitanicus. Macroclinium robustum is
the highest-growing species in Costa Rica and is
unique for having both elongate inflorescences
and flowers with papillose-hirsute calli. The long
and stout claw easily distinguishes it from its clos-
est relative, M. ramonense (Schltr.) Dodson.
Mesospinidium Rchb. f.
(D. E. Mora de Retana)
REFERENCE — N. H. Williams, A Reconsidera-
tion of Ada and the glumaceous brassias (Orchi-
daceae). Brittonia 24:93-1 10. 1972.
Epiphytic, cespitose herb of shady cloud and elfin
forests. Pseudobulbs lanceolate to ovoid, little com-
pressed laterally, subtended by 2 or more foliaceous
sheaths; apex 1-2-foliate. Leaves light green, flexible,
conduplicate, petiolate, acute to acuminate. Inflores-
cence arcuate or pendent, lateral at the pseudobulb base,
usually paniculate, a raceme in young plants. Flowers
fleshy, campanulate, greenish spotted with brown, the lip
usually white or cream spotted with reddish brown. Se-
pals dissimilar, the laterals partially united. Petals ovate,
acute, spreading. Lip simple, ovate to obovate, convex
apically, margins usually revolute; callus a pair of fleshy
keels at the base forming a concave depression. Column
stout, without foot; pollinia 2, waxy, with relatively
large stipe. Fruit a capsule.
A small genus of six to eight species ranging
from southern Nicaragua (Volcan Mombacho) to
Andean South America. Mesospinidium is rather
easily recognized by the small, brown-spotted,
greenish flowers on arcuate to pendent panicles.
Plants are grayish green.
According to Williams, Mesospinidium is relat-
ed to Ada and Brassia. No studies of DNA have
been published that include this genus.
Key to the Species of Mesospinidium
la. Lip 6-8 mm long, dorsal sepal oblong
Ib. Lip 4-6 mm long, dorsal sepal suborbicular
M. warscewiczii
. M. horichii
Mesospinidium horichii Bock, Orchidee (Ham-
burg) 47:246. 1996. TYPE: Costa Rica, Atlantic
rain forest, Sarapiqui jungles, near Cariblanco
(holotype: HAL 75286, not seen). Figure 33C.
Plant a cespitose epiphyte to 30 cm tall. Rhizome short;
roots 1-2 mm in diameter. Pseudobulbs small, lanceolate
to ovoid, compressed, 3.5-4.5 cm long, 1.8-2 cm wide,
subtended by 2-3 foliaceous, bracts, apically l(2)-foliate.
Leaves rather thin, carinate beneath, oblong-lanceolate to
elliptic, 15-30 cm long, 3 cm wide, acute. Inflorescence a
panicle of 15-20 flowers, lateral, 25-30 cm long with
scape, pendent or arcuate, the peduncle covered by trian-
gular, membranaceous bracts 7-9 mm long, 2-3 mm wide;
ovary with pedicel 6-7 mm long; floral bract about 1 mm
long. Flowers somewhat fleshy, 8-10 mm tall, 6-7 mm
wide, yellowish with brownish spots. Sepals dissimilar;
dorsal adnate to the petals at the base, papillose above,
suborbicular, concave, apiculate, 5-6 mm long, 4-5 mm
wide; lateral sepals elliptic, obtuse, % connate, prominently
carinate, papillose above, 6-7 mm long, 3-4 mm wide.
Petals lightly concave, acute to obtuse, recurved, 4-5 mm
long, 2-3 mm wide. Lip simple, fleshy, articulate to the
column base; blade ovate to suborbicular, or rhombic, 4-5
mm long, 3 mm wide; apex with reflexed apicule appearing
emarginate in natural position; margin revolute; callus at
the base fleshy, concave, bilamellate, pubescent between
the lamellae. Column stout, 4 mm long, broadened at the
base forming a prominent, rounded nectary, the margins
with 2 lateral fleshy projections; pollinia 2, globose, with
prominent stipe and viscidium.
This species grows in premontane rain forests
at 800-1000 m. Flowering September and Octo-
132
FIELDIANA: BOTANY
her. Costa Rica; also reported from Panama (R.
L. Dressier, pers. cornm.)-
This species is distinguished from similar M.
warscewiczii Rchb. f. by the smaller, fleshier flow-
ers with suborbicular dorsal sepal and by the
shorter lip. The name, M. endresii (Kraenzl.) Gar-
ay, has been misapplied to M. horichii.
Mesospinidium warscewiczii Rchb. f., Bot. Zei-
tung (Berlin) 10:929-930. 1852. Central Amer-
ica, Warscewicz (holotype: w). IMesospinidium
endresii (Kraenzl.) Garay, Orchidee (Hamburg)
24:187. 1973. ISolenidium endresii Kraenzl.,
Pflanzenr. 80:317. 1922. TYPE: probably Costa
Rica, Endres (holotype: w). Figure 33D.
Plant a cespitose epiphyte to 25 cm tall. Rhizome
short; roots to 1 mm in diameter. Pseudobulbs elliptic-
ovoid, compressed, 2.3-3.5 cm long, 1.5-2 cm wide,
subtended by 2-4 foliaceous bracts; apex l(2)-foliate.
Leaves petiolate; blade elliptic-lanceolate, subcoria-
ceous, carinate beneath, 10-25 cm long, 2.8-3.5 cm
wide, acute. Inflorescence a panicle with 10-15 flowers,
lateral, arcuate or subpendent, 20-25 cm long including
the scape; scape concealed by lanceolate bracts to 7 mm
long; ovary with pedicel 5-7 mm long. Flowers 9-13
mm tall, 6-8 mm wide, greenish yellow, tepals with red-
maroon spots; lip white with orange-brown spots. Sepals
dissimilar; dorsal free, oblong, concave, acute, 7-8 mm
long, 4-5 mm wide; lateral sepals elliptic-ovate, carinate
beneath, 11-12 mm long, 4-5 mm wide, the base %
connate, the apices acute. Petals elliptic-lanceolate, por-
rect, somewhat concave, 5-6 mm long, 2-3 mm wide,
acute. Lip simple, fleshy, obovate to subpandurate, 6-8
mm long, 4-5 mm wide, with rounded nectary at the
base, the apex acute and reflexed appearing retuse, the
margin somewhat plicate; callus at the base a pair of
fleshy keels to about % the lip, pubescent. Column stout,
5 mm long, broadly canaliculate; pollinia 2, waxy,
rounded, with viscidium and triangular stipe.
This species grows usually as small populations
in premontane cloud forests at 600-1200 m.
Flowering October to December. Reported from
Nicaragua to Panama.
Mesospinidium warscewiczii differs from sim-
ilar M. horichii Bock by the larger, less fleshy
flower, the oblong dorsal sepal, and the notably
larger lip.
Miltoniopsis Godefroy-Leb.
(D. E. Mora de Retana)
REFERENCE — L. A. Garay and G. C. K. Duns-
terville, Venez. Orch. 111. 6:276-278. 1976.
Epiphytic, cespitose herb of rather wet, usually mid-
elevation habitats. Pseudobulbs ovoid, strongly com-
pressed, subtended by 2 or more foliate sheaths, apex 1-
foliate. Leaves conduplicate, soft and flexible, articulate.
Inflorescence a lateral raceme at the base of the pseudo-
bulb. Flowers large and showy. Sepals free, spreading.
Petals similar to the sepals, recurved or reflexed, spread-
ing. Lip larger than sepals or petals, united to column
base by a central keel, with a pair of sharp, short, lateral
lobes at the base. Column stout, without auricles or foot;
pollinia 2, with stipe and viscidium. Fruit a capsule.
A genus of about six species ranging from Cos-
ta Rica to Andean South America. Two species
are known in Panama, but only one is known in
Costa Rica.
Little is known of the close relationships of
Miltoniopsis. The 1 -foliate pseudobulb, lip with
both central keel adnate to the column, and the
sharp lateral lobes are clearly inconsistent with
Miltonia. Still, Miltoniopsis seems to belong to
the Oncidium altissimum clade (see Chase &
Palmer, 1992), including Odontoglossum and On-
cidium (in part).
Miltoniopsis warscewiczii (Rchb. f.) Garay &
Dunsterville, Venez. Orch. 111. 6:278. 1976.
Odontoglossum warscewiczii Rchb. f., Bot. Zei-
tung (Berlin) 10:692-693. 1852. TYPE: Pana-
ma, Chiriqui at 8000 ft, Warscewicz (holotype:
w). Odontoglossum warscewiczianum Hemsley,
Biol. Centr.-Amer. 3:277. 1883. Miltonia endre-
sii Nicholson 111. Diet. Gard. 2:368. 1888. Mil-
tonia superba Schltr., Repert. Spec. Nov. Regni
Veg. 3:249. 1907. Figure 34A.
Plant cespitose, epiphytic, gray-green, 20-30 cm tall.
Rhizome short; roots 2 mm in diameter. Pseudobulbs
ovoid-ellipsoid, 3.8-4.3 cm long, 1.5-2.5 cm wide, api-
cally 1 -foliate, subtended and nearly concealed by 4-6
foliaceous bracts arranged in a fan. Leaves and folia-
ceous bracts articulate, petiolate; blades abaxially cari-
nate, oblong-elliptic, acute to acuminate, to 30 cm long,
2-3 cm wide. Inflorescence a lateral, 3-5-flowered ra-
ceme 25-30 cm long including the scape; scape with
translucent, triangular bracts; ovary and pedicel 3.5-4.5
cm long subtended by short, acute bracts 4-6 mm long.
Flowers very delicate, 4.5-6 cm high and 4.2-6 cm
wide, white to pink with a wine red to salmon spot
around the yellow callus; column white to magenta with
white anther. Sepals 2.5-3.2 cm long, 1.2-1.6 cm wide;
dorsal elliptic-obovate, erect, concave, obtuse and apic-
ulate; lateral sepals elliptic to obovate, lightly reflexed,
obtuse and apiculate when spread. Petals similar to the
sepals, obovate, 2.8-3 cm long, 1.3-1.4 cm wide; apex
recurved, obtuse, apiculate. Lip pandurate, broad, 3.5-
4 cm long, about as wide; apex deeply emarginate, unit-
ed to the column base by a narrow keel; callus at the
base semicircular with fleshy and lightly elevated border.
Column stout, 4-5 mm long; pollinia 2, waxy, with vis-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
133
cidium and short stipe. Capsules ellipsoid, 6-7 cm long
including the beak; pedicel ca. 2 cm long.
Miltoniopsis warscewiczii grows in high tree
crowns in premontane and lower montane cloud
forests (the "rainbow zone") at 1000-1800 m,
rarely at lower elevations in the area of La Virgen
de Sarapiqui. Flowering October to December.
Reported from Costa Rica and Panama.
In flower, this species is easily recognized by
the large, white, rather flat flowers with a propor-
tionately large, biblike lip. Sterile plants could be
confused with Ada chlorops (Endr. & Rchb. f.) N.
H. Williams, a species with smaller, nearly obso-
lete pseudobulbs and deeper-green leaves. They
often grow together.
This species is most closely related to the South
American M. vexillaria (Rchb. f.) Godefroy-Le-
beuf, a species with somewhat larger and much
more variably colored flowers. Both Odontoglos-
sum \varsce\viczianum Hemsley and Miltonia en-
dresii Nicholson are based on the same type as
Odontoglossum warscewiczii Rchb. f.
Notylia Lindl.
(D. E. Mora de Retana, with help from R.
L. Dressier)
Plant a small cespitose, sympodial, pseudobulbous
epiphyte; rhizomes short; roots usually white, thin, cy-
Key to the Species of Notylia
lindric. Pseudobulbs conspicuous or hidden by 1 or
more foliaceous sheaths, apex 1 -foliate. Leaves condu-
plicate, usually broad and leathery. Inflorescence a lat-
eral, pendent, or arcuate raceme borne at the base of the
pseudobulb, usually many-flowered. Flowers small, usu-
ally white to cream or pinkish, often faintly spotted, sub-
tended by narrow, acute bracts. Sepals narrow and acute,
variously spreading; the lateral sepals usually variously
connate, sometimes completely so. Petals similar to the
sepals but narrower. Lip arrowhead-shaped or trulliform,
acute and often attenuate, sometimes united to the base
of the column. Column variously elongate, cylindric,
straight, without a foot, without wings, with variously
elongate rostellum, anther dorsal, enlarged; pollinia 2,
waxy, supported on an elongate stipe and viscidium.
Fruit a capsule.
A perplexing Neotropical genus of perhaps 50
species, many of which are extremely similar and
difficult to distinguish. Three species are here
treated for Costa Rica, although there may be at
least one more (see note under N. lankesteri).
Notylia is most closely related to Macrocliniwn
Barb. Rodr. (see Chase & Palmer, 1992), which is
distinguished by the equitant leaf, often mono-
podial habit, usually subumbellate inflorescence,
and more complex flowers. Notylia species have
1 -foliate pseudobulbs and pendent lateral inflores-
cences with usually many flowers that have a
characteristic appearance, even when out of flow-
er.
la. Lateral sepals free to base
2a. Lip narrowly acute
2b. Lip subovate, obtuse
Ib. Lateral sepals at least Vi connate
N. trisepala
N. lankesteri
. . N. pittieri
Notylia lankesteri Ames, Sched. Orch. 5:34. 1923.
TYPE: Costa Rica, Province of Limon, Rio Es-
trella, sea level, Lankester & Sancho 373 (AMES,
photo seen). IN. brenesii Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:249. 1923. TYPE: Cos-
ta Rica, San Pedro de San Ramon, Brenes 173
(lectotype: CR; isorype: AMES, photo seen).
Plant cespitose, to about 15 cm tall. Pseudobulb
ovoid, ca. 1.5 cm long, 1 cm wide. Leaves oblong-el-
liptic, rounded at the base, obtuse, 11-14 cm long, 2-
3.5 cm wide. Inflorescence a lateral raceme 4-16 cm
long, densely flowered; floral bracts 4-5 mm long.
Flowers aromatic, fleshy, white; petals with orange
spots. Sepals similar, ca. 5 mm long, 3 mm wide; dorsal
concave; lateral free, elliptic-ovate. Petals elliptic, ob-
tuse, ca. 4 mm long, 2.5 mm wide, thickened at the
unguiculate base. Lip short-unguiculate; ca. 4 mm long,
2-2.5 mm wide; blade ovate to lanceolate, acute, the tip
compressed; margins ascending; callus triangular, erect
near the base. Column fleshy, stout, ca. 3 mm long.
Epiphytic in moist forests at 0-1500 m. Flow-
ering at least November. Apparently endemic to
Costa Rica.
Notylia lankesteri is distinguished from other
species by the free sepals and obtuse lip. The se-
pals and petals are notably fleshy. The original
description indicates that the flowers are "pun-
gently spicy." Much of the above description is
taken from the original description.
One of our reviewers analyzed a flower from
the type of Notylia brenesii Schltr. and found that
134
FIELDIANA: BOTANY
the lip is long-attenuate as described and similar
in shape to an elephant's trunk. This may prove
to be different from the species here described.
Notylia pittieri Schltr., Beih. Bot. Centralb.
36(2):418. 1918. TYPE: Costa Rica, Sur les
Crescentia a Boruca, Pittieri 6850 (AMES, draw-
ing of type). Figures 34C,D.
Plant epiphytic, cespitose, forming small clumps ca. 12
cm tall. Rhizome very short; roots slender, ^ 1 mm in
diameter. Pseudobulbs ovate to oblong, apically 1 -foliate,
somewhat flattened, sulcate, 0.5-3.5 cm long, 0.4-1 cm
wide, subtended by 2-3 foliaceous bracts. Leaves some-
what fleshy, oblong to elliptic-lanceolate, 3-17 cm long,
0.8-3.5 cm wide, shortly petiolate, acute. Inflorescence a
densely and successively flowered, pendent raceme, 3-20
cm long; ovary with pedicel 3 mm long, subtended by
shorter, triangular, acute bracts. Flowers somewhat spread-
ing, with tannish cream sepals; petals cream with brown-
orange spots; lip white. Sepals dissimilar; dorsal concave,
ligular-lanceolate, 4-5.5 mm long, 1.6-1.7 mm wide; syn-
sepal 3.7-4.5 mm long, 1.2-2.2 mm wide, minutely biden-
tate apically. Petals oblique, falcate-lanceolate, 3.3-4.5 mm
long, 0.6-1.2 mm wide; acute and attenuate. Lip 3-3.7 mm
long, 1-2.2 mm wide, with short claw ca. 0.5 mm long;
blade subsagittate to lanceolate, acute, with prominent keel
1-1.5 mm on claw and base of the blade. Column slender,
erect, 2-3 mm long; anther cucullate, elongate, pollinia 2,
with stipe and small viscidium.
Epiphytic in moist and seasonally dry forests in
Pacific lowlands at 0-500 m. Flowering March,
April, and June to September. Costa Rica; to be
expected in western Panama.
Notylia pittieri is distinguished by the green or
tan-green sepals, the lateral sepals being nearly
totally connate, and the trullate-lanceolate lip.
Notylia trisepala Lindl., Paxton's Fl. Gard. 3:45.
1852. TYPE: Guatemala, Van Houtten (K). IN.
turialbae Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 19:145-146. 1923. TYPE: Costa Rica, Tur-
ialba, A. and C. Erode 1186 (AMES, drawing of
type). N. barken of authors, not Lindl. Figure
34B.
Plant a cespitose epiphyte, ca. 15 cm tall. Rhizome very
short; roots slender, :£ 1 mm in diameter. Pseudobulbs
ovoid, apically 1 -foliate, largely concealed by 2-3 subtend-
ing bracts, 1-2.2 cm long. Leaves somewhat fleshy, abax-
ially carinate, elliptic to ovate-lanceolate, subpetiolate, ob-
tuse, 5-13.5 cm long, 1.2-3 cm wide. Inflorescence a pen-
dent raceme 12-25 cm long; ovary with pedicel 5 mm
long, subtended by shorter, triangular, acute bracts. Flowers
successive, white or greenish white, petals with orange-
yellow bars near the base. Sepals free, similar; dorsal el-
liptic, 5-6 mm long, 1-2.5 mm wide; lateral sepals lance-
elliptic to lance ovate, 4-6 mm long, 1-1.6 mm wide. Pet-
als similar to the sepals. Lip unguiculate, 3.5-5 mm long,
1-2 mm wide; blade triangular-ovate, acute, with prominent
keel on claw and base of the blade. Column stout, 2.5-2.7
mm long; anther cucullate, elongate; pollinia 2, with stipe
and small viscidium.
Epiphytic in moist forest at 100-900 m. Flow-
ering January to April. Reported from Guatemala
to Costa Rica; probably also in western Panama.
Notylia trisepala is distinguished by the white
or greenish white flowers and free lateral sepals.
Oncidium Sw.
(D. E. Mora de Retana)
REFERENCES — Retana, D. E. M. de, and J. T
Atwood, Orchids of Costa Rica. 2. Icones Plan-
tarum Tropicarum 15: pi. 1401-1500. 1992. Re-
tana, D. E. M. de, and J. T. Atwood, Orchids of
Costa Rica. 3. Icones Plantarum Tropicarum 16:
pi. 1501-1600. 1993.
Epiphytic, rarely terrestrial, cespitose or rhizomatous
herb. Pseudobulbs present to obsolete or absent, subtend-
ed by 0-several foliaceous bracts; apex 0-3-foliate. Leaves
conduplicate, chartaceous to fleshy, flat to terete, petiolate
to sessile, acute to obtuse. Inflorescence lateral at the pseu-
dobulb base, a raceme of 1 -several flowers or paniculate.
Flowers small to large, generally showy, usually yellow
with reddish brown spots, sometimes white or with purple
spots, rarely wine red to pink. Sepals similar or the laterals
dissimilar and often variously connate. Petals usually sim-
ilar to the sepals but somewhat smaller, rarely larger. Lip
divergent from the column forming an angle of at least 90°,
usually fleshy, simple or 3-lobate, the midlobe often emar-
ginate or 2-lobate; callus often complex comprising various
plates and teeth. Column stout, footless, with a fleshy plate
below the stigma (tabula infrastigmatica), the apex with or
without conspicuous wings; pollinia 2, waxy, supported on
a conspicuous stipe and viscidium. Fruit a capsule.
A Neotropical genus of about 450 species de-
pending in part on circumscription. Twenty-eight
species are known in Costa Rica, although there
may be others hiding under the synonyms we ac-
cept. Although we exclude O. luridum Lindl., it
may occur in mangroves along the Atlantic coast,
as reported both south and north of Costa Rica.
According to Chase, Oncidium includes several
distinct alliances, some more closely related to other
currently defined genera than to other oncidiums.
The key to genera of the Oncidiinae allows one to
arrive at Oncidium through several couplets reflect-
ing the incoherent diversity of this polyphyletic "ge-
nus."
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
135
Key to the Species of Oncidium
la. Plants with a single, fleshy flat or terete apical leaf, no foliaceous bracts; pseudobulbs small or
essentially obsolete 2
2a. Leaves terete (see Figs. 35A-C) 3
3a. Lateral lobes porrect, column wings linear, curved O. ascendens
3b. Lateral lobes not porrect; column wings otherwise, not linear 4
4a. Callus prominent, noselike in profile, with a pair of teeth near the base
O. cebolleta
4b. Callus not noselike, with 3 ridges apically but lacking teeth O. teres
2b. Leaves flat, fleshy but never terete 5
5a. Flowers white with reddish purple spots, lip ^ 12 mm wide O. carthagenense
5b. Flowers brown to yellow spotted with brown, lip > 13 mm wide
O. luridum [see excluded taxa]
Ib. Plants with 2 or more flat leaves including foliaceous bracts, never fleshy except perhaps in O.
ampliatum 6
6a. Lip white or cream when young, sometimes turning yellow, never marked with red to brown
7
7a. Lip 2: 1.6 cm long 8
8a. Lip > 2.5 cm long O. schroederianum
8b. Lip < 2.4 cm long 9
9a. Lip apex anchor-shaped, basal callus prominent O. cariniferum
9a. Lip simple, without basal callus O. stenoglossum
7b. Lip ^1.5 cm long 10
lOa. Inflorescence very large, > 1.5 m long O. ochmatochilum
lOb. Inflorescence relatively short, < 1 m long 11
11 a. Inflorescence bracts subtending primary branches < 1.5 cm long, not cym-
biform O. panduriforme
1 Ib. Inflorescence bracts subtending primary branches > 2.5 cm long, cymbiform
O. storkii
6b. Lip yellow or greenish yellow when young, often marked with red to brown 12
12a. Inflorescence a raceme, or if not, pseudobulb never with purple spots (O. planilabre
may produce a branch in robust specimens 13
13a. Plant basically a vine; shoots at least 6 cm apart on wiry rhizome lacking bracts
between shoots O. globuliferum
13b. Plant cespitose, never a vine 14
14a. Foliaceous bracts subtending pseudobulbs 5-8 O. cristagalli
14b. Foliaceous bracts subtending pseudobulbs 2-3 15
15a. Tepals yellow; with cymbiform floral bracts enclosing the ovary mostly
> 1.5 cm long O. warscewiczii
15b. Tepals brown with yellow margin; floral bracts lanceolate, < 1.2 cm
long, never enclosing ovary O. planilabre [see excluded taxa]
12b. Inflorescence a panicle, pseudobulb various 16
16a. Inflorescence with small, infertile flowers in addition to the single fertile flower
per primary branch O. bryolophotum
16b. Inflorescence lacking small infertile flowers 17
17a. Primary inflorescence branches subtended by spreading, cymbiform bracts
> 3 cm long O. bracteatum
17b. Bracts subtending primary branches not as above 18
18a. Distance across lateral lobes > 2X greater than midlobe 19
19a. Leaves 5-7 per shoot, > 1.5 cm wide O. luteum
19b. Leaves 3-4 per shoot, < 1.5 cm wide O. cheirophorum
18b. Distance across lateral lobes < 2X width of midlobe, midlobe usually
broader 20
136 FIELDIANA: BOTANY
20a. Pseudobulbs spotted with purplish brown 21
2 la. Foliaceous sheaths lacking, leaves somewhat fleshy
O. ampliation
21b. Foliaceous sheaths present, leaves not fleshy 22
22a. Tepals yellow with brownish midveins
O. obryzatoides
22b. Tepals not as above 23
23a. Petals never spatulate, heavily blotched with
brown or purple to > % from the base; pseudo-
bulb usually with 2 leaves O. dichromaticwn
23b. Petals distinctly spatulate, spotted only to < Vi
from the base O. klotzschianum
20b. Pseudobulbs not spotted with purplish brown 24
24a. Flowers ^1.5 cm tall O. parviflorwn
24b. Flowers > 1.5 cm tall 25
25a. Tepals yellow stained with brown along midveins O. obryzatoides
25b. Tepals dark brown or yellow spotted with brown 26
26a. Pseudobulbs suborbicular to elliptic or ovate, strongly compressed, sharp-edged 27
27a. Tepals nearly solid dark brown with yellow tips O. ansiferum
27b. Tepals yellow with few brown spots O. stenobulbon
26b. Pseudobulbs various, neither strongly compressed, nor suborbicular, rather lanceolate, ovoid
or ellipsoid 28
28a. Plants terrestrial; foliaceous sheaths without articulations, inflorescence erect
O. ensatum
28b. Plants epiphytic; leaves articulate, inflorescence pendent 29
29a. Ovary with pedicel ^ 2.5 cm long, pseudobulb ellipsoid with 1 apical leaf ....
O. stenotis
29b. Ovary with pedicel < 2.5 cm long, pseudobulbs lanceolate, with 2-3 (rarely 1)
apical leaves 30
30a. Midlobe of lip about 2X as long as base below narrowest part of isthmus;
lateral lobes pointing laterally, callus at least as broad as long O. isthmi
30b. Midlobe about as long as base, lateral lobes somewhat porrect; callus nar-
rower than long O. polycladium
Oncidium ampliatum Lindl., Gen. Sp. Orch. PI.
202. 1833. TYPE: Panama and Colombia, Cum-
ing 1312 (holotype: K). Oncidium bernoullian-
um Kraenzl., Pflanzenr. IV. 50(Heft 80):231.
1922. TYPE: Guatemala, Bernouli 339 (holo-
type: w). Figure 38D.
Plant epiphytic, cespitose or shortly creeping. Rhi-
zome short; roots 1.5-4 mm in diameter. Pseudobulbs
green with reddish brown markings, ovoid to orbicular,
laterally flattened, coarsely rugose, 3.5-10 cm long, 3.5-
8 cm wide, concealed when young by nonfoliaceous
bracts, apically (l)2-3-foliate. Leaves somewhat petio-
late; blades coriaceous, elliptic, 6-35 cm long, 3-8.5 cm
wide, subacute to obtuse. Inflorescence lateral, a raceme
when small, panicle when large, 35-70 cm long includ-
ing the scape; ovary and pedicel 2.5-4 cm long. Flow-
ers somewhat successive, 2.5-3.5 cm long, 2.2-3.5 cm
wide, brilliant yellow, the sepals somewhat spotted with
red-brown, also the lip with reddish brown surrounding
the callus. Sepals similar, 5-9 mm long, 5-7 mm wide,
obovate-spatulate, concave. Petals larger than the sepals.
with claw; blade 8-13 mm long, 6-9 mm wide, subor-
bicular. Lip 3-lobate, 1.8-3.5 cm long, 2.5-3.5 cm wide;
lateral lobes small, 5-6 mm wide; midlobe reniform,
emarginate, with undulate margins, forming a narrow
isthmus basally; callus at the isthmus complex, fleshy,
somewhat elevated, consisting of 2 teeth basally and 3
teeth apically. Column short, 3-5 mm long, with dentate
column wings; anther bed dentate; pollinia 2, supported
on a viscidium and very short stipe. Capsules ellipsoid,
retaining floral segments, 2.5-3.5 cm long; pedicel 2-
3.5 cm long.
Uncommon epiphyte in dry, deciduous to sem-
ideciduous forests of the Pacific lowlands at 0-
800 m. Flowering December to April. Broadly
distributed from Guatemala to Venezuela, Trini-
dad, and Peru.
Oncidium ampliatum is easily recognized even
when sterile by the strongly rugose, apically usu-
ally 2-3-foliate pseudobulbs. The flowers are no-
table for their proportionately enormous, yellow
lips.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
137
Oncidium ansiferum Rchb. f., Bot. Zeitung (Ber-
lin) 10:696. 1852. TYPE: Chiriqui, Warscewicz
(holotype: w, photo seen). Oncidium lankesteri
Ames, Sched. Orch. 4:53. 1923. TYPE: Costa
Rica, Irazii Volcano, Lankester & Sandro 415
(holotype: AMES). Figure 40A.
Plant cespitose, epiphytic. Rhizome abbreviated,
shortly climbing; roots white, 1-2 mm in diameter.
Pseudobulbs elliptic-ovoid, strongly flattened laterally,
furrowed when dry, 6-12 cm long, 3-6 cm wide; base
subtended by 2-3 imbricate, foliaceous, bracts; apex 1-
2-foliate. Leaves short to long-petiolate, strongly cari-
nate abaxially, 1-2(3) at the pseudobulb apex; blade el-
liptic-lanceolate, obtuse to subacute, coriaceous, con-
duplicate at the base, 15-40 cm long, 2.5-5.5 cm wide.
Inflorescences lateral, 1-2 at the base of the shoot, an
erect or arcuate panicle, exceeding the leaves, 50-80 cm
long; branches 15-20 cm long, with 4-6 flowers; ovary
with pedicel 2-3.5 cm long. Flowers with tepals red-
brown tipped with yellow, 3-3.5 cm in diameter; lip
brilliant yellow with a brown border around the callus.
Sepals and petals similar, free, reflexed, 1.5-1.7 cm
long, 5-7 mm wide, with undulate margins. Lip 3-lo-
bate, 1.5-2 cm long, 1.2-1.8 cm wide; lateral lobes
small, rounded, 5-7 mm wide; midlobe emarginate, with
denticulate margin; isthmus 3-5 mm wide; callus fleshy,
consisting of 7 teeth, 5 in 1 plane, 2 lower at the apex.
Column short, ca. 5 mm long, with prominent tabula
infrastigmatica; column wings broad, dolabriform, with
crenulate margins; anther papillose; pollinia 2, obovate,
supported on a prominent stipe and viscidium. Capsules
ellipsoid, 4.5 cm long; pedicel 2 cm long.
This species grows as a low-density epiphyte
in cloud forests at (1000)1350-2200 m. Flower-
ing December to June with peak in February to
April. Costa Rica and Panama.
Oncidium ansiferum is easily recognized by the
strongly compressed, sharp-edged pseudobulbs
and by the tepals, which are dark brown with yel-
low apices. It is closely related to O. stenobulbon
Kraenzl., which grows in dryer habitats, usually
at lower elevations (800-1050 m). Plants of that
species are more yellow-green and have shorter
and proportionately broader leaves, yellow tepals
spotted with less intense brown, and less pro-
nounced column wings. From the regions of San
Pedro and Santiago de San Ramon (850 and 1000
m, respectively) are two specimens that appear
somewhat intermediate with O. stenobulbon and
may represent natural hybrids.
The common name of "caite" has been applied
to this species because the odd, flattened pseu-
dobulbs are similar in shape to caite, a kind of
sandal worn by campesinos.
Oncidium ascendens Lindl., Edwards's Bot. Reg.
28: sub t. 4. 1842. TYPE: Guatemala. Figure
35A.
Plant an erect or pendent, cespitose epiphyte; rhizome
short, roots to 2 mm in diameter. Pseudobulbs minute,
suborbicular, 5-8 mm in diameter; subtended by non-
foliaceous bracts; apex 1 -foliate. Leaves terete, fleshy,
dark green with reddish spots, 10-45 cm long, 6-1.2 cm
in diameter, acute. Inflorescence lateral, paniculate, usu-
ally shorter than the leaf, 10-30 cm long including the
scape; branches short; flowers congested; ovary with
pedicel 7-8 mm long. Flowers with tepals greenish
spotted with brown; lip yellow with spotted callus. Se-
pals similar, free, concave, shortly unguiculate, obovate,
obtuse, 6-7 mm long, 3-5 mm wide. Petals similar to
the sepals, shortly unguiculate, concave, obovate, ob-
tuse, 6-8 mm long, 3-5 mm long, with undulate mar-
gins. Lip 3-lobate, 9-10 mm long, 8-9 mm wide; lateral
lobes rounded, porrect, 3-4 mm long; midlobe emargin-
ate to 2-lobate, reniform, forming a distinct isthmus at
the base 2-3 mm wide. Column stout, about 3 mm long,
with a pair of linear, curved wings apically; anther pa-
pillose; pollinia pyriform, supported on viscidium and
oblanceolate stipe. Capsules ellipsoid, shortly beaked,
2.3 cm long; pedicel ca. 1 cm long.
Epiphytic in hot, humid, tropical lowland for-
ests with a short dry season at 0-1000 m. Flow-
ering January to March. Reported from Mexico to
Costa Rica, Colombia, and the Caribbean.
Three terete-leaved species of Oncidium are
recognized in Costa Rica, O. ascendens, O. ce-
bolleta (Jacq.) Sw., and O. teres Ames & C.
Schweinf. Of these species, O. ascendens is
unique in the porrect lateral lobes and in the lin-
ear, incurved column wings.
The entire alliance around O. ascendens needs
to be reexamined, and we make no attempt to re-
peat questionable synonymies published else-
where.
Oncidium bracteatum Warsz. & Rchb. f., Bot. Zei-
tung (Berlin) 10:695. 1852. TYPE: Probably Pan-
ama, Chiriqui, Warscewicz (w). Oncidium palea-
tum Schltr., Repert. Spec. Nov. Regni Veg. Beih/
19:260-261. 1923. TYPE: Costa Rica, San Rafael
de San Ramon, Brenes 206 (drawing of type:
AMES). Oncidium henricigustavi Kraenzl., Pflan-
zenr. IV. 50(Heft 80):242. 1922. TYPE: Costa
Rica, Endres 292 (w). Figure 38A.
Plant a cespitose epiphyte of large branches to 60 cm
high; rhizome short; root 1-1.5 mm in diameter. Pseu-
dobulbs ovate, compressed, sulcate, subtended by 2-4
foliaceous bracts, apex 1-2 foliate. Leaves elliptic-lan-
ceolate, carinate abaxially, 15-40 cm long, 1.5-3.5 cm
wide, conduplicate at the base; apex asymmetrically
acute. Inflorescence lateral, paniculate, s 1.25 m tall,
138
FIELDIANA: BOTANY
peduncle scarious; primary branches subtended by cym-
biform bracts 3.5-4.5 cm long, the base with short in-
ternodes and usually with 2 imbricate bracts; ovary with
pedicel 1.5-2.5 cm long, subtended by glumaceous
bracts 1.5-2.8 cm long, 8-10 mm wide when spread.
Flowers brilliant yellow marked with dark brown, lip
brilliant yellow with brown base and isthmus. Sepals
dissimilar, elliptic, carinate abaxially, narrowed at the
base, with undulate margins; dorsal 1.3-1.5 cm long, 5-
6 mm wide, obtuse, apiculate; lateral sepals falcate, re-
flexed or not, 1.5-1.7 cm long, 5-6 mm wide, acute.
Petals similar to the dorsal sepal, 1.3-1.5 cm long, 5-6
mm wide, obtuse, apiculate, margins undulate. Lip 3-
lobate, 1.3-1.6 cm long, 1.2-1.5 cm wide; lateral lobes
at the base rounded, somewhat retrorse; midlobe reni-
form, deeply emarginate, forming a pair of rounded
lobes; isthmus ca. 5 mm wide; callus fleshy, with a thick
central keel, 4 lateral and 3 apical teeth. Column 6-8
mm long, tabula infrastigmatica with fleshy borders,
wings slender; anther; pollinia, supported on an elongate
stipe and viscidium. Capsules ellipsoid, 4.5-5 cm long;
pedicel ca. 3 cm long.
Epiphytic on shady, larger branches in premon-
tane and lower montane cloud forests at 1000-1700
m. Flowering mostly May to October; sporadically
throughout the year. Costa Rica and Panama.
Oncidiwn bracteatum resembles O. storkii in the
conspicuous inflorescence bracts, but the latter spe-
cies has strictly secund branches and dull yellow
flowers with much less pronounced lateral lobes.
The name O. paleatum has been misapplied to
O. stenotis Rchb. f. (F. Hamer, Icon. PI. Trop. 11:
t. 1059), but the drawing of the type of O. palea-
tum shows a smaller, 2-foliate pseudobulb with
very narrow leaves and long bracts subtending the
primary branches of the inflorescence, which are
consistent with O. bracteatum, not O. stenotis.
Also, the original description of O. paleatum in-
dicates a much smaller flower, consistent with O.
bracteatum.
OIK id in ni bryolophotum Rchb. f., Gard. Chron.
738. 1871. TYPE: Central America, Veitch (w).
Oncidium megalous Schltr., Repert. Spec. Nov.
Regni Veg. 9:30. 1910. TYPE: Costa Rica, Es-
meralda, Barba Massif, Biolley 7256. Oncidium
asparagoides Kraenzl., Pflanzenr. IV. 50(Heft
80): 175. 1922. TYPE: Costa Rica, Endres 127
(w). Oncidium microphyton Kraenzl. Pflanzenr.
IV. 50(Heft 80):204-205. 1922. TYPE: Costa
Rica, Endres s.n. (w). Figure 37D.
Plant most variable in size, 10-25 cm tall; roots slen-
der, about 1 mm in diameter. Pseudobulbs ovoid to lan-
ceolate, little compressed, 2-8 cm long, 1.5-2.5 cm
wide; subtended by 2-3 foliaceous bracts; apex 1-2-fo-
liate. Leaves shortly petiolate, blades elliptic-lanceolate,
7-25 cm long, 1-2.5 cm wide, acute. Inflorescence lat-
eral, paniculate, 6 cm-1 m, complexly branched; pri-
mary branches to 5-10 cm long, with a single fertile and
many aborted flowers per branch with oblong-linear te-
pals; ovary with pedicel 1-1.5 cm long. Fertile flowers
ca. 1.2-2 cm tall, yellow with reddish brown markings
on the tepals and lower half of lip. Sepals dissimilar;
dorsal shortly unguiculate, ovate, acute, 7-9 mm long,
4-5 mm wide; lateral sepals linear-lanceolate, acute, re-
flexed, 9-10 mm long, 2-3 mm wide. Petals similar to
the dorsal sepal, shortly unguiculate, 7-9 mm long, 4-
6 mm wide. Lip 3-lobate, nearly as broad at the base as
across the midlobe, 1-1.5 cm long, 1-1.5 cm wide; lat-
eral lobes bluntly triangular, 3-4 mm long; midlobe 1-
1.5 cm wide, deeply emarginate forming 2 rounded
lobes; isthmus ca. 4-5 mm wide; callus at the base with
a central blunt keel and 4 rows of blunt teeth with ad-
ditional teeth on the lateral lobes. Column ca. 5 mm
long, with tabula infrastigmatica, the apex with a pair of
large, porrect, rounded wings. Capsules 3.5 cm long in-
cluding elongate beak, pedicel 1-1.5 cm long.
Epiphytic on larger branches of open canopies
in premontane and lower montane cloud forests
at 1000-2100 m. Flowering July to December.
Costa Rica and Panama; probably to Peru.
Oncidium bryolophotum is easily distinguished
by the asparagus-like, slender inflorescence,
which is often seen waving in the wind. Each pri-
mary branch has a single fertile and numerous
sterile flowers.
Plants from Costa Rica and Panama have often
been accepted as O. heteranthum Poeppig &
Endl., a species based on a Peruvian type (w) that
has roots of larger diameter, inflorescences usually
with two flowers per branch, and larger aborted
flowers with obovate tepals. Most of the Ecua-
dorian specimens at SEL may be O. bryolophotum
rather than O. heteranthum as labeled, but two
specimens from Bolivia and Peru show root and
inflorescence features consistent with the latter.
All living and preserved plants that we have
seen from Costa Rica represent a single species
that corresponds with O. bryolophotum and ov-
erdescribed by Fritz Kraenzlin and Rudolf Schlech-
ter. Both O. megalous Schltr. and M. asparagoides
Kraenzl. are straightforward O. bryolophytum
Rchb. f. The type of O. microphyton Kraenzl. is
a plant less than 10 cm tall with a few aborted
flowers and a single fertile flower showing the
characteristic lip shape of O. bryolophotum.
Oncidium cariniferum (Rchb. f.) Beer, Pract.
Stud. Fam. Orch. 283. 1854. Odontoglossum
cariniferum Rchb. f., Bot. Zeitung (Berlin) 10:
638. 1852. TYPE: Middle America. Figure
36A.
Plant a cespitose epiphyte. Rhizome short; roots 1-2
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
139
mm in diameter. Pseudobulbs sulcate, ovoid, 7-12 cm
long, 4-6 cm wide, subtended by 2-4 foliaceous bracts
when young, apex 2-foliate. Leaves coriaceous, carinate
abaxially, elliptic-lanceolate, 15-40 cm long, 3-6 cm
wide, base conduplicate, apex acute. Inflorescence a lat-
eral panicle, 50-100 cm long including the scape;
branches primary, 13-20 cm long; ovary with pedicel
2-3 cm long. Flowers 6-10 per branch, about 4 cm
high, 2 cm broad; tepals greenish yellow spotted with
reddish brown; lip white aging yellow, callus clear wine
red, column white spotted with reddish brown. Sepals
free, subequal, elliptic-lanceolate, acuminate; dorsal
erect, carinate externally, 2.4-2.5 cm long, 6 mm wide;
laterals subfalcate, strongly thickened and carinate api-
cally, 2.4-2.6 cm long, 4-5 mm wide. Petals similar to
the dorsal sepal, elliptic-lanceolate, carinate, apiculate,
2.4-2.5 cm long, 5-6 mm wide. Lip anchor-shaped,
1.8-2 cm long, 2.1-2.3 cm wide with conspicuous claw
at the base, the blade reniform, shallowly emarginate,
and apiculate; callus fleshy, elevated, with 2 high, short,
lateral keels at the base, a low central keel, and a pair
of ascending teeth in front. Column stout, a bit sigmoid
in profile, about 1 cm long, broadened at the base, with
2 ascending projections below the stigma; pollinia 2,
obovate, with prominent stipe and viscidium.
Epiphytic and uncommon on the larger branch-
es of premontane and lower montane forests at
1000-2000 m. Flowering September to February.
Costa Rica and Panama; doubtfully reported from
Colombia and Venezuela.
The species is easily recognized by the white,
anchor-shaped lip and chocolate brown tepals.
The callus suggests a relationship with Odonto-
glossum, and we note the angle between the col-
umn and lip would also if the column were not
reflexed at the base.
Oncidium carthagenense (Jacq.) Sw., Kongl. Ve-
tensk. Acad. Nya Handl. 21:240. 1800. Epiden-
drwn carthagenense Jacq. Select. Stirp. Amer.
228, t. 133, fig. 4. 1763. Figure 35D.
Plant epiphytic, cespitose with few shoots. Rhizome
short; roots to 2 mm in diameter. Pseudobulbs nearly
obsolete, about 1 cm long, 1 cm wide, concealed when
young by bracts, apically 1 -foliate. Leaves coriaceous
to fleshy, elliptic, sharply keeled abaxially, conduplicate
at the base, apically acute, 20-30 cm long, 6—8 cm wide.
Inflorescence usually pendent, a lateral panicle 75-125
cm long including the scape, the nodes with triangular
bracts; ovary with pedicel 1.5-2.5 cm long. Flowers
with sepals and petals cream spotted with maroon-
brown; 1.8-2.2 cm in diameter. Sepals similar, reflexed
and unguiculate; blades concave, suborbicular, 9-12 mm
long, 6-8 mm wide, obtuse, the margins undulate. Petals
similar to the sepals but broader, 1.2-1.4 cm long, 7-9
mm wide, the margin strongly undulate. Lip 3-lobate,
subpandurate, constricted on the upper half, 1.2-1.4 cm
long, 8-10 mm wide; midlobe, reniform; lateral lobes
somewhat fleshy with revolute margins; basal callus
consisting of 4 fleshy tubercles, 2 near the base sepa-
rated by a central furrow and 2 near the middle. Column
2 mm long, with a pair of 2-lobate wings apically; anther
papillose; pollinia 2, rounded, sulcate, with viscidium
and short stipe.
Epiphytic in humid but seasonally dry forests
at 300-1100 m on trunks and large branches.
Flowering June to August. Reported from Florida,
West Indies, Central America, and much of trop-
ical South America. The range may depend ulti-
mately on the synonymies accepted.
Oncidium carthagenese is easily recognized in
Costa Rica by the white maroon-spotted flowers,
the relatively small, inconspicuous pseudobulb,
and the large, fleshy leaves, after which it has
earned the common name orejas de burro. Exten-
sive synonymies published by authors reflect
much variation over the range.
Oncidium cebolleta (Jacq.) Sw., Kongl. Vetensk.
Acad. Nya Handl. 21:240. 1800. Epidendrum
cebolleta Jacq., Enum. PI. Carib. 30. 1760.
TYPE: Colombia, Cartajena. Figure 35C.
Plant an erect or pendent, cespitose epiphyte. Rhi-
zome short; roots 1.5-2 mm in diameter. Pseudobulbs
globose, 1 -foliate, 8-10 mm high and wide, mostly con-
cealed by bracts. Leaves terete, canaliculate, green or
spotted with reddish brown, apex acute, 26-40 cm long,
8-15 mm wide. Inflorescence a lateral panicle 25-40
cm long including the scape; ovary and pedicel 1-1.5
cm long. Flowers 1.9-2.2 cm tall, 1.4—1.7 cm wide;
tepals greenish spotted with reddish brown; lip yellow
with callus marked brown. Sepals similar, obovate, con-
cave, obtuse and apiculate, 7-8 mm long, 4.5-5.5 mm
wide. Petals spatulate with undulate margins, obtuse and
apiculate, 8-9 mm long, 3.5-4.5 mm wide. Lip 3-lobate,
pandurate, 1.3-1.5 cm long, 1.4-1.7 cm wide, with cren-
ulate margins; lateral lobes suborbicular to obovate, ob-
tuse or rounded, 5-8 mm long, 3-4 mm wide; midlobe
emarginate; callus consisting of a large nose-shaped cen-
tral keel and 2 lateral teeth. Column 3-4 mm long, with
a pair of 2-lobate wings; anther papillose; pollinia 2,
waxy, with conspicuous oblanceolate stipe and visci-
dium.
Epiphytic on larger branches and widespread in
seasonally dry forests below 600 m. Flowering
late December to March. Reported as widespread
from Mexico through Central and South America.
Of the three species of terete-leaved oncidiums
recognized in Costa Rica, O. cebolleta has the
largest flower. It is most similar to O. teres Ames
& C. Schweinf., but the column wings are 2-lo-
bate, and the callus includes a pair of teeth flank-
ing the central keel.
Oncidium cheirophorum Rchb. f., Bot. Zeitung
(Berlin) 10:695. 1852. TYPE: Panama, Volcan,
140
FIELDIANA: BOTANY
Warscewicz (holotype: w, photo seen). Oncidi-
um dielsianum Kraenzl., Pflanzenr. IV. 50(Heft
80): 197, t. 198F. 1922. TYPE: Costa Rica, En-
dres (holotype: w, photo seen). Oncidium ma-
crorhynchum Kraenzl., l.c.:200, t. 198J. 1922.
TYPE: Costa Rica, San Juan de Candelaria, En-
dres (holotype: w, photo seen). Oncidium chei-
rophorum var. exauriculatum Hamer & Garay,
Las Orquideas de El Salvador 2:176. 1975.
TYPE: Costa Rica, Guanacaste, Tilaran, Stand-
ley & Valeria 46266 (holotype: AMES; Isotype:
us). Oncidium exauriculatum (Hamer & Garay)
R. Jimenez, Orquidea (Mex.) 12:270. 1992.
Figure 38C.
REFERENCE — R. Jimenez Machorro, Oncidium
exauriculatum: Una Orquidea localizada reciente-
mente en Mexico. Orquidea (Mex.) 12:269-276.
1992.
Plant cespitose, sometimes densely so, epiphytic, ^
15 cm tall excluding the inflorescence. Rhizome short;
roots about 1 mm in diameter. Pseudobulbs smooth,
green, marked and suffused with brown, pyriform to su-
borbicular, somewhat compressed, apically 1 -foliate,
1.5-3.5 cm tall, 1.5-2.5 cm wide, subtended by 2-3 fo-
liaceous bracts. Leaves thin, 5-15 cm long, 5-14 mm
wide, elliptic-lanceolate, shortly petiolate and condupli-
cate at the base, apically acute, with a prominent abaxial
keel. Inflorescence a lateral panicle, 10-25 cm long,
including the scape; ovary and pedicel 6-9 mm long.
Flowers greenish yellow, 1-1.5 cm across. Sepals dis-
similar, unguiculate, 5-6 mm long, 4-5 mm wide; dorsal
erect, obovate, strongly concave, obtuse, apiculate; lat-
eral sepals shortly connate, reflexed, subquadrate, retuse.
Petals shortly unguiculate, reflexed, ovate, obtuse, apic-
ulate, 5-6 mm long, 4-5 mm wide. Lip adnate to the
base of the column, strongly 3-lobate, with a pair of
fleshy glands at the base, 1.1-1.2 cm long, 1.2-1.4 cm
wide; lateral lobes subquadrate, lightly incurved, round-
ed; midlobe reniform, emarginate, the margins lightly
undulate; callus complex below the isthmus, fleshy, cru-
ciform. Column 2-3 mm long, with a pair of broad,
subquadrate, porrect wings or wings lacking (var. ex-
auriculatum); anther terminal, much elongate; pollinia
2, with conspicuous, slender stipe ca. 3 mm long and
viscidium. Capsules ellipsoid, with beak and persistent
flower, 1.5-1.6 cm long; pedicel 8-10 mm long.
Epiphytic in open canopies, usually on larger
branches, in premontane and lower montane for-
ests at 600-1700 m. Flowering October to Feb-
ruary (var. cheirophorum, mostly October to De-
cember; var. exauriculatum, mostly January and
February). Reported from Mexico to Colombia.
This species is easily recognized out of flower
by the pyriform pseudobulbs lightly stained with
brown. The waxy yellow flowers with cruciform
callus are distinctive when plants are in flower.
We accept O. exauriculatum, based on the lack
of conspicuous column wings, as a geographic va-
riety of O. cheirophorum. Some herbarium spec-
imens in Costa Rica show intermediate column
wings. We note, however, that specimens from
about central Costa Rica northward mostly lack
the wings, occur at lower elevations (600-1450
m), and flower in late December to February,
while O. cheirophorum occurs at 1000-1700 m
and flowers from October to December and rarely
in January. R. Jimenez Machorro reports that one
in 10 flowers of Mexican var. exauriculatum pro-
duce fruits through self-pollination.
Oncidium cristagalli Rchb. f., Bot. Zeitung 10:
697. 1852. TYPE: Mexico, Oaxaca, Galeotti
5289 (holotype: w, photo seen). Oncidium de-
cipiens Lindl., Fol. Orch. Oncidium 22. 1855.
TYPE: Mexico, Oaxaca, Galeotti 5289. Figure
37B.
Plant epiphytic, cespitose, 6-10 cm high. Rhizome
short; roots 1 mm in diameter. Pseudobulbs elliptic-
ovoid, compressed, 5-15 mm long, 5 mm wide, sub-
tended by 5-8 foliaceous bracts; apex 1 -foliate, or leaf
aborted. Leaf thin, without articulation, linear-lanceo-
late, somewhat carinate beneath, 2-5 cm long, 6-12 mm
wide. Inflorescence a lateral raceme of 3-4 somewhat
successively borne flowers held just above the leaves;
ovary with pedicel 4-8 mm long. Flowers 1.8-3 cm tall,
1.5-2.5 cm wide, yellow; petals and lip base spotted
with brown. Sepals dissimilar; dorsal erect, concave,
ovate, ca. 5 mm long, 3 mm wide; lateral sepals shortly
connate, ca. 5 mm long, 2 mm wide. Petals similar to
the dorsal sepal, ovate to subquadrate, 5-7 mm long, 4-
5 mm wide, acute. Lip deeply 3-lobate, 1.6-2 cm long,
1.4-1.7 cm wide, 5-7 mm across the isthmus, with
somewhat crenulate margins; lateral lobes obovate 5-8
mm long, 3-5 mm wide; midlobe deeply emarginate, 4-
lobate; callus below the isthmus complex, with a more
or less A-shaped, raised, bifid lamina in front. Column
stout, ca. 4 mm long, with a conspicuous, broad pair of
wings in front; anther shortly beaked; pollinia 2 sup-
ported on viscidium and oblanceolate stipe. Capsule el-
liptic-obovoid, ca. 1 cm long; pedicel 5 mm long.
Epiphytic in premontane rain forests, often on
vines at 400-1100 m; reported to 1850 m. Flow-
ering mostly July to January, but also sporadically
throughout the year. Reported from Mexico to
Peru.
The small plant with nonarticulate, nonequitant
leaves and usually aborted leaf at the pseudobulb
apex readily distinguishes this species in Costa
Rica. It resembles Psygmorchis pusilla (L.) Dod-
son & Dressier in the flower and general habit,
but the nonequitant leaves and the presence of a
pseudobulb readily separate it from that species.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
141
Plants from Costa Rica southward seem to have
more strongly spotted petals than those from
Mexico to Nicaragua.
Oncidium dichromaticum Rchb. f., Bonplandia
3:215. 1855. TYPE: New Granada(?), Booth
(holotype: w, photo seen). Oncidium cabagrae
Schltr., Repert. Spec. Nov. Regni Veg. 9:292.
1911. TYPE: Costa Rica, "Cabagra bei Buenos
Aires," Pittier 6589 (drawing of type: AMES).
Oncidium rechingerianum Kraenzl., Pflanzenr.
IV, 50(Heft 80):202, t. 18. 1922. TYPE: Costa
Rica, Endres (holotype: w; Photo of type:
AMES). Oncidium castaneum Rchb. f. ex
Kraenzl., Pflanzenr. IV, 50(Heft 80):278. 1922.
TYPE: Costa Rica and Chiriqui, Endres (holo-
type: w, photo seen). Probably also Oncidium
costaricense Schltr., Repert. Spec. Nov. Regni
Veg. 9:30. 1910. TYPE: Terraba, 2600 m(?),
Pittier 3859. Figure 39C.
Plant epiphytic, cespitose. Rhizome short; roots < 1
mm in diameter. Pseudobulbs green spotted with dark
wine-purple, somewhat laterally flattened, 7.5-12 cm
long, 3.5-5.5 cm wide, ovoid to elliptic -oblong, sub-
tended by 2-3 foliaceous sheaths, apically 2-foliate.
Leaves oblanceolate, acute, with a prominent midvein,
conduplicate at the base, 9-28 cm long, 1.4-3.5 cm
wide. Inflorescence a lateral, slender, laxly flowered,
short-branched panicle 30-80 cm long; ovary and ped-
icel 1.7-2.5 cm long, subtended by much shorter acute
bracts. Flowers 1.8-3.3 cm high, 1.5-2.9 cm wide, col-
ors basically of two patterns, yellow with brown mark-
ings or white with purple markings. Sepals subequal, 7-
13 mm long, 4-8 mm wide, elliptic-oblong, the apex
lightly recurved, apiculate; lateral sepals somewhat fal-
cate. Petals similar to the sepals but larger, 1.2-2 cm
long, 5-1 1 mm wide, margins undulate. Lip 3-lobate,
pandurate, 1.5-2.3 cm long, 1.6-1.8 cm wide; lateral
lobes small; midlobe deeply emarginate; callus at the
base variable, usually of 4 teeth, 3 in front. Column 4-
6 mm long, with dolabriform column wings at the apex,
and with a conspicuous fleshy and smooth tabula infra-
stigmatica; pollinia 2, waxy, with stipe and viscidium.
Capsules 4 cm long; pedicel 3 cm long.
Epiphytic on small to large branches in tropical
evergreen forests at 500-1200 m of the Pacific
slope. Often in disturbed forests. Flowering Jan-
uary to June. Costa Rica and Panama.
The variability of flower size and color in O.
dichromaticum is extreme and at times confusing.
Usually the flower is yellow with dense spots of
dark brown on the sepals and petals as well as the
lip, but rose forms may be frequent (var. rosea
Hort.), and white flowers can be found. With ex-
perience it is recognized by the combination of
the 2-foliate pseudobulbs, lax inflorescence, often
waving in the wind, and the dark sepals and pet-
als.
Oncidium ensatum Lindl., Edwards's Bot. Reg.
28: misc. 17. 1842. Figure 39A. TYPE: Gua-
temala (not seen). Figure 40C.
Plant a robust, somewhat rhizomatous terrestrial to
ca. 60 cm tall excluding the inflorescence. Rhizome
short, s 3 cm between pseudobulbs; roots 1-2 mm in
diameter. Pseudobulbs ellipsoid, pale green, ca. 6-8 cm
long, 2.3-3.5 cm wide, subtended by a 4 foliaceous
sheaths lacking articulations, apically 2-foliate. Leaves
and foliaceous bracts linear-ensiform, conduplicate, car-
inate, acuminate, nonarticulate, 25-40 cm long, 2.5-4
cm wide. Inflorescence lateral, paniculate, arcuate, 1-
1.5 m long including the scape; primary branches 6-8
cm long; ovary and pedicel 1.8-2.5 cm long, subtended
by a shorter floral bract 5-6 mm long. Flowers 2.3-2.8
cm in diameter, yellow, with reddish brown on the se-
pals, petals, and callus. Sepals similar, 1.3-1.5 cm long,
4-6 mm wide, acute; dorsal elliptic-oblong, margins un-
dulate; lateral sepals somewhat larger, oblong, carinate.
Petals similar to the sepals, oblique, elliptic-oblong,
acute, 1.4-1.6 cm long, 3-5 mm wide, margins undulate.
Lip 3-lobate, subpandurate, 1.2-1.5 cm long, 1.2-1.5 cm
across the base, 1 .2 cm across the midlobe; lateral lobes
rounded, spreading, midlobe emarginate, apiculate, isth-
mus ca. 6-7 mm wide; callus fleshy, as wide or wider
than long, with 7 teeth. Column short, 5 mm long, with
undulate wings; pollinia 2, waxy.
Terrestrial on embankments in lowland, dry
tropical, semideciduous forests at 450-850 m.
Flowering March to April. Uncommon from Mex-
ico to Panama.
The usually terrestrial plants with long, ensi-
form leaves lacking articulations (at least the fo-
liaceous sheaths) and more or less typical flowers
of the genus are good diagnostic features for field
recognition.
Oncidium globuliferum Kunth, Nov. Gen. & Sp.
1:347. 1815. TYPE: New Granada, "Andium
Popayanensium . . . ," (?P). Oncidium globuli-
ferum var. costaricense Rchb. f., Gard. Chron
1678. 1871. TYPE: Costa Rica, Endres (lecto- '
type selected here: W13212). Oncidium werck-
lei Schltr., Repert. Spec. Nov. Regni Veg. Beih.
19:68. 1923. New name based on Endres (w
13212). Figure 37 A.
Plant an epiphytic vine; apparent rhizome probably a
modified, indeterminate inflorescence, with lateral
shoots spaced 6-10 cm or more apart. Pseudobulbs of
lateral shoots suborbicular to elliptic-ovoid, strongly
compressed; 1.5-3 cm long, 1.3-2.5 cm wide, subtended
by 1-3 foliaceous sheaths; apex 1 -foliate. Leaves ellip-
tic-ovate, abaxially carinate, the apical leaf with short
142
FIELDIANA: BOTANY
petiole, 5-8 cm long, 1.5-2.5 cm wide; apex acute to
obtuse; foliaceous sheaths smaller. Functional inflores-
cence formed in 2 ways, directly on rhizome, with or
without an accompanying new shoot, or within a folia-
ceous sheath at the pseudobulb base; a solitary flower;
scapes to about 3 cm long (excluding flower and pedi-
cel); ovary with pedicel 2.5-4.5 cm long, subtended by
subulate bract 3-6 mm long. Flower large, 2.5-4 cm
tall, 2-3 cm wide, brilliant yellow with tepals spotted
with brown. Sepals dissimilar, shortly unguiculate; dor-
sal elliptic, acute, apiculate, 1.2-1.5 cm long, 6-7 mm
wide; lateral sepals reflexed, oblong-elliptic, acute, 1.3—
1.6 cm, 5-6 mm wide. Petals oblong-ovate, somewhat
constricted centrally, shortly unguiculate, 1.4-1.6 cm
long, 5-6 mm wide; margins undulate; apex acute to
obtuse. Lip strongly 3-lobate, 2-3 cm long, 2.5-3.5 cm
wide (midlobe), ca. 1 cm across lateral lobes; isthmus
5-6 mm wide; midlobe deeply emarginate, forming 2
rounded lobes; callus fleshy, with 7-9 teeth. Column
erect, 8 mm long, with broad wings; anther papillose,
pollinia 2, with short stipe and viscidium.
Epiphytic in high canopies of premontane and
montane rain forests at 800-1800 m. Flowering
only in the brightest part of the canopy January
to August. Costa Rica to Ecuador; probably also
to Venezuela and perhaps to Bolivia.
The viny habit of O. globuliferum is unique to
this species, especially since the "rhizome" ap-
pears to be a modified inflorescence that produces
short vegetative shoots laterally. Nothing is
known about the base of the plant.
Reichenbach attempted to recognize the Costa
Rican plants with consistently smaller flowers as
var. costaricense, but specimens from Ecuador ex-
hibit flowers of similar size.
() IK id him isthmi Schltr., Repert. Spec. Nov. Reg-
ni Veg. Beih. 17:84. 1922. TYPE: Panama,
Sambo River, South Darien, 200 m, Pittier 5569
(drawing of type: AMES). Figure 41B.
Plant a robust, somewhat rhizomatous epiphyte to
about 75 cm excluding inflorescence; rhizome short to
about 5 cm between pseudobulbs; roots 1-2 mm in di-
ameter, some of them directed upward (trash-basket
roots). Pseudobulbs large, 9-15 cm long, 3-4.5 cm
wide, sulcate, oblong-ovoid, apically 2-3-foliate, cov-
ered at the base by 2-4 foliaceous bracts. Leaves cori-
aceous, linear-ligulate, acute, 25-35 (60) cm long, (2)4-
5 cm wide; apex acute. Inflorescence lateral, paniculate,
arcuate or pendulous, to 1.50 m long including the
scape, primary branches 8-15 cm long; ovary and ped-
icel 1.8-2.5 cm long, subtended by bracts 5-6 mm long.
Flowers 2.3-2.8 cm long, 1.8-2.5 cm wide, yellow,
barred with reddish brown on the sepals, petals, and cal-
lus. Sepals dissimilar; dorsal elliptic-oblong, acute, with
undulate margins, 1 1-12 mm long, 5-7 mm wide; lateral
sepals somewhat larger, oblong, acute, 1.2-1.5 cm long,
4-7 mm wide. Petals elliptic-oblong, oblique, subacute,
with undulate margins, 1.1-1.3 cm long, 4-5 mm wide.
Lip 3-lobate, pandurate, 1.4-1.7 cm long, 9-11 mm
across the base, 1.4-2 cm across midlobe; lateral lobes,
rounded, extended laterally; midlobe emarginate, apic-
ulate; isthmus narrow, ca. 4 mm wide with strongly re-
curved margins; callus fleshy, as wide or wider than
broad, with 7 teeth. Column short, 5 mm long, with
serrate, dolabriform wings; pollinia 2, waxy, pyriform,
with stipe and viscidium.
Epiphytic on larger branches in lowland tropi-
cal semideciduous forests at 450-850 m. Flow-
ering December to April. Southeastern Costa Rica
and Panama.
Oncidium isthmi is most similar to O. polyclad-
ium Rchb. f. ex Lindl. but has longer primary
branches in the inflorescence, and the flowers are
a more intense yellow. Also, the midlobe of the
lip is much larger with, consequently, a more con-
spicuously narrow isthmus.
Oncidium klotzschianum Rchb. f., Bot. Zeitung
(Berlin) 10:695. 1852. TYPE: Chiriqui, War-
scewicz (w). Oncidium obryzatum Rchb. f.,
Bonplandia 2:108-109. 1854. TYPE: Peru,
Warscewicz (?holotype: w, photo seen). Oncid-
ium fulgens Schltr., Repert. Spec. Nov. Regni
Veg. Beih. 17:83. 1922. TYPE: Panama, Chi-
riqui, Powell 157 (lectotype selected here: AMES
23918). Oncidium angustisepalum Kraenzl.,
Pflanzenr. IV. 50(Heft 80):203. 1922. TYPE:
Costa Rica, Endres (holotype: w, photo seen).
^.Oncidium sclerophyllum Kraenzl., Pflanzenr.
IV. 50(Heft 80):204. 1922. TYPE: Costa Rica,
San Ramon, Endres (w). Oncidium varians
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
151. 1923. TYPE: Costa Rica, La Palma, A. &
C. Brade 1095. Oncidium graciliforme C.
Schweinf., Bot. Mus. Leafl. 5:96-97. TYPE:
Panama, Chiriqui, Powell 3227 (Hort. Powell
157). Figure 39B.
Plant epiphytic, cespitose, forming small clumps to
40 cm tall excluding inflorescence. Rhizome short; roots
0.5-1.5 mm in diameter. Pseudobulbs smooth to rugose,
usually with purple spots, suborbicular to conic, suffused
and marked with dark brown, 3-8.5 cm long, 1.2-4 cm
wide, subtended by 3-5 foliaceous bracts, apex 1(2)-
foliate. Leaf base conduplicate; blade carinate beneath,
linear-lanceolate to broadly elliptic, coriaceous, 10-35
cm long, 1 .2-6 cm wide, apex acute or subacute. Inflo-
rescence an erect or arcuate raceme (when small) or
panicle 30-80 cm long including the scape with numer-
ous flowers; ovary and pedicel 1.1-2.5 cm long. Flow-
ers variable in size, 2-4 cm high, 1-3.2 cm wide, yel-
low, segments marked basally with brown. Sepals sub-
equal, free, obovate-spatulate, obtuse or truncate, cari-
nate abaxially, 9-15 mm long, 2-7 mm wide. Petals
broader than sepals, obovate-spatulate, obtuse or trun-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
143
cate, 1-1.5 cm long, 4-12 mm wide. Lip 3-lobate, 1-
2.5 cm long, 1-2.5 cm wide, midlobe deeply emarginate;
isthmus 2-5 mm wide; lateral lobes rounded; callus ba-
sal, fleshy, the basal part forming a plate with 4-5 small
teeth at the base and 3 larger teeth in the front. Column
5-6 mm long, with a pair of apical, porrect, dolabriform
wings with serrulate margins; tabula infrastigmatica with
2 fleshy projections; anther beaked; pollinia 2, waxy,
rounded, with elongate stipe and viscidium. Capsules
(only 1 specimen in fruit seen) ellipsoid, ca. 3 cm long
with beak; pedicel about 1 cm long.
Common epiphyte in premontane and lower
montane rain and cloud forests at 700-1900 m.
Flowering throughout the year. Costa Rica to
Peru.
The small-flowered forms of O. klotzschianum
may be distinguished from O. obryzatoides
Kraenzl. by the larger, paniculate inflorescences
and by the color of the tepals, which are spotted
with brown rather than stained along the mid-
veins.
The bewildering range in flower size and plant
form seems to indicate either extreme variation
with expected proliferation of synonyms, or that
there are several closely related species here.
There appears to be no pattern of size variation
according to either horizontal or vertical ranges,
nor of phenology. Indeed, marked differences in
flower size can be seen in the same herbarium
specimen. Of all the synonyms proposed above,
only O. obryzatum corresponds to the large-flow-
ered forms. The smallest-flowered extremes are
represented by the names O. sclerophyllum and
O. graciliforme. In the Manual (MO) treatment,
Dressier is accepting O. sclerophyllum as distinct,
although it is known only by the type specimen.
A rehydrated flower from the type had broad,
spatulate sepals in addition to the petals, but the
callus and column were consistent with O. klot-
schianum.
The plant accepted in Dunsterville and Garay
(Venezuelan Orchids Illustrated 6, p. 303) as O.
klotzschianum exhibits a repent habit, smaller lip
in proportion to the tepals, and shorter column
wings, features that do not agree with the type
from Panama. The illustrated plant may be O. pic-
turatum Rchb. f., described from Venezuela but
not treated in the six-volume work.
Oncidium luteum Rolfe, Bull. Misc. Inform. 172.
1893. Type locality unknown. Oncidium pittieri
Schltr., Repert. Spec. Nov. Regni Veg. 9:30.
1910. TYPE: Costa Rica, by La Palma, Pittier
10310 (drawing of type: AMES, photo seen). On-
cidium cheirophoroides Kraenzl., Pflanzenr. 80:
199, t. 1922. TYPE: Costa Rica, Endres (ho-
lotype: w, photo seen). Figure 38B.
Plant a stout, cespitose epiphyte 10-35 cm tall ex-
cluding inflorescence. Rhizome short; roots ^ 2 mm in
diameter. Pseudobulb nearly obsolete, compressed, ^ 4
cm long, subtended and concealed by 4-6 foliaceous
sheaths, apex 1 -foliate. Leaves arranged in a fan, dupli-
cate, subcoriaceous, narrowly to broadly elliptic, acute,
with conspicuous sheath below the articulation, 6-30 cm
long, 1.5-6.5 cm wide. Inflorescences 1-2 in the foli-
aceous sheath axils, mostly less than twice the longest
leaf, each a panicle with 1-2 or more orders of branch-
ing, 15-55 cm tall including the scape; branches fracti-
flex; ovary with pedicel ca. 1 cm long, subtended by
small triangular bracts < 2 mm long. Flowers greenish
yellow, with brown spot near callus. Sepals similar,
somewhat concave, oblanceolate-obovate, obtuse, 3-5
mm long, 2-3 mm wide; lateral sepals somewhat re-
flexed at the base. Petals broadly elliptic, obtuse, 4-5
mm long, 2-3 mm wide. Lip deeply 3-lobate near the
base, about 9-14 mm long, 1-1.2 cm wide; lateral lobes
winglike, suborbicular; midlobe broadly cuneate, 4-5
mm wide; isthmus ca. 2 mm wide; callus below the isth-
mus of 2 lateral and 3 central teeth. Column complex
and unusual, with a central horn at the base curved up-
ward, with a pair of antrorse hornlike wings at the apex;
anther with conspicuous beak; pollinia 2, with long stipe
about 2 mm long. Capsules ellipsoid, 1.6 cm long in-
cluding the long beak; pedicel 5-7 mm long.
Epiphytic in windy cloud forests at 1200-2500
m. Flowering October to December. Endemic to
Costa Rica.
This rare epiphyte is easily distinguished by the
large fan of leaves concealing the pseudobulb.
Flowers could be confused with those of O. chei-
rophorum Rchb. f., but the plants are much larger
with much broader and longer leaves subtending
the nearly obsolete pseudobulb.
Oncidium obryzatoides Kraenzl., Pflanzenr. IV.
50(Heft 80):240. 1922. TYPE: Costa Rica, En-
dres (w). Oncidium tetraskelidion Kraenzl.,
Pflanzenr. IV. 50(Heft 80):259. 1922. Oncidium
brenesii Schltr., Repert. Spec. Nov. Regni Veg.
19:257-258. 1923. TYPE: Costa Rica, Brenes
240 (AMES). lOncidium calyptostalix Kraenzl.,
Pflanzenr. IV. 50(Heft 80):260. 1922. TYPE:
Costa Rica, Endres (w, photo seen). Figure
39A.
Plant a small cespitose epiphyte to 8-25 cm tall; rhi-
zome short; roots to 1.5 mm in diameter. Pseudobulbs
often with purplish spots, ovoid, sulcate, somewhat com-
pressed laterally, 2.5-5 cm long, 2-3 cm wide; subtend-
ed by 2-4 foliaceous bracts; apex 1 -foliate. Leaves usu-
ally distinctly petiolate; blade subcoriaceous, carinate
beneath, elliptic-lanceolate, 5-25 cm long, 1-3 cm wide;
apex acute, apiculate. Inflorescence a lateral raceme or
144
FIELDIANA: BOTANY
few-branched panicle, 12-35 cm long; ovary with ped-
icel 1.7-2.5 cm long. Flowers variable, 1.5-3 cm high,
greenish yellow; tepals stained with brown along mid-
vein; lip brown around the callus. Sepals similar, shortly
unguiculate, obovate to spatulate, 10-12 mm long, 4-5
mm wide; apex obtuse to rounded, apiculate, carinate
abaxially; lateral sepals sometimes reflexed. Petals sim-
ilar to the sepals, somewhat constricted centrally, 10-12
mm long, 5-6 mm wide; apex obtuse to rounded, apic-
ulate. Lip deeply 3-lobate at the base, 10-17 mm long,
6-16 mm across the subtriangular lateral lobes; midlobe
deeply emarginate, forming an isthmus 3-5 mm wide;
callus fleshy between lateral lobes, with broad projection
centrally, 2 blunt lateral teeth centrally, 2 similar teeth
in front. Column complex, 8 mm long; tabula infrastig-
matica with prominent lateral margins, apical wings sub-
triangular with crenulate margins; pollinia 2, round, sup-
ported on an oblanceolate stipe and viscidium. Capsules
ellipsoid, ca. 4 cm long including elongate beak; pedicel
2-2.5 cm long.
Epiphytic on larger branches of cloud forests at
1050-1900 m. Flowering September to February.
Endemic to Costa Rica from the south side of the
Central Valley to the Monteverde area.
Oncidium obryzatoides is distinguished by the
inflorescence, which has few branches, each with
few greenish yellow flowers with centrally brown-
stained tepals. The similar O. klotzschianum
Rchb. f. has tepals distinctly spotted with brown
and often larger, deeper-yellow flowers. Hybrids
possibly exist between the two species.
Oncidium fulgens Schltr. and O. varians Schltr.
have been considered as synonyms of O. obry-
zatoides (Bot. Mus. Leafl. 6:6-7. 1938), and their
types have similarly small flowers. However, the
types of O. fulgens and O. varians have broader
brown-spotted petals consistent with small forms
of O. klotzschianum Rchb. f. (synonym, O. ob-
ryzatum Rchb. f.) rather than the narrower petals
stained with brown along the midvein as in O.
obryzatoides.
Oncidium ochmatochilum Rchb. f., Bot. Zeitung
(Berlin) 10:698. 1852. TYPE: Probably Chiri-
qui, Panama, Warscewicz s.n. (w). 1 Oncidium
chelidon Kraenzl., Pflanzenr. IV. 50(Heft 80):
216-217. 1922. TYPE: Locality unknown, Wal-
lis. Oncidium chelidonizon Kraenzl., Pflanzenr.
IV. 50(Heft 80):217. 1922. TYPE: Costa Rica,
Endres s.n. (w).
Plant a large, cespitose epiphyte to nearly 1 m with-
out inflorescence; rhizome short, roots 2-4 mm in di-
ameter. Pseudobulbs little compressed laterally, rugose,
ovoid to lanceolate, 10-15 cm long, 3.5-5 cm wide;
subtended by 6-10 foliaceous sheaths; apex usually
2(rarely l)-foliate, somewhat attenuate. Leaves condu-
plicate at the base, the blade linear-ensiform, with raised
veins abaxially, 40-60 cm long, 3-5 cm wide; apex
acute. Inflorescence lateral, paniculate, to about 3 m
long including the scape; branches fractiflex; ovary with
pedicel, 3-4 cm long, subtended by a short, triangular
bracts ca. 2 mm long. Flowers with tepals greenish with
reddish brown spots, apices greenish; lip white with yel-
low callus, with an orange spot at the base and purple
spots at and below the isthmus. Sepals similar, free, re-
flexed, abaxially carinate, linear-lanceolate, 1.5-1.6 cm
long, 3-4 mm wide; apices acute, apiculate; lateral se-
pals reflexed. Petals reflexed, elliptic ovate to lanceo-
late, carinate abaxially, 1.3-1.5 cm long, 5-7 mm wide;
apex acute, apiculate; margins undulate. Lip 3-lobate,
1.2-1.4 cm long, 1-1.2 cm wide; lateral lobes rounded,
ca. 5 mm long, 3 mm wide; midlobe anchor-shaped,
apiculate, not undulate; isthmus ca. 5 mm wide; callus
on lower half, fleshy, with 7 teeth, 4 teeth laterally and
3 apically. Column stout, 6 mm long, with conspicuous
tabula infrastigmatica, without wings; pollinia 2, sup-
ported on an elongate stipe and viscidium.
Low-density epiphyte in evergreen forests on
larger branches at 800-1400 m; usually seen as
single plants. Flowering April to August. Costa
Rica and Panama; reported from Guatemala to
Nicaragua and Andean South America.
Oncidium ochmatochilum is easily distin-
guished by the growth habit, with several long
leaves subtending the pseudobulbs, and the long
inflorescence sometimes exceeding 2 m. Vegeta-
tively, it is similar to O. panduriforme Ames &
C. Schweinf., a species with shorter inflorescences
and somewhat smaller, less complex flowers.
Oncidium cardiochilum Lindl. (1855) may be
the South American equivalent of O. ochmato-
chilum.
Oncidium panduriforme Ames & C. Schweinf.,
Sched. Orch. 8:77-78. 1925. TYPE: Costa
Rica, Cachi, Alfaro (holotype: us, photo seen).
Figure 36C.
Plant epiphytic, cespitose, forming small clumps to
about 50 cm excluding inflorescence. Rhizome short;
roots flexuous, 3-4 mm in diameter. Pseudobulbs ovoid,
6-10 cm long, 3-5 cm wide, apically 1-2-foliate, con-
cealed when young by 6-8 foliaceous sheaths. Leaves
elliptic-lanceolate, acute, coriaceous, with raised veins
abaxially, base conduplicate, 10-40 cm long, 3-5 cm
wide. Inflorescence lateral, paniculate 40-80 cm long
including the scape, arcuate, branches short, fractiflex;
ovary and pedicel 1.5-2.5 cm long. Flowers 2-2.2 cm
long, 1.7-1.9 cm wide; tepals yellowish lightly spotted
with brown; lip white turning yellow. Sepals subequal,
elliptic-lanceolate, acuminate, abaxially carinate, 10-12
mm long, 3-4 mm wide; dorsal erect; lateral sepals
lightly falcate and reflexed. Petals similar to the sepals
but broader, 10-12 mm long, 3.4-4.5 mm wide. Lip
pandurate, 3-lobate, 10-12 mm long, 8-9 mm wide,
isthmus broad, about 5-6 mm wide; lateral lobes small,
subquadrate; midlobe reniform, emarginate, apiculate;
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
145
disk with a fleshy, obscurely 3-dentate, platelike callus
basally. Column erect, without wings, 6-8 mm long;
tabula infrastigmatica with 2 fleshy, elevated, antrorse
borders; pollinia 2, waxy, rounded, with conspicuous
stipe and viscidium.
Rare epiphyte of high canopies of premontane
and lower montane cloud forests at 1400-1800
(2500) m. Flowering July to November. Costa
Rica; also reported from Panama.
Oncidium panduriforme is nearly identical in
habit to O. ochmatochilum Rchb. f., a species with
somewhat larger pseudobulbs, longer inflores-
cences, and larger, more complex flowers.
Oncidium parviflorum L. O. Williams, Amer.
Orchid Soc. Bull. 11:32-33. 1942. TYPE: Pan-
ama, Province of Cocle, region north of El Val-
le de Anton, Allen 2937 (AMES). Figure 39D.
Plant a cespitose epiphyte to about 40 cm tall; rhi-
zome short; roots ^ 2 mm in diameter. Pseudobulbs
spotted with purple, elliptic to ovoid, compressed, sul-
cate, 8-12 cm long, 3-4 cm wide, subtended by 3-4
foliaceous bracts; apex 1 -foliate. Leaves petiolate;
blades coriaceous, carinate beneath, linear lanceolate,
15-40 cm long, 2-3 cm wide; apex acute, somewhat
asymmetric. Inflorescence lateral, paniculate, 75-100
cm long, with short, compact, fractiflex primary branch-
es 1-2 cm long; ovary with pedicel ca. 6 mm long,
subtended by a subulate bract 3-4 mm long. Flowers
spreading, 1.3-1.5 cm tall; tepals brown with yellow api-
ces; lip with yellow lobes, area around callus and isth-
mus brown. Sepals dissimilar, distinctly unguiculate,
with undulate margins; dorsal obovate to spatulate,
somewhat keeled abaxially 5-6 mm long, 2-3 mm wide,
obtuse, apiculate; lateral sepals elliptic, falcate, with
prominent keel abaxially, 6-7 mm long, 2-3 mm wide,
acute. Petals similar to the dorsal sepal, unguiculate,
spatulate, 5-6 mm long, 3-4 mm wide; apex acute to
obtuse, apiculate. Lip 3-lobate from the base, 9-12 mm
long, 6-7 mm wide across lateral lobes, 4-5 mm across
midlobe; lateral lobes rounded; midlobe deeply emar-
ginate; isthmus ca. 3 mm wide; callus between lateral
lobes fleshy, minutely pubescent, with 2 lateral and 2
apical teeth. Column 2-3 mm long, with a pair of
rounded, porrect, acute column wings; anther with short
beak; pollinia 2, obovoid, with viscidium and elongate
stipe. Capsules unknown.
Epiphytic and apparently rare in premontane
rain forests at 800-1000 m. Flowering in Costa
Rica May and June; in Panama, February and
March. Northern Costa Rica to Central Panama.
Oncidium parviflorum is easily recognized by
the purple-spotted pseudobulbs, elongate inflores-
cence with very short branches, and the small
flowers.
Flowers of Costa Rican plants differ somewhat
from the type in the more slender lateral sepals,
the broader midlobe, and the markedly less pu-
bescent callus. Otherwise, the plant habit, with
long, slender inflorescence, the color pattern of
the flowers, and the large, semilunate column
wings provide a very good match for O. parviflo-
rum. The above description is based on Costa Ri-
can material.
Oncidium polycladium Rchb. f. ex Lindl., Fol.
Orch. 6(7):47. 1855. TYPE: Costa Rica (prob-
ably Panama in modern times), Veragua Chiri-
qui, Warscewicz (holotype: K; isotype: w).
Probably Oncidium guttulatum Rchb. f. ex
Lindl. TYPE: Locality unknown, from garden
at Herrenhausen (K). Oncidium tonduzii Schltr.,
Repert. Spec. Nov. Regni Veg. 9:31. 1910.
TYPE: Costa Rica, Santo Domingo de Golfe
Dulce, A. Tonduz 9891 (isotype: us; drawing of
type: AMES). tOncidium peliograma Linden &
Rchb. f., Card. Chron. 1451. 1871. TYPE: Chi-
riqui, Wallis. Figure 41 A.
Plant epiphytic, cespitose to ca. 50 cm high; rhizome
short; roots ^ 2.5 mm in diameter. Pseudobulbs ovate,
sulcate, compressed, 7-15 cm long, 1.5-4 cm wide;
apex (l)2-3-foliate, subtended by 5-6 foliaceous, soon
deciduous bracts. Leaves subcoriaceous, abaxially cari-
nate, 35-50 cm long, 1.5-6 cm wide, base conduplicate,
apex acute. Inflorescence paniculate, ^ 1 m long in-
cluding scape; primary branches somewhat fractiflex, 2-
12 cm long; ovary with pedicel 1.5-2 cm long subtended
by a subulate bract about 7 mm long. Flowers 2-2.7 cm
high, yellow, bases of tepals and lip with brown bands.
Sepals dissimilar, distinctly unguiculate, with undulate
margins; dorsal obovate to ovate, obtuse, apiculate, 9-
1 1 mm long, 4 mm wide; lateral sepals elliptic-oblong,
falcate, 11-13 mm long; 3-4 mm wide. Petals shortly
unguiculate, elliptic to subspatulate, 8-10 mm long, 4-
5 mm wide; margins undulate; apex obtuse. Lip 3-lobate
from the base, 9-13 mm long, 6-8 mm wide across the
base, 8-13 mm wide across midlobe; lateral lobes small,
rounded; midlobe reniform, emarginate; isthmus 3-4
mm wide when spread; callus fleshy, narrower than long,
with about 7 rounded teeth, 3 apical and 4 lateral, the
central tooth largest and obliquely truncate. Column 4-
5 mm long; tabula infrastigmatica with enlarged lateral
margins; column wings dolabriform; anther without
beak; pollinia 2, with stipe and viscidium. Capsules el-
lipsoid, 5 cm long including short beak; pedicel 1 cm.
Epiphytic usually in crotches between larger
branches of trees in semideciduous forests at 0-
1000 m on the Pacific slope. Flowering late Jan-
uary to April. Costa Rica and probably Panama.
Oncidium polycladium has been synonymized
with related O. baueri Lindl. from South America.
However, that species has a more sharply toothed
callus, narrower column wings, and a distinctly
beaked anther. It is easily distinguished from O.
146
FTELDIANA: BOTANY
isthmi Schltr. only with experience. In aspect, the
isthmus of O. isthmi is comparatively much nar-
rower, and the branches of the inflorescence are
statistically longer.
The original description of O. guttulatum men-
tions neither origin nor collector, but a recent
analysis of the flower shows the characteristic lip
that is longer than the lateral sepals and the callus
of O. polycladiwn.
OIK- id in m schroederianum (O'Brien) Garay &
Stacey, Bradea 1:408. 1974. Odontoglossum
schroederianum Rchb. f., nom. illeg. (not Rchb.
f. 1882), Card. Chron. 3,2:364. 1887. TYPE:
Schroeder (holotype: w, photo seen). Miltonia
schroederiana O'Brien, Gard. Chron. (2):210.
1889. Odontoglossum confusum Garay (nom.
nov.), Orchidee (Hamburg) 13:217. 1962. Mil-
tonioides confusa (Garay) Brieger & Liickel,
Orchidee (Hamburg) 34(4):131. 1983. Figure
35E.
Plant epiphytic, cespitose. Rhizome short; roots 2 mm
in diameter. Pseudobulbs elliptic-oblong, coarsely sul-
cate, 5.5-10 cm long, 2-4 cm wide, apically 1-3-foliate,
subtended when young by 4-6 foliaceous bracts. Leaves
subcoriaceous, abaxially carinate, conduplicate at the
base, acute, 10-40 cm long, 2.5-5 cm wide. Inflores-
cence a several-flowered raceme, rarely a panicle with
1-2 branches, erect or arcuate, 20-30 cm long including
the scape; ovary and pedicel 2.5-3 cm long subtended
by a much shorter, triangular bract. Flowers 5-6.3 cm
tall, 2.8-3.3 cm wide; tepals yellowish barred with choc-
olate brown, yellow-tipped; lip white turning yellow
with age, tinted rose basally. Sepals free, similar, linear-
lanceolate, acuminate; dorsal 2.5-3.2 cm long, 5-8 mm
wide; lateral sepals lightly inflexed, 3.2-3.7 cm long, 5-
8 mm wide. Petals falcate, acuminate, curved upward,
2.2-3.2 cm long, 9-12 mm wide. Lip simple, 2.8-3.2
cm long, 2.4-2.6 cm wide, united firmly to the column
base, with a single broadened reniform apical lobe; disk
with a fleshy elevated callus of 3 keels, the middle keel
bifid apically. Column short, 9 mm long, with fleshy
tabula infrastigmatica; pollinia 2, with stipe and con-
spicuous viscidium.
Epiphytic in premontane cloud forests at 900-
1800 m. Flowering July to October. Costa Rica
and Panama.
Oncidium schroederianum is easily recognized
by the chocolate-barred tepals and large, simple,
white lip stained with rose at the base.
Oncidium stenobulbon Kraenzl., Pflanzenr. IV.
50(Heft 80):281. 1922. TYPE: Costa Rica, En-
dres (holotype: w, photo seen). Oncidium nar-
anjense Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 19:259. 1923. TYPE: Costa Rica, Cerro
del Espiritu Santo a Naranjo de Alajuela, Bre-
nes 22 (lectotype selected here: AMES). Figure
40B.
Plant cespitose, epiphytic, yellowish green. Rhizome
short; roots white, 1-2 mm in diameter. Pseudobulbs
elliptic-ovoid to suborbicular, strongly flattened laterally,
3.5-12 cm long, 3.5-5 cm wide, furrowed when dry, the
base covered by 2-4 imbricate, foliaceous bracts.
Leaves sessile to short petiolate, strongly carinate abax-
ially, 1-2(3) at the pseudobulb apex; blade elliptic to
obovate, obtuse, coriaceous, conduplicate at the base, 4-
15 cm long, 2.5-5.5 cm wide. Inflorescences lateral, 1-
2 at the base of the shoot, an erect or arcuate panicle,
exceeding the leaves, 50-80 cm long; branches zigzag,
2-10 cm long, with 1-5 flowers; ovary with pedicel 1.5-
2.5 cm long. Flowers yellow, 2.5-3 cm in diameter;
tepals spotted with light brown, apices yellow; lip yel-
low with brown on isthmus. Sepals dissimilar, free with
undulate margins, elliptic-ovate, with claw at the base;
dorsal erect, 1.1-1.2 mm long, 1-2 mm wide; lateral
sepals reflexed with longer claw, subfalcate, 1.4-1.5 cm
long, 2-3 mm wide. Petals broader than the sepals, with
short claw, elliptic-ovate to lanceolate, 1-1.2 cm long,
5-6 mm wide. Lip 3-lobate, 1.4-1.7 cm long, 1-1.5 cm
wide; lateral lobes, rounded, 5-7 mm long, 4-5 mm
wide; midlobe emarginate, apiculate, with undulate mar-
gin; isthmus ca. 5 mm wide; callus fleshy, consisting of
7 teeth, 5 in one plane, 2 lower at the apex. Column
short, 5-6 mm long, with tabula infrastigmatica; column
wings somewhat rounded, the margins entire; pollinia 2,
obovate, with prominent stipe and viscidium.
Uncommon epiphyte in deciduous forests at
800-1050 m elev. Flowering November to De-
cember; April and June. Apparently endemic to
Costa Rica.
Oncidium stenobulbon closely resembles O. an-
siferum Rchb. f., a species with plants less yellow-
green, usually more elongate leaves, hatchet-
shaped column wings, and, most notably, nearly
solid, dark brown tepals. Oncidium ansiferum is
known mostly above 1 350 m. See additional notes
under that species.
Oncidium stenoglossum (Schltr.) Dressier & N.
H. Williams, Orquidea (Mex.) 4:339. 1975. Mil-
tonia stenoglossa Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:66. 1923. TYPE: Costa
Rica, Tonduz 31. Odontoglossum stenoglossum
(Schltr.) L. O. Williams, Lloydia 10:212. 1947.
Miltonioides stenoglossa (Schltr.) Brieger &
Liickel, Orchidee (Hamburg) 34:131. 1983.
Figure 36B.
Plant a cespitose epiphyte; rhizome short; roots to 2
mm in diameter. Pseudobulbs ovate, sulcate, 5-9 cm
long, 2.5-3.5 cm wide; subtended by 3-4 foliaceous
bracts; apex 1-2 foliate. Leaves coriaceous, condupli-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
147
cate, elliptic to oblong, 15-35 cm long, 2.2-3.3 cm
wide; apex bluntly acute, apiculate. Inflorescence lat-
eral, arcuate, paniculate (Costa Rican plants), 50-80 cm
long, laxly flowered; scape with short bracts and con-
spicuous nodes; primary branches 6-15 cm long; ovary
with pedicel 1-2 cm long, subtended by triangular ovate
bracts 4-6 mm long. Flowers with tepals yellowish
barred with dark green to brown; lip white turning yel-
low, with purple band(s) on lower %. Sepals similar,
elliptic oblanceolate, 1.5-1.9 cm long, 5-7 mm wide;
apex carinate abaxially, acute; lateral sepals close form-
ing an acute angle. Petals similar to the sepals, oblan-
ceolate, subfalcate, curved upward, 1.6-2 cm, 5-6 mm
wide; apex recurved, acute. Lip simple, 1.6-1.7 cm
long, 5-7 mm wide; base canaliculate clasping the col-
umn; apex dilated, fleshy, abaxially carinate; callus es-
sentially absent, base with obscurely thickened nerves.
Column stout, 4 mm long; with narrow, undulate wings;
pollinia 2, with elongate stipe and viscidium. Capsules
ellipsoid, 2 cm long, pedicel 1 cm long.
Epiphytic on larger branches in humid, shady,
cloud forests at 1400-2000 m. Flowering May to
December. Costa Rica and Panama; also reported
from Mexico to Nicaragua, but may be confused
with other species. Plants in cultivation often pro-
duce fruit, suggesting autogamy, which might ex-
plain differences seen in different regions.
We are fortunate that the basionym Miltonia
stenoglossa Schltr. is Costa Rican, because we
cannot verify heterotypic synonyms suggested
by others. The basionym Odontoglossum leu-
comelas Rchb. f. (1864) from Mexico is much
more strongly colored, but is otherwise similar
to the Costa Rican plant. The basionym Odon-
toglossum pauciflorum L. O. Williams (1954) is
probably synonymous, although the type is from
Honduras.
Oncidium stenotis Rchb. f., Linnaea 41:67.
1877. TYPE: Costa Rica, Low (holotype: w,
photo seen). Oncidium turialbae Schltr., Re-
pert. Spec. Nov. 9:32. 1911. TYPE: Costa
Rica, Turialba, Biolley (or Pittierl} 8423
(drawing of type: AMES; ?isotype: Pittier
8423, us). Figure 40D.
Plant epiphytic, somewhat climbing; rhizome dis-
tinct; roots much branched, 1-2 mm in diameter. Pseu-
dobulbs elliptic oblong, 8-15 cm long, 3-5 cm wide,
somewhat compressed laterally, ancipitous, coarsely sul-
cate, subtended by 3-6 foliaceous bracts, apex 1 -foliate.
Leaves petiolate, elliptic-oblong, subcoriaceous, base
conduplicate, strongly carinate abaxially, 20-60 cm
long, 2-6 cm wide, apex obtuse to rounded, apiculate.
Inflorescence lateral, paniculate, 1-1.5 m long with pri-
mary branches < 12 cm long; ovary with pedicel 2.5-
4 cm long. Flowers numerous, 2-3.5 cm in diameter,
usually dull yellow spotted with brown, lip bright yellow
marked at and below the isthmus with brown. Sepals
free, elliptic-lanceolate, apiculate, the margins crenulate;
dorsal 1.3-1.7 cm long, 5-7 mm wide; lateral sepals
unguiculate, strongly carinate abaxially, 1.7-2.2 cm
long, 4-7 mm wide. Petals similar to dorsal sepal, par-
allel with lateral sepals, elliptic-lanceolate, 1.3-1.7 cm
long, 5-7 mm wide, margins crenulate; apex falcate,
apiculate. Lip 3-lobate, 1.3-1.6 cm long and wide; lat-
eral lobes suborbicular; midlobe reniform, emarginate,
apiculate, margins undulate; isthmus 3-4 mm wide, less
than Vi the width of the lip; callus fleshy, with 4-6 teeth
at the base and 3 at the apex. Column 5-7 mm long,
with tabula infrastigmatica, with a pair of narrow, den-
tate, apical wings; pollinia 2, with elongate stipe and
viscidium.
Epiphytic in lowland and premontane rain for-
ests of the Atlantic slope at 50-1300 m. Flower-
ing mostly September to December; rarely to
April. Reported from Honduras to Ecuador; per-
haps also Peru.
Oncidium stenotis is distinguished by the broad
apical leaf on relatively large, sulcate pseudo-
bulbs, the elongate inflorescence with relatively
short primary branches, and the relatively large
flowers with suborbicular lateral lobes. In plant
habit it much resembles O. powellii Schltr. from
Panama, a species with much larger flowers with
nearly solid brown sepals and petals.
Oncidium storkii Ames & C. Schweinf., Sched.
Orch. 10:106. 1930. TYPE: Costa Rica, Santa
Maria de Dota, Lankester 1244 (holotype:
AMES). Figure 36D.
Plant epiphytic, cespitose. Rhizome short; roots to 2
mm in diameter. Pseudobulbs pyriform, somewhat com-
pressed, sulcate, 3-6 cm long, 2.5-4 cm wide, subtended
by 2-4 foliaceous bracts, apically 2-3-foliate. Leaves
coriaceous, elliptic-lanceolate, the larger 18-30 cm long,
2-3 cm wide, conduplicate at the base forming a short
petiole, apically bluntly acute, 2-lobate. Inflorescence a
lateral panicle, erect, 25-45 cm long including scape;
primary branches 3 or more, recurved, secund 4-6 cm
long, subtended by cymbiform bract, 3-3.5 cm long, to
about 1 cm broad when spread; ovary and pedicel 1-2
cm long, subtended by triangular bracts 1.4-2 cm long.
Flowers 3-8 per branch, 1.7-2 cm high, 1.3-1.5 cm
broad, tepals green, unspotted, lip cream turning yellow,
isthmus orange. Sepals dissimilar; dorsal erect, obovate,
margins undulate, 7.5-9 mm long, 4-6 mm wide apic-
ulate; lateral sepals shortly connate, lanceolate, carinate
abaxially, 10-12 mm long, 4-5 mm wide. Petals ob-
long-ovate, obtuse to rounded, 7.5-9.5 mm long, 5-6
mm wide. Lip sessile, pandurate, 12-14 mm long, 7-9
mm wide; lateral lobes subquadrate; midlobe 1.3—1.5 cm
wide, reniform, emarginate, and apiculate, forming a
narrow isthmus at the base; callus at the base fleshy,
elevated without apparent teeth. Column stout, 4-5 mm
long, erect, with fleshy tabula infrastigmatica, apex with
2 porrect wings; pollinia 2, waxy, with prominent stipe
and viscidium.
148
FIELDIANA: BOTANY
Rare epiphyte of premontane and montane
cloud forests at 1150-1800 m. Flowering January
to May. Endemic to Costa Rica and known only
in the southern Central Valley.
Oncidium storkii is characterized by the unusu-
al inflorescence structure, which consists of a pan-
icle of secund racemes, each with several green
and yellow, unspotted flowers. Oncidium bractea-
tum, also with similar large inflorescence bracts,
has yellow flowers with tepals spotted with dark
brown. Also, the lip is deep yellow with dark
brown around the isthmus, and there are more
pronounced lateral lobes in that species.
Oncidium teres Ames & C. Schweinf., Sched.
Orch. 8:78-80, t. 7. 1925. TYPE: Panama, Ver-
aguas, San Francisco, Powell 383 (holotype:
AMES, photo seen). Figure 35B.
Plant an erect to pendent, cespitose epiphyte; rhizome
short; roots to 2 mm in diameter. Pseudobulbs globose,
1-15 mm high, 10-12 mm wide, subtended by nonfol-
iaceous bracts, apex 1 -foliate. Leaves green tinted with
maroon, terete, linear, sulcate, acute, 30-40 cm long,
1.5-1.6 wide. Inflorescence lateral, paniculate, 40-50
cm long including the scape. Flowers with tepals green
spotted with brown, lip yellow, with reddish brown
around the callus. Sepals similar, free, concave; dorsal
suborbicular, rounded, and apiculate, 5-6 mm in diam-
eter; lateral sepals obovate to triangular, acute to obtuse,
5-6 mm long, 3-4 mm wide. Petals obovate to oblong,
obtuse to truncate and apiculate, with undulate margins,
6-8 mm long, 3-4 mm wide. Lip 3-lobate, pandurate,
1.3-1.4 cm long, 1-1.1 cm wide; callus, fleshy, without
teeth, a lunate, horizontal plate basally, with a central
rounded keel apically, and a pair of blunt lateral keels
converging at the central keel apex. Column stout, 3
mm long, with 2 subquadrate to somewhat rounded
wings apically; anther papillose, pollinia 2.
Epiphytic in the Golfito area, at 200-300 m.
Flowering in April. Costa Rica and Panama;
doubtfully reported from Nicaragua.
Oncidium warscewiczii Rchb. f., Bot. Zeitung
(Berlin) 10:693-694. 1852. TYPE: Veragua:
Chiriqui Vulkan, Warscewicz (lectotype select-
ed here: w). Figure 37C.
Plant a cespitose epiphyte 10-30 cm tall excluding
inflorescence; rhizome short; roots to 1-2 mm long.
Pseudobulb shiny, ovoid to suborbicular, little com-
pressed laterally, 4-9 cm long, 2.5-5 cm wide; subtend-
ed by 2-3 foliaceous bracts; apex 2-foliate. Leaves co-
riaceous, carinate beneath, petiolate; blades 6-30 cm
long, 1.8-3 cm wide; apex asymmetrically obtuse. In-
florescence an erect, lateral, secund raceme, somewhat
recurved apically, 25-50 cm tall including the scape;
ovary with pedicel 1-2.3 cm long, subtended and ex-
ceeded by a cymbiform, acute, and acuminate bract 1.5-
3.5 cm long. Flowers campanulate (autogamous?), gre-
garious, lemon yellow with white callus, rarely marked
with red. Sepals dissimilar; dorsal shortly unguiculate,
obovate, concave, erect, 14-16 mm long, 5-7 mm wide;
lateral sepals connate forming an emarginate blade 1.8-
2 cm long, 1-1.2 cm wide. Petals ovate, 1.4-1.6 cm
long, 8-9 mm wide, margins undulate, apex emarginate.
Lip pandurate, concave 1.8-2 cm long, 1.5-1.6 cm
wide, ca. 8 mm across lateral lobes; midlobe reniform,
emarginate; isthmus ca. 5 mm wide; callus at the base
fleshy, complex, farinose, broadened apically. Column
large, 8-11 mm long, tabula infrastigmatica lacking,
wings narrow, elongate, sometimes forming a lobe at the
base. Capsules ellipsoid, 2-3 cm long including the
beak; pedicel 1-1.5 cm long.
Epiphytic in cold, windy, montane cloud forests
at 1500-2800 m. Flowering August to December.
The quantities of capsules formed per inflores-
cence and the campanulate flowers suggest that at
least some plants are autogamous, perhaps self-
pollinated when shaken by strong winds. Costa
Rica and Panama.
Oncidium warscewiczii is easily recognized by
the shiny pseudobulbs, secund racemes with yel-
low flowers, and large, cymbiform floral bracts. It
is closely related to O. bifrons Lindl. (1857), de-
scribed presumably from Mexico, and may be
conspecific.
Excluded Taxa
Oncidium advena Rchb. f., Hamburger Garten-
Blumenzeitung 16:422. 1860. Kraenzlin in his
monograph cited this species for Costa Rica,
but no specimens have been seen. The type
from Venezuela includes a painting of the flow-
er showing a broad isthmus similar to that of
O. ensatum Lindl. or O. sphacelatum Lindl.
Oncidium lintriculus Kraenzl., Pflanzenr. IV.
50(Heft 80):282. 1922. TYPE: Costa Rica, En-
dres (w). This entity is known only from the
type, consisting of a fragment with flowers that
appear immature. The drawing published with
the original description looks similar to an im-
mature flower of O. polycladium with midlobe
not yet expanded, but the anther is clearly
beaked.
Oncidium luridum Lindl., Ed wards 's Bot. Reg.
9: t. 727. 1823. This is reported from Mexico
to Nicaragua, the Caribbean region, and north-
ern South America. It might occur in mangrove
swamps of the Caribbean coast, which have
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
149
been poorly collected, but neither herbarium
specimens nor recent observations exist to sub-
stantiate its occurrence in Costa Rica.
Osmoglossum Schltr.
(D. E. Mora de Retana)
Oncidium ornithorhynchum Kunth, Nov. Gen.
et Sp. PL 1:345, t. 80. 1815. This species was
listed for Costa Rica by Kraenzl. (Pflanzenr. IV.
50(Heft 80): 201. 1922), citing a specimen col-
lected by Endres. R. L. Dressier (pers. comm.)
has indicated that this specimen is O. klotz-
schianum Rchb. f.; however, he has also heard
of a superior form of O. ornithorhynchum said
to have originated in Costa Rica.
Oncidium planilabre Lindl., J. Hort. Soc. Lon-
don 6:59. 1851. This species is known from
southern Nicaragua (Maderas Volcano) and
from Panama southward. It may occur in Costa
Rica, but no specimens have been seen. It
should be recognized by the relatively short
plant (ca. 30 cm tall) with ovoid, 2-foliate pseu-
dobulbs and raceme of mostly brown flowers
with yellow-tipped perianth segments.
Oncidium pumilum Lindl., Edwards's Bot. Reg.
t. 920. 1825. This species is recorded for Costa
Rica from a single herbarium (SEL 10440) pre-
pared by C. Lankester from cultivated material
claimed to have been collected by A. Skutch in
the San Isidro area. Lankester often failed to
label his plants and their origins, so the plant
given him by Skutch was probably something
else, perhaps a vegetatively similar species of
Trichocentrum. Because O. pumilum is other-
wise known only from South America, it is ex-
cluded from the flora.
REFERENCES — E. S. Ayensu and N. H. Wil-
liams, Leaf anatomy of Palumbina and Odonto-
glossum subgenus Osmoglossum. Amer. Orch.
Soc. Bull. 41:687-696. 1972. F. Halbinger, Odon-
toglossum and related genera in Mexico and Cen-
tral America. Orquidea (Mex.) 8:242-282. 1982.
Plant an epiphytic, cespitose herb. Pseudobulbs
ovoid usually congested, compressed, subtended by sev-
eral, distichous, imbricate bracts, apex 1 -2-foliate.
Leaves linear-ligulate, coriaceous, flexible, basally con-
duplicate, acute. Inflorescence a lateral raceme borne at
the base of the uppermost leaf subtending the pseudo-
bulb; peduncle compressed and provided with distichous
bracts. Flowers nonresupinate, fleshy, spreading or cam-
panulate. Sepals dissimilar; dorsal free, often cucullate;
lateral free to variously connate forming a concave syn-
sepal. Petals free, similar to the sepals, membranaceous
or fleshy, usually somewhat concave. Lip sessile, porrect
or geniculate, simple, adnate to column foot, about the
same size as the sepals and petals; callus fleshy. Column
short, with short foot, apex winged; pollinia 2, waxy,
with stipe and viscidium. Fruit a capsule.
A small genus of about five species ranging
from Mexico to Ecuador. Two species are recog-
nized in Costa Rica.
The close relationship of Osmoglossum with
Palumbina cannot be disputed on the basis of veg-
etative anatomy and gross morphology as found
by Ayensu and Williams. Analysis of DNA by
Chase and Palmer suggests that related Palumbina
is a member of the Lophiaris clade, including the
mule-ear oncidiums.
Key to the Species of Osmoglossum
la. Lateral sepals about !/5 connate, inflorescence rachis straight S. convallarioides
Ib. Lateral sepals about ¥2. connate, inflorescence rachis zigzag S. egertonii
Osmoglossum convallarioides Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:148. 1923.
TYPE: Costa Rica, forests of Santa Rosa de Co-
pey, Tonduz, s.n. (drawing of type: AMES).
Odontoglossum convallarioides (Schltr.) Ames
& Correll, Bot. Mus. Leafl. 11:19. 1943. Figure
41C.
Plant epiphytic, cespitose, 30-40 cm tall. Rhizome
short; roots slender, to 2 mm in diameter. Pseudobulbs
shiny, elliptic-ovoid, sulcate, compressed, 4-8 cm tall,
1-4 cm wide, concealed at the base by 3-5 foliaceous
bracts, 1 -2-foliate apically. Leaves coriaceous, flexuous,
linear to narrowly oblanceolate, 25-42 cm long, 0.7-1.4
cm wide, forming a conduplicate petiole at the base,
apically acute. Inflorescence a lateral raceme 25-30 cm
long including the peduncle, borne in the uppermost fo-
liaceous bract; ovary and pedicel 1.5 cm long, rarely
exceeded by the subtending floral bract. Flowers 3-7,
fragrant, small, nonresupinate, campanulate, 10-13 mm
150
FIELDIANA: BOTANY
high, 5-7 mm broad, white suffused with rose; callus
and column yellowish orange. Sepals subequal, elliptic-
lanceolate, concave, 7-9 mm long, 4-5 mm wide, acute,
with a fleshy apicule; lateral sepals ca. !/5 connate. Petals
elliptic-oblanceolate, concave, 7-9 mm long, 5-6 mm
wide, apiculate. Lip sessile, adnate to column base, ob-
ovate to subpandurate, markedly narrower in lower half,
7-9 mm long, 5-6 mm wide, lightly emarginate; callus
basal, fleshy, with 2 short keels extended apically as
teeth. Column stout, 3-4 mm long, with 2 porrect wings
apically; pollinia 2, waxy, with well-developed stipe and
viscidium. Capsule ellipsoid, 2.4-3 cm long; pedicel
0.8-1 cm long.
Epiphytic on the larger, shady trunks of trees
of premontane and lower montane cloud forests
at 1400-2000 (2700) m, often forming large
masses. Flowering December to April. Reported
Mexico to Panama.
Osmoglossum convallarioides is similar to O.
egertonii (Lindl.) Schltr., which has larger flow-
ers, nearly totally connate lateral sepals, and non-
constricted lips.
Osmoglossum egertonii (Lindl.) Schltr., Orchis
10:166. 1916. Odontoglossum egertonii Lindl.,
Edwards's Bot. Reg. 31: misc. 50. 1845. TYPE:
Rucker s.n. (K). Osmoglossum acuminatum
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 17:
79-80. 1922. TYPE: Panama, Chiriqui, Powell
255 (isotype: AMES). Osmoglossum anceps
Schltr., Repert. Spec. Nov. Regni Veg. Beih. 19:
147-148. 1923. TYPE: Costa Rica, without pre-
cise locality, A. and C. Brade 1180 (isotype:
AMES). Not Odontoglossum anceps Klotzsch,
1851. Not Odontoglossum anceps Lem., 1856.
Figure 4 ID.
Plant epiphytic, cespitose, 30-40 cm tall. Rhizome
short; roots slender, to 2 mm in diameter. Pseudobulbs
shiny, elliptic-ovoid, compressed, sulcate, 6-10 cm long,
1.3-3 cm wide, subtended by 3-5 foliaceous bracts, 1-
2-foliate apically. Leaves coriaceous, flexuous, linear to
narrowly oblanceolate, the larger 25-45 cm long, 1-1.5
cm wide, forming a conduplicate petiole, acute. Inflo-
rescence a lateral raceme 30-40 cm long including the
peduncle, usually shorter than the subtending foliaceous
bract, rarely 1.2 times as long; rachis somewhat zigzag;
ovary with pedicel 1-2 cm long. Flowers small, non-
resupinate, white, callus yellowish surrounded with
brown, column yellowish. Sepals subequal, obovate,
apiculate, 15-16 mm long, 5-6 mm wide; dorsal erect,
lateral sepals about Vi connate. Petals similar to the se-
pals but smaller, obovate. Lip sessile, subrhombic,
somewhat concave, apiculate, 13-14 mm long, 5-6 mm
wide; callus fleshy, with 2 low keels converging and
ending in front as 2 erect teeth, with a broad tooth at
column base. Column stout, 5-7 mm long, with 2 por-
rect, fimbriate wings apically; anther cucullate; pollinia
2, waxy, with well-developed stipe and viscidium. Cap-
sule ellipsoid, to 3.2 cm long; pedicel to 0.8 cm long.
Epiphytic in premontane and lower montane
rain forests at 1000-1700 m (ca. 150 m in Tala-
manca). Flowering October to December. Report-
ed from Mexico to Panama.
Osmoglossum egertonii closely resembles O.
convallarioides but has larger flowers, lateral se-
pals that are connate for about half their length
(see previous description), and a lip that is slightly
narrower in the lower half. The name O. egertonii
may prove not to fit the Costa Rican plants, be-
cause the presumed type (Rucker s.n. at K) with
drawing shows a more acute lip and apparently
less connate lateral sepals. The type locality is in-
dicated as "Guatemala?" in the original description.
Otoglossum (Schltr.) Garay & Dunsterv.
(D. E. Mora de Retana)
REFERENCE — G. C. K. Dunsterville and L. A.
Garay, Venez. Orch. 111. 6:41. 1976.
Plant epiphytic, repent to somewhat viny. Pseudo-
bulbs distant, oblong-ovate, subtended by conduplicate
sheaths foliaceous or not, apex 1-2 foliate. Leaves
fleshy to subcoriaceous, articulate, conduplicate. Inflo-
rescence a lateral raceme. Flowers showy, large, flat or
somewhat concave, yellow and reddish brown. Sepals
dissimilar, spreading, ovate to obovate, obtuse, forming
a short, obtuse mentum at the base. Petals similar to the
dorsal sepal, ovate to obovate, obtuse. Lip adnate to
column foot, deflexed near the base, 3-lobate, often pan-
durate, the midlobe sometimes 2-lobate. Column short,
stout, with short, broad, recurved foot at base, winged
above; pollinia 2 attached to subquadrate stipe by very
short caudicles; viscidium small. Fruit a capsule.
A small genus of several (perhaps eight) spe-
cies ranging from Costa Rica into South America.
Only a single species is known from Costa Rica.
Otoglossum is easily recognized by the widely
separated and flattened pseudobulbs hidden by
sheathing leaves and showy flowers.
The relationships of Otoglossum were considered
by M. Chase to be with various species of Oncidium
and with Odontoglossum sect. Myanthium (Syst.
Bot. 11:477-491, 1986). More recent evidence from
DNA analysis (Chase and Palmer 1992) suggests
even more complex relationships involving Ada, As-
pasia, Brassia, and Symphyglossum.
Otoglossum chiriquense (Rchb. f.) Garay &
Dunsterv., Venezuelan Orchids 111. 6:41. 1976.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
151
Odontoglossum chiriquense Rchb. f., Bot. Zei-
tung 10:692. 1852. TYPE: Warscewicz (holo-
type: w, photo seen). Odontoglossum coronar-
ium var. chiriquense (Rchb. f.) Veitch, Man.
Orch. PI. 1:23. 1886. Figure 41E.
to September. Costa Rica and Panama; possibly
also into South America.
The very scandent habit of the plant and large,
round, reddish brown flowers with mostly yellow
lip are good field characters for recognition.
Plant epiphytic, scandent. Rhizome very elongate,
covered with rather large foliaceous bracts; roots 3-5
mm in diameter. Pseudobulbs oblong-ovoid, 4-8 cm
long, 3-4 cm wide, concealed by 2 or more foliaceous
bracts, apically 1-2-foliate. Leaves subcoriaceous, ellip-
tic-ovate, the blade 10-20 cm long, 5-8 cm wide, peti-
olate at the base, obtuse to rounded apically, apiculate.
Inflorescence a raceme of 5-10 flowers borne from the
pseudobulb base, 30-45 cm long including the scape;
ovary with pedicel 2-4 cm long. Flowers somewhat
concave, 3-4 cm high, 5.5-5.8 cm broad; sepals and
petals dull red spotted with yellow toward the recepta-
cle; lip yellow at and below the isthmus. Sepals dissim-
ilar with crisped margins; dorsal concave, suborbicular
to obovate, rounded and apiculate, thickened abaxially
near the base, 2-2.5 cm long, 2-2.3 cm wide; laterals
oblong-oblanceolate, rounded and apiculate, 2-3 cm
long, 1.5-1.7 cm wide. Petals similar to the lateral se-
pals, slightly constricted on the lower Vi, 2-2.5 cm long,
2-2.3 cm wide. Lip 3-lobate, 2.5-2.7 cm long, 1.5-1.7
cm wide; midlobe obovate, forming an isthmus, rounded
at the apex; lateral lobes much smaller than the midlobe;
callus fleshy with numerous teeth. Column erect, 1 cm
long, with small, irregularly toothed wings; pollinia 2,
obovate, supported on a viscidium and distinct stipe.
Epiphytic and often rambling over canopies of
premontane and lower montane rain forests at
1050-1800 m. R. L. Dressier reports (pers.
comm.) that on Cerro Jefe, Panama, it may be
found below 1000 m crawling about in open for-
ests where there is sufficient light. Flowering June
Pachyphyllum Kunth
(J. T. Atwood & D. E. Mora de Retana)
Plant a monopodial, few-branched epiphyte. Stems
erect, creeping or pendulous, concealed by leaf sheaths;
pseudobulbs lacking. Leaves conduplicate, distichous,
articulate, chartaceous, coriaceous to fleshy, usually el-
liptic to oblong, with variously toothed margins. Inflo-
rescence lateral, short, a few-flowered raceme; pedicel
with articulation leaving a stubble behind on abscission.
Flowers inconspicuous, green to whitish or yellowish,
never red, orange, or purple, campanulate. Sepals sim-
ilar, sometimes partially connate into a tube. Petals sim-
ilar to the sepals, often adnate to the sepals forming part
of the cup of the tube. Lip usually simple, sometimes
somewhat 3-lobate, usually with a pair of tuberculate
calli. Column simple, often with broad, dentate wings
apically; pollinia 2, with simple stipe, or with a pair of
stipes. Fruit a capsule, often 3-alate.
A Neotropical and mostly Andean genus of about
35 species. Three species are known in Costa Rica.
Pachyphyllum is clearly related to Fernandezia
Ruiz & Pavon, which differs 'in the much larger
and much more colorful flowers with more com-
plex pollinaria. Dressier (1993) places Pachy-
phyllum and Fernandezia in subtribe Oncidiinae
on the basis of similarity in plant habits with
Lockhartia and also on the similarity in pollinaria
between Fernandezia and Lockhartia.
Key to the Species of Pachyphyllum
la. Plants small and mosslike, the stems rarely exceeding 6 cm long; sepals, petals, and lip connate
into a distinct cup or tube P. hispidulum
Ib. Plants larger, the stems usually exceeding 6 cm long; sepals and petals not at all connate, not forming
a cup or tube 2
2a. Sepals > 3 mm long P. costaricense
2b. Sepals < 3 mm long P. crystallinum
Pachyphyllum costaricense (Ames & C.
Schweinf.) L. O. Williams, Lilloa 3:481. 1938.
Centropetalum costaricense Ames & C.
Schweinf., Sched. Orch. 10:110. 1930. TYPE:
Costa Rica, San Jose, near Finca La Cima,
above Los Lotes, North of El Copey, Standley
42600 (holotype: AMES, photo seen; drawing of
type: SEL).
Plant epiphytic, small and inconspicuous. Stems to
12 cm long, concealed by loose, imbricating leaf
sheaths; roots large and stout. Leaves densely disti-
152
FIELDIANA: BOTANY
chous, oblong, to 1.6 cm long, 4.5 mm wide, acute or
shortly apiculate, spreading, apparently fleshy in the
fresh plant, minutely ciliolate especially on the apical
margins. Inflorescence a lateral, short raceme of 3-4
flowers; ovary 3-alate; floral bracts funnel-shaped.
Flowers white. Sepals fleshy, rigid, oblong-lanceolate or
triangular-oblong; laterals ca. 4 mm long, 1.9 mm wide,
acute, prominently keeled on the outer surface; dorsal
sepal ca. 3.5 mm long, 1.4 mm wide. Petals oblong,
similar to the sepals, ca. 1.15 mm wide. Lip rigidly at-
tached to the short column foot, sigmoid when viewed
laterally, oblong-lanceolate, with a saccate base, ca. 3.6
mm long, 1.8 mm wide, abruptly acute, apparently
somewhat reflexed on the lower Vt, with a pair of calli.
Column short, 1.5 mm long, terminating in a broad,
fleshy, lobulate wing; apparently each pollinium with
separate stipe.
Endemic to Costa Rica at 2000-2400 m. The
holotype was collected in flower in late Decem-
ber. The above description is based on the orig-
inal description, on floral diagnoses from the
type, and on a painting of a dissection showing
separate stipes (Rodriguez, Mora, Barahona, &
Williams, Generos de orquideas de Costa Rica,
p. 255. 1986).
Pachyphyllum crystallinum Lindl., Orch. Lin-
den. 18. 1846. TYPE: Venezuela, Merida,
10,000 ft, /. Linden 686 (holotype: K, photo
seen). 1 Pachyphyllum pastii Rchb. f., Bonpl. 3:
239. 1855. Xen. Orch. 1:165, t. 58, fig. 1(1-4).
1856. TYPE: Ecuador, Jameson 439 (holotype:
w, photo seen). ? Pachyphyllum stuebelii Schltr.,
Repert. Spec. Nov. Regni Veg. Beih. 7:204.
1920. TYPE: Colombia, Tolima, Volcan de To-
lima, A. Stuebel 213 (holotype: B, destroyed;
drawing of holotype: AMES). Figure 42A.
Plant a scandent monopodial epiphyte of moss-cov-
ered shrubs and dwarfed trees. Stems to about 30 cm
long, rarely branching below, covered by the sheaths of
leaves about 5 mm apart; roots white, to 1.3 mm in
diameter. Leaves fleshy, elliptic-ovate, apiculate, to 1.5
cm long and 8 mm wide, the margins hispidulous. In-
florescence a short, lateral raceme of 3-5 flowers in the
leaf axils, supported by a peduncle to 6 mm long; ovary
with jointed pedicel to 1 .5 mm long, exceeding the sub-
tending ovate, hispidulous floral bract. Flowers minute,
light green, the perianth parts with denticulate margins.
Sepals similar, cucullate; dorsal elliptic-ovate, acute,
1.7-1.8 mm long, 0.8-1.3 mm wide; lateral somewhat
concave, 2-2.3 mm long, 1-1.3 mm wide. Petals ellip-
tic-ovate, acute, 1.4-1.8 mm long, 0.7-1.1 mm wide.
Lip narrowly ovate with the sides inflexed toward the
column when young, becoming subsaccate at the base
with age, slightly constricted just below the middle, pro-
vided with a pair of calli just below the constriction,
apex reflexed with maturity, ca. 2 mm long and 1 mm
wide in natural position. Column stout, becoming au-
riculate with maturity, ca. 1 mm long when young, ca.
1.5 mm long at maturity.
Epiphytic on shrubs in cold, wind-swept elfin
forests at 3000-4000 m. Flowering season at least
July and September and probably throughout the
year. Costa Rica, Colombia, and Ecuador.
The small flowers and dentate margins of the
separate perianth parts are good features for field
recognition with a hand lens. Louis O. Williams
used the name P. pastii Rchb. f. (Ceiba 5:251.
1956.) for these plants, a name adopted by At-
wood (Icon. PI. Trop. 14: t. 1375. 1989). We thank
C. H. Dodson for identifying this species by the
earlier name of P. crystallinum.
Pachyphyllum hispidulum (Rchb. f.) Garay &
Dunsterville, Venez. Orch. 111. 3:236. 1965.
Aeranthus hispidulus Rchb. f., Linnaea 41:31.
1877. TYPE: Colombia, S. Martha, Purdie (ho-
lotype: w, photo seen). Campylocentrum hispi-
dulum Rolfe, Orch. Rev. 11:246. 1903. Orchi-
dotypus hispidulus (Rchb. f.) Senghas, Orchi-
deen (ed. 3). 1923. 1995. Wrchidotypus mus-
coides Kraenzl., Bot. Jahrb. Syst. 37:383. 1906;
Pflanzenr. IV. 50(Heft 83):51, t. 3C. TYPE:
Peru, Province of Chota, Department of Caja-
marca, Weberbauer 4189. (^Pachyphyllum
muscoides (Kraenzl.) Schltr., Repert. Spec.
Nov. Regni Veg. 15:216. 1918. Figure 42B.
Plant an inconspicuous and usually pendent epiphyte
2-6 cm long. Stems weak, branching only at the base,
concealed by the sheaths of 10-25 leaves; roots slender,
to 1 mm in diameter. Leaves fleshy, distichous, 2-5 mm
apart on each side of the stem, the sheaths with hispi-
dulous margins and a pair of stipule-like appendages on
each side of the articulation; blades falcate-lanceolate to
elliptic, acute, diminishing in size apically, 2.5-8 mm
long, 1-2 mm wide, with hispidulous margins. Inflores-
cence axillary, 1-3-flowered, shorter than the leaves;
ovary with rudimentary pedicel about 1.5 mm long.
Flowers minute, greenish yellow; the perianth ca. Vi
connate forming a tube at the base about 1 mm long.
Sepals with free portions ovate, abaxially keeled, to
about 1 mm long, < 1 mm wide. Petals with free por-
tions ovate, subsagittate, to about 0.8 mm long, ca. 0.7
mm wide. Lip similar to the petals, with claw united to
the perianth tube, the free portion suborbicular, about 1
mm long, with a pair of rounded calli on each side below
the base of the blade. Column to about 1.2 mm long,
with a broad, cucullate anther bed; pollinia 2, with dis-
tinct stipe and viscidium. Capsule suborbicular, ca. 3
mm long, 2 mm wide.
Epiphytic on smaller branches at 2000-
3150(3500) m. Flowering at least March, May,
September and probably throughout the year. Cos-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
153
ta Rica, Panama (Chiriqui), and at least northern
Andean South America.
Plant habit and size match the type of Aeran-
thus hispidulus, and the flowers match those il-
lustrated by Dunsterville and Garay (Venez. Orch.
111. 3:237) and identified as P. hispidulum.
Of the three species of Pachyphyllum known in
Costa Rica, this is the smallest, with stems less
than 5 cm long, and is easily overlooked for a moss.
Plectrophora Focke
(D. E. Mora de Retana)
REFERENCE — M. W. Chase, Obligate twig epi-
phytism in the Oncidiinae and other Neotropical
orchids. Selbyana 10:24-30. 1987.
Epiphytic, cespitose herb. Pseudobulbs small, subor-
bicular, subtended by conduplicate foliaceous sheaths,
apex 1 -foliate. Leaves articulate, fleshy, carinate, con-
duplicate, sometimes equitant. Inflorescence lateral, ra-
cemes or panicles with successively borne flowers; ova-
ries trialate in the Costa Rican species. Flowers cam-
panulate, with conspicuous spur. Sepals subequal, the
laterals forming an external spur. Petals usually broader
than the sepals. Lip forming a spur at the base within
the sepaline spur; blade simple, often with sides em-
bracing the column. Column stout, usually without con-
spicuous wings or auricles; anther subglobose; pollinia
2, waxy with stipe and viscidium. Fruit a capsule.
A Neotropical genus of about nine species,
most of which are twig epiphytes. A single spe-
cies is known from Central America.
Little is known of the relationships of Plectro-
phora within the Oncidiinae. Chase (1986) places
it within one of two major groups of the Oncidi-
inae characterized by the usual presence of a spur
or nectary.
Plectrophora alata (Rolfe) Garay, Bot. Mus.
Lean". 21:261. 1967. Trichocentrum alatum
Rolfe. Bull. Misc. Inform. 140:197. 1898.
TYPE: Colombia, Milican (not seen). Figure
42C.
Plant epiphytic, cespitose, forming small clumps, ca.
15 cm tall. Rhizome very short; roots slender, < 1 mm
in diameter. Pseudobulbs suborbicular, apically 1 -foli-
ate, 5-8 mm long, 6-7 mm wide, concealed at the base
by 2-3 papery, triangular bracts. Leaves somewhat
fleshy, elliptic-oblong, acute, abaxially carinate, 6-8 cm
long, 1-1.3 cm wide, conduplicate at the base forming
a short petiole. Inflorescence a lateral, short raceme of
flowers opening successively; ovary and pedicel trialate,
2-2.2 cm long, subtended by much shorter, triangular,
acute bracts. Flowers 2.5-3 cm high and 1.9-2.5 cm
wide; sepals greenish white, petals white, lip white with
pale yellow-orange nectar guides. Sepals subequal, el-
liptic-lanceolate, acuminate, carinate, directed forward
but with the apex lightly recurved, 1.4-1.6 cm long, 4-
6 mm wide; lateral sepals forming an external spur 1.9-
2 cm long enclosing the spur formed by the lip. Petals
much broader than the sepals, elliptic-obovate, 1.6-1.8
cm long, 1-1.2 cm wide, embracing the lip dorsally, the
apex somewhat reflexed, retuse, apiculate; margins un-
dulate and erose. Lip narrowed at the base into a spur;
blade simple, subrhombic to ovate, adnate to the col-
umn, 1.5-1.7 cm long, 1.8-2 cm wide; erose margins
undulate, with the sides upturned forming a trumpet and
embracing the column; callus of 2 short, parallel lamel-
lae or blunt keels. Column 4-5 mm long, erect, cylin-
dric, narrowed above, without wings; anther subglobose;
pollinia 2, waxy, with stipe and prominent viscidium.
An uncommon twig epiphyte in premontane for-
ests in shady, windy habitats at 670-1400 m. Flow-
ering June and September. Costa Rica to northern
South America; also reported from Guatemala.
Plectrophora alata is easily recognized by the
light-colored, campanulate flowers with conspic-
uous spur. Unlike species of Trichocentrum, it has
a sepaline spur enclosing a spur formed by the lip.
Psychopsis Raf.
(D. E. Mora de Retana)
Cespitose epiphytic herb. Pseudobulbs ovoid to sub-
orbicular, compressed, coarsely rugose, 1 -foliate. Leaf
coriaceous, elliptic to oblong, dark green, often flushed
or spotted with purple. Inflorescence a lateral raceme or
few-branched panicle with successively borne flowers.
Flowers yellow with red-brown markings, rarely pure
yellow or yellow and white. Dorsal sepal and petals
linear, reflexed; lateral sepals falcate, much shorter and
broader than the petals. Lip 3-lobate, provided with a
complex basal callus, midlobe broad and deeply emar-
ginate. Column short, with complex auricles; pollinia
waxy, 2, with viscidium and stipe. Fruit a capsule.
A Neotropical genus of four species, only one
in Costa Rica. Psychopsis is easily distinguished
by the antenna-like dorsal sepal and petals and by
the broad, falcate lateral sepals.
Evidence from DNA analysis by Chase and
Palmer suggests that P. sanderae is related to
fleshy-leaved species of Oncidium and Trichocen-
trum.
Psychopsis krameriana (Rchb. f.) H. G. Jones,
Novosti Sist. Nyssh. Rast. 12:141. 1975. On-
cidium kramerianum Rchb. f., Allg. Gartenzei-
154
FIELDIANA: BOTANY
tung 23:9. 1855. TYPE: Ecuador, "Chimbor-
asso," Warscewicz (holotype: w, photo seen).
Oncidium papilio Lindl. var. kramerianum
(Rchb. f.) Lindl., Fol. Orch. Oncidium 56. 1855.
Papiliopsis krameriana (Rchb. f.) Morren ex
Cogn. & Marsch, PL Feuill. Ornam. 2:55. 1874.
Oncidium kramerianum var. resplendens Rchb.
f., Card. Chron. 3(3):360. 1888. TYPE: Bull
1036 (holotype: w, photo seen). Figure 42D.
Plant epiphytic, cespitose. Rhizome short; roots grayish,
1-2 mm in diameter. Pseudobulbs suborbicular, somewhat
compressed, coarsely rugose, 1.9-4 cm long, 2-3.2 cm
wide, subtended by nonfoliaceous imbricate bracts, apically
1 -foliate. Leaf coriaceous, minutely spotted with dark green
or marbled with dull red, elliptic-oblong, 15-30 cm long,
4-6 cm wide, forming a short conduplicate petiole at the
base, the apex acute. Inflorescence a lateral, erect or lightly
arcuate, successively flowering raceme about 80 cm tall
including the scape; ovary with pedicel about 2 cm long,
subtended by somewhat shorter lanceolate bracts. Flowers
large and spectacular, the dorsal sepal and petals reddish or
purplish brown, the lateral sepals and lip yellow spotted
with reddish brown and with a large yellow region on the
lip. Dorsal sepal and petals similar, linear-filiform, 5-7 cm
long, 4-6 mm wide, the apical half broadened and with
undulate margins; lateral sepals very different from the dor-
sal sepal, narrowly ovate, falcate, 4-5 cm long, 1.3-1.6 cm
wide, the apex subobtuse. Lip 3-lobate, pandurate, with
undulate margins, 3.5-5 cm long, 3.2-5 cm wide; lateral
lobes subquadrate; midlobe reniform; apex deeply emar-
ginate, contracted at the base into a narrow isthmus; callus
between lateral lobes basically of 3 ridges each terminating
in a tooth, the central ridge transversely rugose. Column
erect, complex, 9-1 1 mm long, with tabula infrastigmatica
below, the apex broadly winged and with a pair of filiform
appendages apically; pollinia 2, waxy, with fleshy stipe and
viscidium; stigma entire.
Rare epiphyte on large branches in tropical low-
land and premontane rain forests at 50-750 m.
Flowering mostly September and October; year-
round in cultivation. Costa Rica to western Ecuador.
Psychopsis krameriana is easily distinguished
by the large successive flowers borne on long
scapes and the antenna-like dorsal sepal and lat-
eral petals.
With regard to conservation, P. krameriana
probably has never been a common plant. Cur-
rently, it is an extremely rare species in Costa
Rica, known populations having been severely
decimated by commercial collectors. Wild-col-
lected plants are often hosts for a species of rust,
and few plants are allowed through quarantine
when brought into the United States. Clearly,
mass propagation of selected clones to satiate
markets are required to help curb indiscriminate
and futile overcollection from wild populations.
Psygmorchis Dodson & Dressier
(D. E. Mora de Retana)
REFERENCE — C. H. Dodson and R. L. Dressier,
Two undescribed genera in the Orchidaceae — On-
cidiinae. Phytologia 24(4):285-292. 1972.
Plant epiphytic usually on twigs, iris-like, short-lived,
without pseudobulbs. Leaves usually numerous, ensi-
form, remaining conduplicate to the apex, arranged in a
fan, without articulations. Peduncle axillary, 1-12-flow-
ered borne in succession. Flowers yellow or with red
dots, the lip proportionately large. Sepals and petals
similar, yellow, free and spreading or lateral sepals par-
tially connate. Lip strongly 4-lobed and provided with
a complex fleshy callus at the base. Column short, 2-
winged; pollinia 2, waxy, supported on a stipe and vis-
cidium. Fruit a capsule.
A Neotropical genus of perhaps six species,
with two species known in Costa Rica. The plant
habit and yellow flowers with large lip are the best
characters for field recognition.
Chase and Palmer suggest that Psygmorchis is
related to several Brazilian taxa, including Oncidium
crispum, O. pubes, and Gomesa planifolia. The
unique psygmoid plant habit and reduced chromo-
some numbers (2n =10) suggest that Psygmorchis
is basically a paedomorphic oncidium.
Key to the Species of Psygmorchis
la. Flowers yellow with reddish spots; lip ^ 10 mm wide P. pusilla
Ib. Flowers entirely yellow; lip < 9 mm wide P. pumilio
Psygmorchis pumilio (Rchb. f.) Dodson &
Dressier, Phytologia 24(4):288. 1972. Oncidium
pumilio Rchb. f., Bot. Zeitung (Berlin) 10:697.
1852. TYPE: not seen. Oncidium gnoma
Kraenzl., Pflanzenr. IV. 50(Heft 80):98-99.
1922. TYPE: Locality unknown, Sander (photo
of type: AMES). Oncidium titania Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:67. 1923.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
155
TYPE: Costa Rica, "Carillo" (Carrillo?), Werc-
kle 63 (drawing of type: AMES). lOncidium ob-
eronia Schltr., Repert. Spec. Nov. Regni Veg.
Beih. 27:113. 1924. TYPE: Colombia, Mocoa
(Territorio del Caqueta), Hopp 207. Oncidium
hondurense Ames, Bot. Mus. Leafl. l(5):31-35,
t. 1933. Tolumnia pumilio (Rchb. f.) Hoehne,
Iconografia Orch. Bras. 231. 1949. Psygmorchis
gnoma (Kraenzl.) Dodson & Dressier, Phyto-
logia 24(4):289. 1972. Figure 43A.
Plant an epiphyte with 1-3 shoots, very small, 1-3.5
cm high excluding inflorescence, rhizome short; roots <
1 mm in diameter. Leaves equitant, fleshy, 10-15 per
shoot distributed in a fan; each elliptic-lanceolate and
subfalcate in profile, 8-35 mm long, 2-5 mm deep. In-
florescences lateral, 1 -several per shoot; each a succes-
sively flowering raceme of 2-4 flowers 2-4 cm long
including the filiform peduncle; ovary with pedicel 5-7
mm long, subtended by a much shorter ovate, acute flo-
ral bract. Flowers yellow, without spots, 8-14 mm high.
Sepals dissimilar, dorsal elliptic-ovate, acuminate, 2.5
mm long, 1.7 mm wide; lateral sepals ca. Vi connate,
reflexed at the base; each linear-lanceolate, acute, 2.8-3
mm long, 0.6-0.8 mm wide. Petals obliquely ovate,
acute, 2.8-3 mm long, 2-2.3 mm wide. Lip 3-lobate,
pandurate, 5-6 mm long, 6-7 mm wide; margins un-
dulate; basal lobes broad, dolabriform; midlobe deeply
emarginate, obscurely reniform; callus fleshy, cuneate at
the base, elevated and papillose apically. Column short,
fleshy, 2-3 mm long including conspicuous dolabriform
wings beside the stigma; pollinia 2, waxy, with conspic-
uous stipe and viscidium. Capsules ca. 2 cm; pedicel 5
mm long.
Epiphytic on twigs and vines in tropical low-
land and premontane rain forests at 300-1200 m.
R. L. Dressier (pers. comm.) relates its occurrence
on cultivated azaleas; it is also common in aban-
doned coffee plantations. Flowering December to
February. Reported from Guatemala to northern
and Andean South America.
Psygmorchis pumilio appears similar to a min-
iature P. pusilla, but the flowers are entirely yel-
low, and the callus has a fimbriate margin. Plants
may flower when less than 1 cm tall.
Psygmorchis glossomystax (Rchb. f.) Dodson
& Dressier has been reported from Costa Rica. It
is similar to P. pumilio in its callus, but generally
has larger flowers with reddish brown spots, and
the lobes of the lip are more rounded. No herbar-
ium specimens identifiable as O. glossomystax
have surfaced from Costa Rica.
Psygmorchis pusilla (L.) Dodson & Dressier,
Phytologia 24:288. 1972. Epidendrum pusillum
L., Sp. PI. (ed. 2): 1352. 1763. TYPE: not seen.
Cymbidium pusillum (L.) Sw., Nov. Act. Upsal.
6:74. 1799. Oncidium pusillum (L.) Rchb. f.
Ann. Bot. Syst. 6:714. 1863. Tolumnia pusilla
(L.) Hoehne, Icon. Orch. Bras. 231. 1949.
Plant a cespitose twig epiphyte 2.5-8 cm tall, fanlike;
rhizome nearly absent; roots to 1 mm in diameter. Pseu-
dobulbs lacking, the stem concealed by leaf bases.
Leaves coriaceous to fleshy, conduplicate, distichous,
imbricate, falcate-lanceolate in profile, lacking articula-
tions, 2-8 cm long, 3-10 mm deep or high (measured
from margin to midrib). Inflorescences lateral in the leaf
axils, < 12 per shoot; each a successively flowered, con-
densed raceme (rarely with 1 branch) to 1.5 cm long
supported on a peduncle 1.5-6 cm long covered by 2-
3 compressed sheaths each producing a subulate, in-
curved blade; ovary with pedicel glabrous, 8-10 mm
long, subtended by a laterally flattened subulate bract 2-
3 mm long. Flowers bright yellow, the petals and base
of the lip spotted with brick red; callus and column usu-
ally white marked with red. Sepals dissimilar, incon-
spicuous, about 5 mm long; dorsal ovate, apiculate; lat-
eral sepals lanceolate-falcate, acute. Petals larger than
the sepals, ovate-elliptic, 6-8 mm long, 3-5 mm wide.
Lip 3-lobate but the midlobe 4-lobulate, 10-18 mm
long; 9-20 mm wide, the lateral lobes broadly rounded;
midlobe with 4 lobules similarly rounded; margins un-
dulate; callus basal, fleshy, consisting of 1 basal plate,
2 lateral plates, and 1 folded plate at the apex. Column
stout, 2.5-3.5 mm long; wings broad, coarsely dentate.
Capsules ellipsoid to obovoid, 2-3 cm long; beak short;
pedicel short.
Epiphytic mostly on twigs and vines in tropical
lowland and premontane rain forests at 0-800 m.
Flowering throughout the year. Locally common
in Costa Rica; reported throughout most of the
Neotropics.
Psygmorchis pusilla has the larger plants and
flowers among Costa Rican Psygmorchis. The
yellow flower spotted with red and nonfimbriate
callus further distinguish it.
Rhynchostele Rchb. f.
(Including the following synonyms:
Odontoglossum Kunth sect. Rhynchostele
(Rchb. f.) Halb., Cymbiglossum Halb., and
Lemboglossum Halb.)
(D. E. Mora de Retana)
REFERENCE — M. Soto Arenas, G. A. Salazar,
and A. Rojas, Nomenclatural changes in Rhyn-
chostele, Mesoglossum, and Lemboglossum (Or-
chidaceae, Oncidiinae). Orquidea (Mex.) 13:145-
152. 1993.
Epiphytic or terrestrial cespitose to rhizomatous herb.
156
FIELDIANA: BOTANY
Pseudobulbs ovoid, usually strongly compressed later-
ally, subtended by 1-4 foliaceous bracts; apex 1-2 fo-
liate. Leaves articulate, conduplicate, elliptic to lanceo-
late, chartaceous to coriaceous. Inflorescence a scapose
raceme borne laterally at the pseudobulb base, of 1-20
flowers. Flowers usually spreading, large and colorful.
Sepals lanceolate, usually attenuate. Petals often shorter
and broader than the sepals, ovate to lanceolate, atten-
uate. Lip free, with claw with bilamellate callus. Col-
umn slender, straight to slightly arcuate, broadened near
the stigma; pollinia 2, supported on a viscidium and
stipe. Fruit a capsule.
A genus of about 16 species ranging from Mex-
ico to Panama. Three species are confirmed from
Costa Rica, although there are reports of five (see
notes under R. hortensiae). The plants with char-
acteristically large and attractive flowers are valu-
able to collectors. Probably because of their value,
few plants have been preserved as herbarium
specimens, making documentation difficult.
The relationships of Rhynchostele are unresolved,
although the work of Chase and Palmer suggests
an alliance of R. cervantesii with Rossioglossum
and Ticoglosswn. To minimize confusion, we are
following R. L. Dressier (pers. comm.) and the
reference cited above in recognizing the species
included here under the genus Rhynchostele.
These have been variously accepted under the
genera Odontoglossum, Cymbiglossum, and Lem-
boglossum, and we predict more transfers in the
future.
Key to the Species of Rhynchostele
la. Inflorescence of 10-20 flowers; lip cordate, lavender R. bictoniensis
Ib. Inflorescence of 1-5 flowers; lip trulliform or triangular, whitish, suffused or marked with reddish
brown 2
2a. Inflorescence with 2-5 flowers at a time R. hortensiae
2b. Inflorescence with a single flower at a time R. stellata
Rhynchostele bictoniensis (Bateman) Soto Are-
nas & Salazar, Orquidea (Mex.) 13:147. 1993.
Cyrtochilum bictoniense Bateman in Orch.
Mex. and Guat. t. 6. 1837. Odontoglossum bic-
toniense (Bateman) Lindl. in Edwards's Bot.
Reg. 26:66. 1840. Cymbiglossum bictoniense
(Bateman) Halb., Orquidea (Mex.) 9:2. 1983
(nom. illeg.). Lemboglossum bictoniense (Bate-
man) Halb. ex Christenson, Lindleyana 3:222.
1988. Figure 43B.
Plant epiphytic, somewhat climbing. Rhizome short;
roots grayish, 3-5 mm in diameter. Pseudobulbs elliptic,
compressed, 5.5-7 cm long, 2.7-3 cm wide, subtended
by 2-3 foliaceous bracts; apex 2-3-foliate. Leaves sub-
coriaceous, elliptic-lanceolate, acute, carinate, condupli-
cate and articulated to the sheath; blade 17-38 cm long,
2.5-3.6 cm wide. Inflorescence lateral from the base of
the pseudobulb, a raceme, 25-30 cm long including the
scape; ovary with pedicel 3-5 cm long. Flowers 2-3 cm
long, 2.5-3.5 cm wide, with tepals greenish yellow spot-
ted with red, lavender lip, column red-maroon. Sepals
free, dissimilar, apically and abaxially carinate; dorsal
cucullate, elliptic-oblong, acute, apiculate, 1.6-1.8 cm
long, 9-11 mm wide; lateral subfalcate, elliptic-lanceo-
late, acute, apiculate, the margins lightly undulate, 1.9-
2.1 cm long, 6-7 mm wide. Petals similar to the sepals
but smaller, 1.7-1.9 cm long, 6-7 mm wide. Lip sub-
cordate, united by the claw to the base of the column,
the margins undulate, 2.3-2.5 cm long and broad; callus
a pair of fleshy, elevated keels at the base, with numer-
ous translucent hairs and with 2 lamellae prolonged to
the apex. Column semiterete, 1.2-1.5 cm long, apex
with a pair of porrect dolabriform wings; pollinia 2,
waxy, pyriform, with elongate stipe and viscidium.
In Costa Rica this species grows terrestrially in
humus with Rubus in montane rain forests at
2000-2500 m. Flowering according to collectors
in May, June, September, and October. Reported
from Mexico, Guatemala, El Salvador, Costa
Rica, and Panama.
Rhynchostele bictoniensis is easily recognized
by the elongate raceme with maroon-spotted
green tepals and large, subcordate, lavender lip.
Rhynchostele hortensiae (R. L. Rodr. C.) Soto
Arenas & Salazar, Orquidea (Mex.) 13:149
1993. Odontoglossum hortensiae R. L. Rodr.
C., Orquidea (Mex.) 7:150-154, fig. 1979.
TYPE: Costa Rica, Province of Cartago, Gli-
censtein sub R. L. Rodriguez 1560 (holotype:
usj; isotypes: AMES, F). Lemboglossum horten-
siae (R. L. Rodr. C.) Halb., Orquidea (Mex.) 9:
353. 1984. Figure 43C.
Plant a shortly creeping epiphyte. Rhizome short;
roots to 3 mm in diameter. Pseudobulbs elliptic-ovoid,
strongly compressed laterally, 4-8 cm long, 2.5-4 cm
wide, subtended by 2-4 foliaceous bracts, apically 1-
foliate. Leaves elliptic, softly coriaceous, acute, apicu-
late, with a prominent abaxial keel, to 8-20 cm long, 2-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
157
4 cm wide. Inflorescence a lateral, 2-5 flowered raceme
borne laterally at the base of the mature pseudobulb, 20-
45 cm long including the scape, scape compressed, part-
ly concealed by triangular, carinate bracts; ovary with
pedicel about 3 cm long. Flowers large, 5-7 cm in di-
ameter, greenish white with maroon spots; lip bordered
and spotted with dark reddish brown. Sepals free, sim-
ilar, lanceolate, attenuate, 3.5-4.5 (5.2) cm long, 8-12
mm wide, the margins smooth; dorsal erect; lateral with
incurved apices. Petals shorter and broader than the se-
pals, ovate, attenuate, with undulate margins, 3-4(5) cm
long, 1-1.4 cm wide. Lip trulliform to rhombic, united
to the base of the column by a short claw, attenuate,
2.6-3.5 cm long, 1.8-2.3 cm wide, the margins coarsely
dentate; callus at the base, fleshy, pilose to velutinous,
of 2 short, lateral keels and a longer central keel. Col-
umn subterete, wingless, with broadened apex, 1.5 cm
long; another subglobose; pollinia 2, obpyriform, with
elongate stipe and elongate viscidium.
Rhynchostele hortensiae is a species of moist
and windy montane cloud forests at 1700-2500
m. It grows upward on trunks and larger branches,
appearing repent despite the short rhizome. Flow-
ering March to May. Endemic to Costa Rica.
This species is easily recognized by the pale
yellowish green plant, the strongly compressed
pseudobulbs formed successively in one direction,
and the large, spotted flowers with trulliform lip
and dentate margins.
Both R. cordata (Lindl.) Soto Arenas & Salazar
and R. maculata (La Llave & Lex.) Soto Arenas
& Salazer have been reported from Costa Rica,
but no reliable vouchers of these species have
been seen. We believe that R. hortensiae has been
confused with both species. Rhynchostele corda-
ta, otherwise known from Mexico to Nicaragua
and reported from Venezuela, is most similar to
R. hortensiae, but the flowers have a more spotted
color pattern, and the lip is clearly basally cordate
with less dentate margins and a more poorly de-
fined red-purple border. One collection at F (Roy
W. Lent 1618) has been repeatedly identified as
R. cordata, but the lip base is clearly broadly cu-
neate with coarsely dentate margin, and the color
notes of the collector indicate the "lip edged in
the same color [brownish wine-red]." Another
specimen (C. H. Lankester s.n. at SEL) is clearly
R. cordata and indicates "Candelaria," a Costa
Rican locality, but is based on cultivated material
that could have easily been confused in cultiva-
tion. Charles Lankester is known to have imported
and grown orchids from around the world, not just
from Costa Rica, and his living plants were not
well-labeled.
The occurrence of R. maculata is based on a
report by C. Horich (Orchideologia 10:265-278.
1975), who included a photograph of R. horten-
siae (not then described) as Odontoglossum ma-
culatum. Rodrigo Escobar (Orquideologia 11:21-
49. 1976), when writing about Odontoglossum
maculatum, stated, "but a variety exists in Costa
Rica, with white, instead of yellow petals." This
probably was of R. hortensiae. Without verifiable
herbarium specimens, we cautiously exclude both
R. cordatum and R. maculatum from the flora.
Rhynchostele stellata (Lindl.) Soto Arenas & Sa-
lazar, Orquidea (Mex.) 13:151. 1993. Odonto-
glossum stellatum Lindl., Edwards's Bot. Reg.
27: misc. 9. 1841. TYPE: Mexico, Hartweg (K,
not seen) Cymbiglossum stellatum (Lindl.)
Halb., Orquidea (Mex.) 9:4. 1983. Lemboglos-
sum stellatum (Lindl.) Halb., Orquidea (Mex.)
349. 1984. Figure 43D.
Plant a cespitose epiphyte, 10-15 cm high; rhizome
short; roots to 2 mm in diameter. Pseudobulbs ovoid to
lanceolate, compressed, 2.5-8 cm long, 1.3-1.5 cm
wide, subtended by nonfoliaceous sheaths, apex 1 -foli-
ate. Leaves petiolate; blade subcoriaceous, carinate, el-
liptic to lanceolate or oblanceolate, 10-14 cm long, 1.5-
2 cm wide; apex acute. Inflorescences 1-2 per shoot,
scapose, lateral, with 1 rarely 2 successive flowers; tri-
quetrous ovary with pedicel 4-7 cm long, subtended by
a much shorter lanceolate bract 7-10 mm long. Flowers
large and attractive; sepals and petals yellowish brown
with yellow apices, lip white suffused with lavender. Se-
pals free, similar, linear elliptic to lanceolate, attenuate,
1.9-2.2 cm long, 3-4 mm wide. Petals similar to the
sepals, 1.9-2.2 cm long, 3-4 mm wide. Lip with a claw
about 8 mm long, parallel with and embracing the col-
umn; blade reflexed, simple to lightly 3-lobate, concave,
ovate to rhombic, 2.8-3.2 cm long, 1.7-2 cm wide; apex
acute to obtuse; margins coarsely dentate and undulate;
callus at the base fleshy, projected beyond the claw, as
a pair of small, lateral teeth, in front of which is an
additional, emarginate callus. Column straight, broad-
ened at the apex, lightly verrucose, 1.3-1.5 cm long;
anther shortly beaked; pollinia 2, with elongate stipe and
viscidium. Capsules from Costa Rica unknown, in one
specimen from Mexico 3.1 cm long including the beak.
Rhynchostele stellata grows in cloud forests at
about 1500 m. Flowering May and June. Reported
from Mexico to Panama (not Nicaragua) and Ven-
ezuela.
Rhynchostele stellata is easily recognized by
the single, proportionately large flower per inflo-
rescence supported on a long pedicellate, trique-
trous ovary. It is known in Costa Rica by a single
collection near Ciudad Quesada made by an in-
spector from the Costa Rican wildlife office. It
was to be expected in Costa Rica because it is
158
FIELDIANA: BOTANY
known from both northern Central America and
Panama.
Rodriguezia Ruiz & Pavon
(D. E. Mora de Retana)
Plants epiphytic, cespitose to rhizomatous, often
forming large clumps; rhizome short or elongate, cov-
ered by persistent bracts. Pseudobulbs compressed,
apex 1-2-foliate. Leaves conduplicate, coriaceous, lig-
ulate to elliptic. Inflorescences 1 -several, each a raceme
formed in the axils of the foliaceous bracts. Flowers
usually showy. Sepals dissimilar, the dorsal erect, the
lateral sepals variously connate forming a spur. Petals
subequal to the dorsal sepal. Lip usually exceeding the
sepals, entire or emarginate. Column erect, the apex di-
lated and often with a pair of auricles, the base footless;
pollinia 2, waxy. Fruit a capsule.
A Neotropical genus of about 34 species, but
with only one species in Costa Rica. In Mesoam-
erica, a second species, R. lanceolata Ruiz & Pa-
von, occurs in neighboring Panama, and R. dress-
leriana R. Gonzalez has been described from
Mexico.
Cladistic analysis based on DNA strongly sup-
ports a relationship with Scelochilus and Com-
parettia.
Rodriguezia compacta Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:144. 1923. TYPE:
Costa Rica, Atlantic coastal hills, A. & C. Brade
1309, 1310 (drawing of type: AMES, photo
seen). Figure 44 A.
Plant epiphytic, never forming large clumps, to about
15 cm tall. Rhizome short; roots rather large, to 1.5 mm
in diameter. Pseudobulbs compressed, ovoid, 1.5-3.5
cm long, 1-1.3 cm wide, nearly covered by the sheaths
of 3-6 foliaceous bracts; apex l(2)-foliate. Leaves co-
riaceous, articulate, oblong to elliptic, broadly rounded
or obtuse apically, appearing unequally 2-lobed in
pressed specimens, 3.5-15 cm long, 1.2-3 cm wide. In-
florescences lateral, 1 -several per shoot, each a raceme
of 1-5 flowers, 5-8 cm long including the peduncle,
much shorter than the leaves; ovary with pedicel to
about 1.5 cm long, each subtended by a conduplicate,
subulate bract 5-15 mm long. Flowers white to light
yellow, the lip yellow with 2 orange lines opposite the
column apex. Sepals dissimilar; dorsal oblanceolate,
1.8-3.3 cm long, 8-13 mm wide, obtuse; lateral sepals
connate forming an oblong synsepal, more or less conic-
saccate at the base concealing the nectary, bifid apically,
1.5-3 cm long, about 10 mm wide. Petals oblanceolate
to subspatulate, rounded to lightly emarginate when flat-
tened and apiculate, 2-3 cm long, 8-15 mm wide, the
margin minutely dentate. Lip cuneate, obovate to spat-
ulate when flattened, 2-3.5 cm long, 1.3-1.5 cm wide,
with a recurved spur at the base, with margins inrolled
forming a tube, emarginate apically; callus with 2 fleshy
keels. Column clavate, 1.5-1.7 cm long with a pair of
falcate auricles apically; anther with beak, pollinia 2,
with long stipe.
Epiphytic in lowland rain forests at 15-100 m;
in Nicaragua recorded to 200 m. Flowering June
to November. Locally common from Nicaragua to
Panama.
This species is easily recognized in Costa Rica
by the shape of the column with apical horns and
the synsepal forming a recurved spur.
Rossioglossum (Schltr.) Garay & Kennedy
(D. E. Mora de Retana)
REFERENCE — L. A. Garay and G. Kennedy, The
genus Rossioglossum. Orch. Dig. 40:139-143. 1976.
Plant a cespitose epiphyte; rhizome short; roots thick.
Pseudobulbs suborbicular to ovoid or obovoid; sub-
tended by nonfoliaceous bracts; apex 2-foliate. Leaves
petiolate, conduplicate. Inflorescence a lateral raceme of
3-20 flowers. Flowers large and attractive, yellow
marked variously with reddish brown. Sepals similar.
Petals usually broader than the sepals. Lip 3-lobate; cal-
lus fleshy, with several teeth. Column stout, with lan-
ceolate wings apically; pollinia 2, supported on a stipe
and viscidium. Fruit a capsule.
A small Mesoamerican genus of five to six spe-
cies, one in Costa Rica. Relationships are unclear
but are apparently with Ticoglossum and possibly
Palumbina and Rhynchostele.
Rossioglossum schlieperianum (Rchb. f.) Garay
& Kennedy, Orch. Dig. 40:143. 1976. Odonto-
glossum schlieperianum Rchb. f., Gard. Chron.
1082, (fig.). 1865. TYPE: Costa Rica, IWend-
land (w, not seen). Figure 44B.
Plant an erect, cespitose epiphyte to about 40 cm tall;
rhizome short; roots < 4 mm in diameter. Pseudobulbs
grayish green, biconvex, lightly sulcate, ovoid to obo-
void, 4-9 cm long, 2.5-6.5 cm wide; subtended by non-
foliaceous bracts; apex sometimes emarginate, 2(3)-fo-
liate. Leaves deep gray-green, base forming a condupli-
cate petiole; blade subcoriaceous, carinate abaxially, el-
liptic-oblanceolate, 10-30 cm long, 3-7 cm wide; apex
acute. Inflorescence a lateral raceme of 3-8 flowers 25-
40 cm high including scape; ovary with pedicel 3-6 cm
long, subtended by a membranaceous, cymbiform bract
1.5-2.5 cm long. Flowers sulfur yellow; tepals barred
with rust red on lower half; lip whitish with reddish
brown isthmus; callus yellow and red. Sepals subequal,
elliptic, acute or apiculate, margins undulate; dorsal
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
159
erect, 4.5-5.2 cm long, 1 .5-2. 1 cm wide; lateral connate
basally, subfalcate, 4.5-5.2 cm long, 1.4-1.7 cm wide.
Petals much broader than sepals, elliptic ovate to obo-
vate, 4.8-5 cm long, 2-2.3 cm wide, acute to obtuse and
apiculate, margins undulate. Lip 3-lobate, 3.1-4 cm
long, ca. 1 cm across the lateral lobes, 1 .9-2.4 cm wide
at the apex; lateral lobes rounded; midlobe obovate, con-
cave, the apex lightly to deeply emarginate; callus at the
base prominent and fleshy, of 4 diverging teeth, the api-
cal pair particularly thickened and mammillate. Column
short, erect, about 1 cm long, broadened apically and
with 2 pubescent, lanceolate auricles; pollinia 2, waxy,
with stipe about 1.5 mm long and viscidium. Capsule
5-6 (or more?) cm long.
Trunk epiphyte in moderate shade of very tall
trees in humid evergreen cloud forests at 1200-
1800 m. Flowering May to July. Costa Rica and
Panama.
Rossioglossum schlieperianum is easily distin-
guished in Costa Rica by the large yellow flowers
marked with reddish brown. Pure yellow forms
have been reported.
Excluded Taxon
Rossioglossum williamsianum Rchb. f., Card.
Chron. 16:134. 1881. TYPE: B. S. Williams.
This was reported by B. S. Williams (Orchid
Grower's Manual, 1894) as having been im-
ported from Costa Rica in a shipment with
Odontoglossum (Rossioglossum) schlieperian-
um. The report has since been repeated by oth-
ers, but no recent specimens nor reliable reports
support Williams' claim. It is most likely that
the species originated from northern Central
America and the labels were confused in culti-
vation.
Scelochilus Klotzsch
(Including Neokoehleria Schltr.)
(D. E. Mora de Retana)
REFERENCE — K. Senghas, Die Gattung Scelo-
chilus, mit einer neuen Art, Scelochilus rubriflora,
aus Peru. Orchidee (Hamburg) 38:114-123. 1987.
Cespitose, usually small, epiphytic herb. Pseudob-
ulbs small, sometimes hidden by subtending, rarely fo-
liaceous sheaths; apex 1 -foliate. Leaves conduplicate,
coriaceous. Inflorescence a lateral raceme or panicle.
Flowers colorful, small, campanulate, with conspicuous
spur. Sepals dissimilar, the laterals variously connate,
forming a single spur at the base. Petals simple, similar
to the dorsal sepal. Lip complex, elongate; base with a
pair of spurs included within the sepaline spur; apex
broadened, often with involute margins; lamina usually
adorned with 1-2 pairs of horns and variously pubes-
cent. Column usually clavate; pollinarium variable, pol-
linia 2, with short to elongate, rarely furcate stipe and
viscidium. Fruit a capsule.
A genus of perhaps 50 montane species with
small, very odd, colorful flowers superficially
similar to those of Corydalis. The center of dis-
tribution is in Andean South America with pos-
sibly two in Central America, with only one
known from Costa Rica.
Floral morphology as well as DNA evidence of
Chase and Palmer suggest that Scelochilus is re-
lated to Comparettia and Rodriguezia.
Scelochilus aureus Schltr., Repert. Spec. Nov.
Regni Veg. Beih. 19:144-145. 1923. TYPE:
Costa Rica, Tabbazo (Tablazo), A. & C. Erode
1063 (drawing of type: AMES). Figure 44C.
Plant a small, cespitose, pendent or arcuate epiphyte;
rhizome short; roots to 1 mm in diameter. Pseudobulbs
subcylindric to fusiform, compressed, 2.2-2.4 cm long,
4-6 mm wide; subtended by 2-3 subulate, nonfolia-
ceous bracts; apex 1 -foliate. Leaves coriaceous to fleshy,
petiolate; blade elliptic-lanceolate or oblanceolate, cari-
nate beneath, 8-12 cm long, 1.5-3.7 cm wide; apex re-
curved and acuminate. Inflorescence lateral, arcuate or
pendent, a raceme or panicle, 12-15 cm long including
scape; each branch with 4-6 successively borne flowers;
ovary with pedicel 8-1 1 mm long subtended by subulate
bracts ca. 5 mm long. Flowers campanulate, 1.2-1.6 cm
long, yellow to orange; petals and lip lined with wine
red. Sepals dissimilar; dorsal free, concave, oblong-ob-
ovate, obtuse, 1.2-1.4 cm long, 8-9 mm wide; laterals
connate forming a synsepal 1.4-1.6 cm long, 8-10 mm
wide, spur at base 4 mm long, apex emarginate. Petals
similar to dorsal sepal in shape and size. Lip complex,
elongate, 1.4-1.6 cm long, 4-5 mm wide, with 2 spurs
inserted within the sepaline spur; blade with a pair of
slender, acute, retrorse, hornlike auricles near the middle
and a second pair near base of apical lobe, lightly ve-
lutinous; apex suborbicular and emarginate with involute
margins. Column subterete, dilated apically, lightly ve-
lutinous ventrally, 8 mm long; pollinia 2, globose, sup-
ported on a furcate stipe and viscidium.
Rare epiphyte on small branches of windy pre-
montane and lower montane cloud forests at
1500-2000 m. Flowering season imperfectly
known, at least August to October; possibly June
to December during the rainy season. Apparently
endemic to Costa Rica.
The yellow to orange, campanulate flowers ap-
pearing similar to those of Corydalis (Fumari-
aceae) are good features for field recognition.
We exclude S. aureus from synonymy with S.
160
FIELDIANA: BOTANY
tuerckheimii Schltr., a species described from
Guatemala with smaller flowers and a narrower,
rounded midlobe. The larger flower size, emar-
ginate midlobe of lip, and the shorter notch pro-
duced by the lateral sepals agree with 5. aureus,
not 5. tuerckheimii; both are otherwise very close-
ly related species. Schlechter's descriptions of the
column and ovary ("c. 2 cm longo") in 5. aureus
match nothing seen from Costa Rica and are prob-
ably erroneous. The above description is based in
part on plants recollected in the type locality.
Sigmatostalix Rchb. f.
(D. E. Mora de Retana)
Plant a cespitose epiphyte. Rhizome short, rarely creep-
ing; roots slender. Pseudobulbs small, elliptic to oblong to
ovoid, compressed, subtended by several imbricate, folia-
ceous bracts; apex 1-2-foliate. Leaves linear to elliptic-
lanceolate, acute, acuminate or asymmetrically 2-lobate. In-
florescence a lateral raceme or panicle with few-many
flowers; usually each shoot with several inflorescences at
the same time. Flowers small, often accompanied by sev-
eral bracts beneath the floral bract; floral bracts membra-
naceous, triangular, often funnel-shaped. Sepals subequal,
spreading or reflexed, free or connate at the base. Petals
similar to the sepals. Lip conspicuous, subsessile or long-
clawed, often united to the base of the column, entire or
3-lobate, usually with fleshy callus. Column slender and
arched or stout, terete, erect, often dilated at the apex; pol-
linia 2, waxy, ovoid on a single stipe and with filiform
caudicles. Fruit a capsule.
A Neotropical genus of about 35 species; six spe-
cies are here treated for Costa Rica. The genus is
easily recognized by the usually small flowers (often
brightly colored) with bizarre shapes and usually
very slender, arcuate column and the tendency to
produce several bracts per flower. Sigmatostalix is
apparently related to several sections of Oncidium,
but little more is known of its relationships.
Key to the Species of Sigmatostalix
la. Lip with claw 2
2a. Claw of the same length as the blade of the lip; lateral lobes of lip prominent and slender . . .
5. unguiculata
2b. Claw shorter than the blade of the lip; lateral lobes neither prominent nor slender 3
3a. Claw > Y4 the width of the lip; blade reniform, lateral lobes reflexed 5. picta
3b. Claw < VA the width of the lip; blade subpandurate, with 2 teeth in front of callus
S. brownii
Ib. Lip without claw 4
4a. Lateral sepals free or shortly connate, lip suborbicular M. macrobulbon
4b. Lateral sepals ca. Va connate; lip obovate 5
5 a. Inflorescence about as long as the leaves or shorter, callus prominently 2-lobate, with a
reddish brown band at the base of the lip S. hymenantha
5b. Inflorescence much longer than the leaves, callus obscurely 3-lobate, lip without band at
base S. adamsii
Sigmatostalix adamsii Dodson, Selbyana 2:54-
56. 1977. TYPE: Ecuador, Pichincha, Santo Do-
mingo, 650 m, Dodson et al. (holotype: SEL).
Figure 46 A.
Plant a cespitose epiphyte 12-15 cm high. Rhizome
short; roots to 1 mm in diameter. Pseudobulbs elliptic-
ovoid, compressed, 1.5-3 cm long, 1.3-1.5 cm wide,
subtended by 2-3 foliaceous bracts, apically 1 -foliate.
Leaves subcoriaceous, linear-lanceolate, carinate be-
neath, acute, 10-15 cm long, 1-1.3 cm wide. Inflores-
cence a lateral, slender, many-flowered panicle longer
than the leaves, 15-20 cm long including the scape; ova-
ry with pedicel 5 mm long subtended by numerous
membranaceous, obovate bracts. Flowers white to yel-
low, the lip yellow, column white. Sepals dissimilar,
membranaceous, concave, ovate-lanceolate, acute, 2-2.2
mm long, 0.5-0.6 mm wide, the dorsal free, the laterals
at least V3 connate. Petals similar to the dorsal sepal,
ovate-lanceolate, acute, incurved, 2-2.2 mm long, 0.5-
0.65 mm wide. Lip sessile, ovate, acute, 3-4 mm long,
1.2-1.5 mm wide; margins undulate. Column rather
stout for the genus, dilated at the apex, 2 mm long, an-
ther cucullate; pollinia 2, with viscidium and stipe.
An uncommon epiphyte on twigs and smaller
branches, usually in disturbed evergreen forests.
Ecuador and possibly in Costa Rica, where it has
been seen in living private collections.
This species is easily distinguished in Costa
Rica by the slender inflorescence with numerous
brown bracts at the base of the flowers.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
161
Sigmatostalix brownii Garay, Caldasia 10:236.
1968. TYPE: Panama, Brown (AMES). Figure 45C.
Plant cespitose or shortly creeping, epiphytic, 12-15
cm high. Rhizome short; roots ca. 1 mm in diameter.
Pseudobulbs red-brown, ellipsoid-ovoid, compressed,
2-4 cm long, 1-1.5 cm wide, subtended by 2-4 folia-
ceous bracts when young, apically l(2)-foliate. Leaves
subcoriaceous, lanceolate, carinate beneath, 8-15 cm
long, 6-10 mm wide, acute. Inflorescence lateral, many-
flowered, a condensed panicle with primary branches
with a single flower, appearing as a raceme, 8-15 cm
long including the scape, subtended by a membrana-
ceous, obovate bract; ovary with pedicel 2 mm long sub-
tended by 2 or more bracts. Flowers pale green; sepals
with red-brown, transverse bars (especially the laterals),
lip purplish red. Sepals subequal, oblong to lanceolate,
carinate, 3-3.3 mm long, 1-1.5 mm wide, acute and
apiculate; dorsal concave; lateral sepals recurved and
subfalcate; margins undulate. Petals lanceolate, falcate,
2.5-2.8 mm long, to about 1 mm wide, acute. Lip ellip-
tic-ovate, 2.5-3 mm long, 2-2.3 mm wide; margins un-
dulate; callus on lower Vi, fleshy, massive, with two pro-
jecting teeth in front. Column erect, dilated apically, 2
mm long; pollinia 2, with clavate stipe and viscidium.
Uncommon epiphytes on twigs and smaller
branches, usually in disturbed evergreen forests at
50-600 m. Flowering September to December.
Costa Rica and western Panama.
Sigmatostalix hymenantha Schltr., Beih. Bot.
Centralb. 36:419. 1918. TYPE: Costa Rica,
Curillo, 300 m Werckle (AMES, a drawing). Fig-
ure 45D.
Plant cespitose, epiphytic, 12-18 cm high. Rhizome
short; roots to 1 mm in diameter. Pseudobulbs ellipsoid-
ovoid, compressed, 1.5-3.5 cm long, 1.5-2 cm wide, sub-
tended by 2-4 foliaceous bracts, apically 1 -foliate. Leaves
subcoriaceous, linear-lanceolate, carinate beneath, 10-24
cm long, 4-13 mm wide, acute. Inflorescence a lateral,
many-flowered panicle about as long as the leaves, 4-20
cm long including the scape; ovary with pedicel 5 mm
long, subtended by numerous membranaceous, obovate
bracts. Flowers white to yellow, the lip darker yellow with
red band at the base of the callus, column white. Sepals
similar, membranaceous, concave, ovate-lanceolate, acute
3-3.3 mm long, 1-1.1 mm wide; the dorsal free, the laterals
shortly connate at the base. Petals similar to the sepals,
ovate-lanceolate, concave. 3-3.2 mm long, 1-1.1 mm wide,
acute. Lip sessile, suborbicular to subquadrate, reflexed, 3-
3.2 mm long, 1.2-1.5 mm wide, acute; margins undulate;
callus fleshy, elevated, transversely 2-lobate, concave, the
cavity containing an oil in live plants. Column stout, di-
lated apically, 1.5-2 mm long; anther cucullate; pollinia 2,
with stipe and viscidium. Capsule globose, ca. 5-6 mm
long, pedicel ca. 5 mm long.
An epiphyte on twigs and smaller branches,
usually in disturbed evergreen forests at (50)350-
950 m; often seen growing on citrus, hibiscus, and
coffee. Flowering December to May. Costa Rica,
Panama, and South America.
Sigmatostalix hymenantha is easily distin-
guished by the paniculate inflorescence with a
cluster of brown bracts at the base of each flower.
Sigmatostalix macrobulbon Kraenzl., Pflanzenr.
80:307. 1922. TYPE: Costa Rica, Endres (w).
Figure 45B.
Plant cespitose, epiphytic. Rhizome short; roots
white, to 1 mm in diameter. Pseudobulbs elliptic-ovate,
strongly compressed, 3-3.5 cm long, 1-1.5 cm wide,
subtended by 2-3 foliaceous bracts when young. Leaves
subcoriaceous with conduplicate petioles; blade linear-
lanceolate to elliptic, carinate beneath, 6-15 cm long, 6-
17 mm wide, acute. Inflorescence a lateral, many-flow-
ered raceme much longer than the leaves, 5-20 cm long
including the scape; ovary with pedicel 5 mm long sub-
tended by a membranaceous, obovate bract. Flowers
greenish, pale, or bright yellow, the lip with orange-yel-
low callus, the column white. Sepals dissimilar, mem-
branaceous, the dorsal free, erect, ovate-lanceolate, 3
mm long, 1.5 mm wide, concave at the base, apiculate
at the apex; lateral shortly connate, strongly reflexed,
ovate-lanceolate, 3.5-4 mm long, 1.5-1.7 mm wide, car-
inate beneath, apiculate. Petals elliptic -ovate, 3.5-4.2
mm long, 2-2.2 mm wide, acuminate, with the margins
undulate and strongly recurved. Lip sessile, suborbicular
to subquadrate, strongly convex, 3-3.2 mm long, 5-5.1
mm wide, the apex emarginate, the margins undulate;
callus on lower Vz fleshy, elevated, round with a trian-
gular cavity containing an oil in the live plant. Column
elongate, dilated at the apex, 4 mm long; anther cucul-
late; pollinia 2, with stipe and viscidium. Capsule ellip-
soid, ca. 5 mm long; pedicel about 7 mm.
Epiphytic in tropical lowland and premontane
rain forests at (400)850-1700 m, where it usually
grows on smaller branches in windy habitats.
Flowering September to February. Costa Rica and
Panama.
Sigmatostalix macrobulbon can be easily dis-
tinguished by the flowers, which have a strongly
convex lip with fleshy, muffin-shaped callus with
triangular cavity. When held under water, the oil
droplets within the cavity float to the surface.
Sigmatostalix picta Rchb. f., Ann. Bot. Syst. 6:
859. 1864. TYPE: Ecuador, Quito, Jamieson
(w). IS. costaricensis Rolfe, Bull. Misc. Inform.
78. 1916. 5. poikilostalix Kraenzl., Pflanzenr.
IV. 50(Heft 80):310. 1922. TYPE: Costa Rica,
Endres 38, 97 (syntypes: w). Figure 45A.
Plant cespitose, epiphytic. Rhizome short; roots 1-1.2
mm in diameter. Pseudobulbs oblong, compressed, 2-3 cm
long, 1-2 cm wide, subtended by 2-4 foliaceous bracts,
162
FIELDIANA: BOTANY
apically 1 -foliate. Leaves with blades oblong-lanceolate to
elliptic, keeled abaxially, 1.5-13 cm long, 0.7-2 cm wide,
acute. Inflorescence a slender, lateral, many-flowered ra-
ceme or condensed panicle with single-flowered primary
branches, to 30 cm long; ovary with pedicel 8 mm long.
Flowers yellow with brown marks on sepals, petals, and
lip; lip apical margin yellow. Sepals ovate-lanceolate, re-
flexed, 8-11 mm long, 2-3 mm wide, acute; dorsal free,
the lateral shortly connate. Petals similar to the sepals,
ovate-lanceolate, reflexed, 8-10 mm long, 2-2.2 mm wide,
acute. Lip spreading, clawed; blade anchor-shaped, some-
what convex, 7-7.2 mm long, 7-7.2 mm wide, apex shal-
lowly emarginate, apiculate; lateral lobes recurved; callus
erect, with 3 more or less rounded teeth. Column elongate,
arcuate, apically dilated, 6 mm long; anther cucullate; pol-
linia 2, oblong with elongate stipe and viscidium. Capsule
ellipsoid, 1-1.5 cm long with beak, pedicel 3-4 mm long.
Epiphytic and uncommon in evergreen forests
at 500-1800 m. Flowering August to February.
Reported from Nicaragua to South America.
Sigmatostalix picta is easily distinguished by
the relatively large, red and yellow flowers with
long claw and recurved lateral lobes. As used
here, the name represents a complex of species in
need of revision. In a narrower sense, S. picta,
based on Ecuadorian material, may apply to any
of several South American species with differing
color patterns, in which case S. costaricensis
Rolfe is probably the next available name. Sig-
matostalix guatemalensis Schltr. may be a later
synonym (1911) but differs in flower color, which
lacks red pigmentation.
Sigmatostalix unguiculata C. Schweinf., Bot.
Mus. Leafl. 8:55-57. 1940. TYPE: Costa Rica,
Province of San Jose, vicinity of El General,
Skutch 3020 (holotype: AMES). Figure 44D.
Plant cespitose, epiphytic, to 15 cm high. Rhizome
short; roots flexuous, to 1 mm in diameter. Pseudobulbs
ellipsoid to ovoid, compressed, 1 .2-2 cm long, 1 .2-2 cm
wide, subtended by 8-10 foliaceous bracts, apically 1-
foliate. Leaves subcoriaceous, linear-oblong to elliptic,
carinate beneath, 4-7 cm long, 5-8 mm wide, apically
2-lobate. Inflorescence a lateral, many-flowered raceme
about as long as the leaves, 6-7 cm long including the
scape; ovary with pedicel 7 mm long. Flowers small,
spreading, yellow to yellow-green. Sepals similar, re-
flexed, membranaceous, ovate-lanceolate, acute, 5.8-6
mm long, 1.8-2 mm wide, acute; dorsal free; lateral
slightly oblique. Petals similar to the sepals, ovate-lan-
ceolate, oblique, 5.8-6 mm long, 1.8-2 mm wide, acute.
Lip long unguiculate, the claw linear, the blade 3-lobate,
abruptly deflexed, 6-6.1 mm long, 2-2.1 mm wide; lat-
eral lobes falcate and hornlike, fleshy, pubescent; callus
fleshy, 2-lobate at the intersection of the 3 lobes and
containing an oil. Column slender, elongate, arcuate, di-
lated apically, 5 mm long; anther cucullate; pollinia 2,
with stipe and viscidium.
Epiphytic and rare in evergreen forests at 500-
975 m. Flowering October to December. Appar-
ently endemic to Costa Rica.
Sigmatostalix unguiculata is easily recognized
by the long, slender claw and the arcuate, narrow,
porrect, lateral lobes of the lip.
Systeloglossum Schltr.
(D. E. Mora de Retana)
REFERENCE — R. L. Dressier and N. H. Williams,
The genus Systeloglossum. Amer. Orch. Soc. Bull.
39:323-329. 1970.
Epiphytic, cespitose, or rhizomatous and somewhat
creeping herb. Pseudobulbs present, strongly compressed,
subtended by 3-4 foliaceous sheaths; apex 1 -foliate.
Leaves and foliaceous sheaths subcoriaceous, conduplicate.
Inflorescence a raceme or panicle with primary branches.
Flowers green, yellowish, or suffused with purple. Sepals
dissimilar; dorsal concave; lateral connate and adnate to the
column foot apically as well as laterally. Petals united lat-
erally to base of lateral sepals. Lip simple, united with
column foot forming a nectary; blade unguiculate, subor-
bicular to ovate, with simple calli usually of a pair of swell-
ings. Column stout with broad wings; pollinia 2 supported
on a squarrose stripe and viscidium (Costa Rican species).
Fruit a capsule.
A Neotropical genus of five species native to
rain forests from Costa Rica (two species) to An-
dean South America.
Systeloglossum forms a rather isolated group
with Diadenium and Oliveriana. No evidence
from DNA studies is available to link this group
with others.
Key to the Species of Systeloglossum
la. Column foot about as long as column; floral bracts 5-6 mm long; pseudobulbs hidden by subtending
foliaceous sheaths 5. acuminatum
Ib. Column foot much shorter than column; floral bracts 2-4 mm long; pseudobulbs clearly visible . .
.5. costaricense
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
163
Systeloglossum acuminatum Ames & C.
Schweinf., Sched. Orch. 10:105-106. 1930.
TYPE: Costa Rica, La Estrella, Lankester 1091
(holotype: AMES, photo seen). Figure 46B.
Plant epiphytic, cespitose to somewhat repent; rhi-
zome segments 3-5 cm long per shoot; roots < 2 mm
in diameter. Pseudobulbs cylindric, narrow, smooth,
compressed, 2-2.6 cm long, ca. 5 mm wide, usually con-
cealed and subtended by 3-7 oblong-lanceolate folia-
ceous bracts; apex 1 -foliate. Leaf linear-elliptic, base
conduplicate, thin, 8-25 cm long, 0.6-1.5 cm wide; apex
asymmetric, attenuate. Inflorescence lateral, a raceme or
few-branched panicle, 10-30 cm long including scape,
successively flowered; ovary with pedicel ca. 1 cm long,
subtended by an acuminate floral bract 5-6 mm long.
Flowers green. Sepals somewhat fleshy, dissimilar; dor-
sal oblong-elliptic, adnate to the base of the petals, 9-
12 mm long, 3-4 mm wide, apex obtuse and thickened;
lateral sepals connate forming an abaxially 2-keeled, el-
liptic-ovate, deeply emarginate synsepal 1.1-1.8 cm
long, 4-6 mm wide, united to the end of the column
foot. Petals adnate to the base of sepals, triangular-lan-
ceolate, carinate, lightly reflexed, 7-8 mm long, 4 mm
wide, acute. Lip shortly unguiculate, forming a broad
nectary with foot, adnate to base of column; blade sub-
orbicular, convex, ca. 6 mm long, 6 mm wide, emargin-
ate; callus of 2 semiglobose protuberances below the
middle. Column erect, fleshy, with broad, crenulate
wings apically, 3 mm long, 7 mm including foot; foot 4
mm long; pollinia 2, fleshy, translucent, with squarrose
stipe and viscidium. Capsules ellipsoid, 2-3.5 cm long
including beak, pedicel ca. 1 cm long.
Epiphytic in montane rain forests at 1 100-2000
m. Flowering October to April, perhaps through-
out the year. Endemic to Costa Rica.
Systeloglossum acuminatum is distinguished
from S. costaricense Schltr. by the smaller pseu-
dobulb, the somewhat longer column foot, and the
less dense inflorescence with longer acuminate
floral bracts. Plants and flowers of S. acuminatum
are less suffused with brown or purple, and the
leaves are generally more slender.
Systeloglossum costaricense Schltr., Repert.
Spec. Nov. Regni Veg. Beih. 19:252-253. 1923.
TYPE: Costa Rica, Alto de Santiago de San Ra-
mon, Brenes 19 (drawing of type: AMES). Figure
46C.
Plant epiphytic, cespitose to distinctly repent; rhi-
zome segments < 3.5 cm long per shoot; roots < 2.5
mm in diameter. Pseudobulbs conspicuous, elliptic to
ovate, smooth, strongly compressed laterally, 3-7 cm
long, 1-2.1 cm wide, subtended by 3-6 oblong-lanceo-
late foliaceous bracts; apex 1 -foliate. Leaf brownish or
purplish green, conduplicate at the base; blade thin, 8-
25 cm long, 1-2.4 cm wide, oblong-lanceolate; apex
asymmetric, acute. Inflorescence a lateral panicle with
1 or more fractiflex, densely and successively flowered
primary branches, 15-35 cm long including the pedun-
cle; ovary with pedicel 1-1.3 cm long, subtended by
subulate bracts 2-4 mm long. Flowers green or brown-
ish, ca. 2.2 cm high, 1.3 cm wide. Sepals somewhat
fleshy, dissimilar; dorsal elliptic, obtuse, 9-13 mm long,
4 mm wide; lateral connate forming an abaxially 2-
keeled, elliptic-ovate, emarginate synsepal 1-1.8 cm
long, ca. 5 mm wide. Petals adnate to base of sepals,
triangular-lanceolate, carinate, lightly reflexed, ca. 9 mm
long, 3 mm wide. Lip obovate, convex, lightly retuse,
apiculate, ca. 6-7 mm long, 4-5 mm wide; callus of 2
rounded pads near the middle. Column erect, fleshy,
broad, 5-6 mm long, forming a nectary at the base; foot
2-3 mm long; apex with a pair of concave wings; anther
papillose; pollinia 2, fleshy, translucent, with intensely
red squarrose stipe and viscidium.
Epiphytic in lowland tropical and premontane
rain forests at (50)750-1300 m. Flowering mostly
January to April, September, and perhaps
throughout the year. Endemic to Costa Rica.
Systeloglossum costaricense differs from S. ac-
uminatum Ames & C. Schweinf. in the more ex-
posed pseudobulb, more densely flowered inflo-
rescence branches with shorter, subulate floral
bracts, the column foot less than half the column
length, the tendency for broader leaves, and the
plants suffused with purple or brown.
Ticoglossum Halb.
(D. E. Mora de Retana)
REFERENCE — F. Halbinger, Cymbiglossum, Ti-
coglossum, and Rhynchostele. Orquidea (Mex.) 8:
155-282. 1983.
Epiphytic, cespitose to rhizomatous herb, roots gray-
ish, somewhat velvety. Pseudobulbs compressed, ancip-
itous, discoid to ovoid, subtended by 2-4 foliaceous
bracts; apex 1 -foliate. Leaves conduplicate, elliptic-lan-
ceolate, petiolate, acute. Inflorescence a lateral, scapose
raceme borne at the base of the pseudobulb, with 1-5
flowers; ovaries terete, pedicellate, subtended by ovate,
acute floral bracts. Flowers showy, white or rose. Sepals
elliptic to ovate, apiculate. Petals subequal to the sepals,
usually with a short, wide claw. Lip free, with claw and
a fleshy callus. Column stout, straight; pollinia 2, conic-
reniform, with stipe and viscidium. Fruit an ellipsoid
capsule.
A small genus of but two recognized species,
both occurring in Costa Rica. Chase and Palmer
suggest that Ticoglossum is related to the Lophiar-
is clade (mule ear oncidiums) of subtribe Onci-
diinae and especially to Palumbina and Rossiog-
lossum.
164
FIELDIANA: BOTANY
Key to the Species of Ticoglossum
la. Pseudobulbs strongly compressed with sharp edges, inflorescence with 2-5 white or pink flowers
T. krameri
Ib. Pseudobulbs thick with rounded edges; inflorescence with 1-2 white flowers T. oerstedii
Ticoglossum krameri (Rchb. f.) Rodriguez ex
Halb., Orquidea (Mex.) 9:5. Odontoglossum kra-
meri Rchb. f., Card. Chron. 98, t. 1868. TYPE:
Costa Rica, Kramer sub Veitch (holotype: w, pho-
to seen). The following names are based on
white-flowered specimens (types not seen). Tico-
glossum krameri var. smithianum (Rchb. f.) E. A.
Christenson, Lindleyana 6:47. 1991. Odontoglos-
sum krameri var. smithianum Rchb. f., Gard.
Chron. 19(ser. 1):242. 1883. Odontoglossum kra-
meri var. album Rolfe, Orch. Rev. 1:200. 1893.
Ticoglossum krameri var. album (R. L. Rodr. C.
ex Halb.) Halb. Odontoglossum krameri var. al-
bum R. L. Rodr. C. ex Halb. Orquidea (Mex.) 8:
186. 1982. Figure 46D.
Plant epiphytic, cespitose to 25(30) cm high. Rhi-
zome short; roots to 2 mm in diameter. Pseudobulbs
ovate-elliptic, strongly compressed laterally edges sharp,
3-5 cm long, to 4 cm wide, subtended by 3-4 subulate,
nonfoliaceous bracts, 1 -foliate apically. Leaves coria-
ceous, the base with petiole to about 4-5 cm long; blade
elliptic-oblong, 13-22(28) cm long, 1.5-4 cm wide, the
apex acute, somewhat apiculate. Inflorescence a lateral
raceme of 2-4(5) flowers, 20-25 cm long including the
scape; ovary with pedicel (3.5)4-5 cm long. Flowers
white or light lavender with yellow callus, the dorsal
sepal with yellow spot at the base. Sepals subequal, ses-
sile, elliptic-ovate, apiculate, 1.3-1.7 cm long, 5-7 mm
wide. Petals elliptic-oblong, united to the base of the
column with a short, broad claw, 1.5-2 cm long, 8-10
mm wide; apex obtuse, lightly apiculate. Lip suborbi-
cular to trapezoid, somewhat concave, about 1.5 cm
long, 1.4 cm wide; apex emarginate; callus basal, some-
what elevated, base with 2 projections, apex with 2
teeth. Column stout, 8-9 mm long, somewhat broad-
ened apically, without wings, but variously auriculate;
pollinia 2, ovate, waxy, with stipe and viscidium. Capsule
ellipsoid, without beak, 5 cm long, pedicel 3 cm long.
Epiphytic in premontane and montane cloud
forests at 850-1100 m, but Halbinger reports
600-1200 m. Flowering February to September
and November. Costa Rica; also reported from
Nicaragua and Panama.
This species is easily distinguished from high-
er-elevation T. oerstedii (Rchb. f.) R. L. Rodri-
guez ex Halb. by the usually 3-5 -flowered inflo-
rescence and flowers with lip usually shorter than
the sepals. Even when not in flower, the strongly
ancipitous pseudobulbs and relatively short peti-
ole easily distinguish T. krameri from T. oerstedii.
Ticoglossum oerstedii (Rchb. f.) R. L. Rodr. C. ex
Halb., Orquidea (Mex.) 9:5. 1983. Odontoglos-
sum oerstedii Rchb. f., Bonplandia 3:214. 1855.
TYPE: Costa Rica, Volcan Irazu, Warscewicz 689
(holotype w, photo seen). Figure 47A.
Plant epiphytic, occasionally terrestrial, cespitose
herbs. Rhizome short; roots to 2 mm in diameter. Pseu-
dobulbs ovoid, suborbicular, thick 1-3.5 cm tall, 1-1.5
cm wide, subtended by 2-4 nonfoliaceous bracts, api-
cally 1 -foliate. Leaves with petiole to ca. 1/2 the blade
length; blade coriaceous, elliptic, acute, strongly keeled
abaxially, 3-13 cm long, 1-3 cm wide. Inflorescences
lateral, with 1 (rarely to 3) flowers, the scape 2-5 cm
tall; ovary with pedicel 3-5 cm long, much longer than
the subtending acute floral bract 6-12 mm long. Flowers
white, callus spotted with orange, lip with orange in
front of callus. Sepals similar, elliptic, 9-16(20) mm
long, 6-9 mm wide, acute to obtuse. Petals obovate,
obtuse, 1-1.7(1.9) cm long, 5-8(14) mm wide. Lip ba-
sally adnate to the column, obovate-flabellate, cuneate,
deeply emarginate, 2-2.5(2.7) cm long, 1.2-1.6(1.8) cm
wide, with an elevated W-shaped callus, the central point
pubescent. Column stout, 6 mm long; pollinia 2, with
long stipe and viscidium. Capsule ellipsoid, with short
beak, 2-2.5 cm long with pedicel 1.8-3 cm long.
Epiphytic in premontane and lower montane
rain forests at 1400-3000 m, where it grows on
somewhat shaded median branches or in duff in
deep shade beneath bamboos. Flowering throughout
the year. Costa Rica; also reported from Panama.
Ticoglossum oerstedii differs from related T.
krameri (Rchb. f.) R. L. Rodr. C. ex Halb. in the
less compressed pseudobulbs, long petiolate leaf,
and the single (rarely two), differently shaped
flowers of the inflorescence. Flower size is ex-
tremely variable. One herbarium specimen (AMES
32911) includes one flower more than 5 cm tall.
Apparently there is little or no overlap in the ver-
tical distribution of the two species.
Trichocentrum Poeppig & Endl.
(F. Pupulin & D. E. Mora de Retana)
Herb a cespitose epiphyte with reduced rhizomes.
Pseudobulbs usually inconspicuous, ovoid to suborbi-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
165
cular, generally concealed by scarious bracts. Leaves
single, occasionally in pairs, articulate, sessile, coria-
ceous to fleshy, ovate, oblong-elliptic to ligulate. Inflo-
rescence a lateral raceme or few-branched panicle with
1-many flowers, often successive. Flowers often large,
showy, and sometimes fragrant. Sepals and petals free,
similar, spreading. Lip adnate to the column base pro-
ducing a conic or gibbous spur; blade with or without
calli or lobules near the base. Column short, stout, ad-
nate to the lip, auriculate or winged above, without a
foot; pollinia 2, on a triangular stipe with reflexed mar-
gins. Fruit a capsule.
A Neotropical genus of about 25 species dis-
tributed from southern Mexico to Brazil, Bolivia,
and Peru. Ten species are reported from Central
America, eight of which occur in Costa Rica.
Cladistic analysis of data based on DNA (Chase
& Palmer, 1992) strongly suggests close relation-
ships within the Lophiaris clade, including Oncid-
ium sect. Cebolletae Lindl. and O. sect. Pluritu-
berculata Lindl.
Key to the Species of Trichocentrum
la. Spur conic or cylindric, not lobed 2
2a. Spur > 5 mm long; column wings with longitudinal brown stripes T. dianthum
2b. Spur < 5 mm long; column wings white with brown spots on margins 3
3a. Lip flabellate, with very crisped apical margin; lateral lobes of lip digitate .... T. pfavii
3b. Lip cuneate, somewhat obovate, fleshy and with plain apical margin; lateral lobes short
and rounded T. estrellense
Ib. Spur 2- or 4-lobed 4
4a. Anther cap glabrous T. caloceras
4b. Anther cap papillose or hirsute 5
5a. Column wings obtuse with incurved apices T. costaricense
5b. Column wings with ascending or porrect apices, acute, usually somewhat revolute, never
incurved 6
6a. Inflorescence erect to spreading, flowers small, tepals < 6 mm long; lip lightly nar-
rowed in the middle T. brenesii
6b. Inflorescence pendent, flowers medium to large, tepals ^ 15 mm long, lip never nar-
rowed in the middle 7
7a. Lip shorter than sepals, elliptic-ovate T. capistratum
7b. Lip longer than sepals, broadly rhombic T. cymbiglossum
Trichocentrum brenesii Schltr., Repert. Spec.
Nov. Regni Veg. Beih. 19:248. 1923. TYPE:
Costa Rica, San Pedro de San Ramon, 1200 m,
Brenes 116 (drawing: AMES). Figure 48C.
Plant epiphytic, small, to ca. 10 cm high. Rhizome
short; roots filiform, flexuous, glabrous. Pseudobulbs
nearly obsolete, scarcely 3 mm long, 1-foliate. Leaf sub-
erect, obliquely ligulate, subacute, abruptly subpetiolate-
narrowed toward the base, to 9 cm long, 1.5 cm wide.
Inflorescence single, erect-spreading or spreading, with
a single flower; peduncle concealed by a few sheaths at
the base, the rest naked; ovary with pedicel glabrous,
1.2 cm long, floral bract ovate, shortly acuminate, ^ %
as long as the ovary. Flowers generally small, glabrous,
thin in texture. Sepals ligulate, acute, 3-nerved, 6 mm
long; lateral sepals oblique. Petals obliquely ligulate,
somewhat acute, 3-nerved, 5 mm long. Lip oblong, sub-
obtuse, somewhat narrowed in the middle, rounded at
the base, smooth, glabrous, with short subtruncate spur
6 mm long, 2.75 mm wide at the base, 3.25 mm wide
above the middle. Column short, 2 mm long, with 2
upward, obliquely subfalcate-elliptic, apiculate, entire
wings; anther dorsally densely papillose-verruculose,
rounded-cucullate.
Epiphytic in the moist premontane forest of
Cordillera de Tilaran, San Ramon, and San Jeron-
imo at 1000-1200 m. Flowering time incomplete-
ly known; at least September (type). Endemic to
Costa Rica, where it has been collected only three
times. The above description is translated from
Schlechter's original.
Although the type of T. brenesii was destroyed,
a copy of Schlechter's drawing is kept at AMES
and clearly shows the critical characters of the
species: the lip slightly narrowed in the middle,
not emarginate, and not longer than the sepals; the
narrow sepals; and the apiculate column wings.
Lip shape should prevent any confusion with T.
capistratum, which has an elliptic-ovate, concave,
carinate lip with undulate margins, whereas T.
brenesii has a perfectly flat lip, lacking any keel
or callosity.
166
FIELDIANA: BOTANY
Trichocentrum caloceras Endres & Rchb. f.,
Card. Chron. 1257. 1871. TYPE: Costa Rica,
Endres, not found (neotype: usi Pupulin & Cas-
telfranco 1; designated in Lindleyana 10:188.
1995). Figure 48A.
Plant a cespitose, pendent epiphyte. Rhizome short;
roots filiform, glabrous. Pseudobulbs minute, rounded,
1 -foliate, 2-2.5 mm long. Leaf fleshy, green spotted
with purple, sessile, oblong to oblong-elliptic, 7 cm
long, 1.4 cm wide, bluntly acute, somewhat minutely
mucronate. Inflorescence a raceme 3-3.5 cm long in-
cluding the terete scape and short, zigzag rachis, suc-
cessively 5-6-flowered; floral bracts conspicuous, im-
bricate, ovate, cucullate, 4 mm long, acute; ovary with
pedicel 8-10 mm long. Flowers small, spreading, with
yellowish brown tepals; lip white marked with large pur-
ple blotches; anther white. Sepals dissimilar, elliptic-lan-
ceolate, bluntly acute; dorsal erect, slightly concave, 1.1
cm long, 4 mm wide; laterals somewhat oblique, 1.2 cm
long, 3 mm wide, apically carinate. Petals oblong-ellip-
tic, obtuse, 1.2 cm long, 4-5 mm wide. Lip ovate-pan-
durate, adnate to the column base, carinate, 1 .6 cm long,
8 mm wide; base cuneate forming a short, subquadrate,
thickened spur of 4 lobes; apex retuse; callus of 2 thick
keels converging at the base. Column 5 mm long, with
2 fleshy, rhombic, concave wings with revolute margins;
anther cucullate, glabrous; pollinia 2, pyriform, with
elongate, triangular stipe and a brown, peltate viscidium.
Trichocentrum caloceras grows either on twigs
or on larger, shadier branches in premontane
cloud forests at 900-1300 m. Flowering generally
May to July. Southeastern Costa Rica and Pana-
ma.
Trichocentrum caloceras may be easily distin-
guished from its other Costa Rican relatives by
the glabrous anther.
Trichocentrum capistratum Linden & Rchb. f.,
Card. Chron. 1257. 1871. TYPE: Costa Rica,
Wallis (w). Trichocentrum panamense Rolfe,
Bull. Misc. Inform. 341-342. 1913. TYPE:
Panama, Canal Zone, Lipscomb s.n. (K). Tri-
chocentrum pusillum Lehmann, name only (K,
MS). Figure 48D.
Plant epiphytic, pendent, cespitose. Rhizome short;
roots slender, to about 1 mm in diameter. Pseudobulbs
very short, 1 -foliate. Leaf fleshy, oblong-lanceolate to
elliptic-lanceolate, 4-7 cm long, 2 cm wide, gradually
narrowed to a conduplicate, sessile base, apex acute. In-
florescence a pendent raceme of 3-7 successive flowers,
2-4 cm long including the zigzag rachis; ovary with
pedicel 5-7 mm long, subtended by concave, triangular,
acute bracts to 1.5 mm long. Flowers small with sepals
and petals pale green or greenish yellow and lip white
marked near the base with 2-4 reddish brown blotches.
Sepals subequal, free, spreading, subcarinate, oblong-
elliptic to elliptic, 1.6-1.8 cm long, 3-4 mm wide, acute.
Petals oblong-elliptic, 1.4-1.6 cm long, 4-5 mm wide,
acute. Lip adnate to the column, elliptic-ovate, concave,
carinate, 1.6 cm long including spur, ca. 9 mm wide,
with a short, obscurely 4-lobate, dorso-ventrally flat-
tened spur at the base, acute, the basal margins some-
what undulate. Column short, stout, ca. 5 mm long, with
a pair of porrect, fleshy, subquadrate, acute wings; an-
ther cucullate, hirsute; pollinia 2, pyriform, compressed-
concave; stipe flat, elongate, triangular; viscidium pel-
tate, brown.
Trichocentrum capistratum is found in lowland
and premontane evergreen forests at 500-1000 m,
often growing low on twigs with roots in dense
moss. It sometimes is found in bright light on the
very smooth and dry bark of Psidium guayava.
Plants have been observed in flower April to June,
August, October, and November; to be expected
throughout the year. Costa Rica to Colombia and
Venezuela.
This species, while variable in leaf size and
shape as well as scape length, is easily distin-
guished from similar T. caloceras and T. costari-
cense by the lack of spots on the lip and the pres-
ence of conspicuous trichomes on the anther.
Trichocentrum costaricense Mora-Retana &
Pupulin, Se:byana 15(2):94. 1994. TYPE: Costa
Rica, Alajuela, near Ciudad Quesada, about 700
m, Horich s.n. (holotype: us;). Figure 48B.
Plant epiphytic, cespitose, pendent. Rhizome short;
roots filiform, glabrous. Pseudobulbs short, rounded,
cespitose, 1 -foliate, ca. 3 mm long. Leaf fleshy, from a
cuneate base, obovate-elliptic to elliptic-lanceolate,
acute, sessile, 4 cm long, 1.7 cm wide. Inflorescence a
raceme with green, terete peduncle, 2-3 cm long; rachis
abbreviated, producing 2-3 flowers consecutively; ovary
with pedicel linear-clavate, 8-10 mm long; floral bracts
distichous, short, ovate, cucullate, 2 mm long, acute.
Flowers spreading; tepals greenish white, lip white with
lilac spots. Sepals similar, subcarinate, to 14 mm long;
dorsal erect, obovate-oblong, acute to acuminate, 4 mm
wide; lateral spreading, obliquely linear-lanceolate,
acute, 2-2.5 mm wide. Petals obovate-oblong, 13 mm
long, 4 mm wide, obtuse to acute. Lip elliptic, adnate
to the column, concave, obtuse to retuse, carinate api-
cally, 1.6 cm long, 10 mm wide, producing with the
column base a short, flattened, truncate, obscurely 2-4-
lobed spur; callus consisting of a pair of obscure keels
toward the base. Column short, stout, without a foot,
ca. 5 mm long, with a pair of fleshy, incurved, subfal-
cate, obtuse wings; anther white, cucullate, papillose;
pollinia 2, pyriform, concave, with elongate, triangular
stipe; viscidium peltate, brown.
Epiphytic in the premontane rain and wet for-
ests of Cordillera de Tilaran, both along the Cen-
tral Pacific watershed and the Rio San Carlos
drainage. Trichocentrum costaricense is generally
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
167
found on shady, constantly moist trunks and inner
branches. Flowering October to January. Endemic
to northern Costa Rica.
Trichocentrum costaricense differs from T.
capistratum and T. caloceras in the fine purple
spotting of the lip and in the incurved, obtuse,
subfalcate wings of the column. The only other
Costa Rican species with purple lip color is T.
caloceras, which has an obovate-subpandurate
lip, revolute margins of the column wings, and
glabrous anther cap. Both T. capistratum and T.
caloceras are more southerly species.
Trichocentrum cymbiglossum Pupulin, Lindley-
ana 9:51. 1994. TYPE: Costa Rica, Alajuela,
Laguna Bosque Alegre, 750 m, Pupulin 5 (ho-
lotype: usj). Figure 49A.
Plant epiphytic, cespitose. Rhizome short; roots fili-
form, glabrous. Pseudobulbs minute, rounded, 1 -foliate,
about 4 mm long. Leaf dark green, fleshy, sessile, linear-
elliptic to elliptic, 5.3-7.5 cm long, 1.7-2.5 cm wide,
obtuse to acute. Inflorescence a pendent raceme to 7 cm
long; rachis short, zigzag, of 1-7 successively borne
flowers; ovary with pedicel 4-7 mm long; floral bracts
papery, distichous, ovate, cucullate, 6-7 mm long, acute.
Flowers rather large, to 4.2 cm in diameter; tepals
greenish white, free, spreading; lip white. Dorsal sepal
elliptic-lanceolate, subcarinate, 1.5-1.9 cm long, 3-5
mm wide; apex acute, somewhat recurved; lateral sepals
obliquely subfalcate, somewhat shorter than dorsal se-
pal, 1.35-1.8 cm long, 3 mm wide, acute. Petals oblan-
ceolate-oblong, 1 .87 cm long, 4.2-5 mm wide, apex
acute and subcarinate. Lip adnate basally to the column,
concave, carinate, broadly rhombic, 2-2.6 cm long in-
cluding the short, flat, truncate, 4-lobed spur, 1.8 cm
wide at the middle, apex retuse, margins undulate. Col-
umn stout, ca. 5 mm long, with a pair of erect-spreading,
triangular, acute wings, apical margins erose; anther cu-
cullate, white, papillose; pollinia 2, pyriform, concave,
supported on a short triangular stipe; viscidium peltate,
brown.
Usually epiphytic in medium or deep shade in
very moist sites on small branches or trunks cov-
ered by dense moss mats in evergreen forests at
600-750 m. Flowering September to November.
Endemic to Costa Rica; Atlantic drainage of
northern and central Cordilleras and the high ba-
sin of the Rio Reventaz6n.
The rhombic, concave lip, which is much lon-
ger than the sepals, is unique among species of
Costa Rican Trichocentrum. Before its descrip-
tion, T. cymbiglossum had been accepted as T.
candidum Lindl. (e.g., Icon. PL Trop. 15: t. 1495)
but has a rhombic lip among other differences.
Trichocentrum dianthum Pupulin & Mora-Re-
tana, Selbyana 15(2):90. 1994. TYPE: Costa
Rica, Province of San Jose, Las Nubes de Qui-
zarra, 1988, flowered in cultivation March
1989, /. Cambronero s.n. (holotype: usj). Fig-
ure 47B.
Plant epiphytic, cespitose, pendent. Rhizome short; roots
filiform, glabrous. Pseudobulbs minute, rounded, to 5 mm
long, 1 -foliate. Leaf light green, sessile from a conduplicate
base, oblong-elliptic to oblong-obovate, to 9.7 cm long, 3
cm wide, apex acute to minutely retuse. Inflorescence
erect, simple or more commonly branched, 2-flowered in
each branch; peduncle terete, green, 3.5-4 cm long; ovary
with pedicel 2.3 cm long, linear-clavate; floral bracts con-
spicuous, ovate, concave, spreading, brownish, to 5 mm
long. Flowers spreading, with tepals yellow covered by a
very large brown blotch; lip white, marked near the base
by 2 rose-purple blotches; column wings brown-striped, an-
ther white. Sepals dissimilar, carinate; dorsal erect, elliptic-
oblanceolate, carinate, to 1.65 cm long, 6.3 mm wide, ob-
tuse to subacute; lateral sepals spreading, obliquely oblan-
ceolate, to 1.7 cm long, 4 mm wide. Petals linear-oblong,
subcarinate, to 1.65 cm long, 5 mm wide, acute. Lip spat-
ulate, adnate to column base, 2.5 cm long, 9.2 mm wide;
lateral lobes at base narrow, falcate; midlobe rounded in
front and with crisped margins; spur at the base elongate,
slender, conic, 1 . 1 cm long. Column short, stout, to 5 mm
long, with a pair of fleshy, erect, subquadrate wings; anther
cucullate, hirsute; pollinia 2, pyriform, compressed, with
short, triangular stipe; viscidium peltate, brown.
Rare epiphyte restricted to 1000-2000 m in the
northern part of the Rio General Valley, both in
the watershed of the Fila Costera and the slopes
of the Cordillera de Talamanca, and to the high
western intermountain valleys of the Cerro Vuel-
tas. Sometimes found on Citrus growing in thick
moss of shady branches. Flowering February to
April. Endemic to Costa Rica.
Trichocentrum dianthum may be easily distin-
guished by its long, conic spur and the heavy
brown striping on the column wings.
Trichocentrum estrellense Pupulin & J. B. Gar-
cia, Lindleyana 10:195-197. 1995. TYPE: Cos-
ta Rica, Cartago, El Guarco, along a minor trib-
utary of Rio Reventazon, F. Pupulin 209 (ho-
lotype: USJ). Figure 47D.
Plant epiphytic, cespitose, pendent. Rhizome short;
roots filiform, glabrous, silvery white. Pseudobulb cy-
lindric, to 10 mm long, 5 mm wide, 1-foliate. Leaf light
green, sessile from a conduplicate base, linear-elliptic to
elliptic-oblong, 8-12(20) cm long, 2.5-3 cm wide, ob-
tuse to retuse. Inflorescence pendent, 3-many-flowered,
successive, sometimes branched at the base; peduncle
terete, 4-1 1.5 cm long, concealed by 2-3 concave, ovate
bracts; ovary linear-clavate, ca. 3 cm long including the
pedicel. Flowers large for the genus, tepals free, ivory
168
FIELDIANA: BOTANY
white with narrow chestnut brown blotch on the lower
Vy, lip white marked near the base by 2 rose-purple bars;
column wing margins spotted and blotched with brown;
anther white. Sepals dissimilar, obtuse to retuse; dorsal
obovate-elliptic, to 1.6 cm long, 8.5 mm wide, slightly
concave apically; lateral sepals obliquely elliptic, to 1.6
cm long, 6.5 mm wide, apex subcarinate and slightly
concave, with short, rounded apicule. Petals linear-ellip-
tic, 1.6 cm long, 6.5 mm wide, slightly concave apically.
Lip adnate to the column base, obovate-subpandurate,
2.1 cm long, with short linear claw 2 mm long abruptly
expanding to the blade, with 2 broad, subquadrate, pu-
bescent basal lateral lobes; apex deeply emarginate; mar-
gins smooth, entire, producing with the column base a
very short, blunt, saccate spur 2.6 mm long, 3.5 mm
wide. Column short, stout, 3 mm long, with a pair of
short, erect, subfalcate, lacerate wings; anther cucullate,
papillose; pollinia 2, pyriform, rugulose, with short tri-
angular stipe; viscidium elliptic, brown.
Epiphytic in premontane or tropical lowland
rain forests at 450-1450 m. Known from the At-
lantic slopes of Cordillera de Talamanca, where it
grows on shady branches and moss-covered
trunks mainly over streams. Flowering June to
September. Endemic to Costa Rica.
The obovate, fleshy lip with plain apex, the short
and rounded lateral lobes, and successive inflores-
cence distinguish T. estrellense from close relatives.
Trichocentrum pfavii Rchb. f., Gard. Chron.
(n.s.) 16:70. 1881. TYPE: Chiriqui (probably
Panama), Pfau 60 (w). Trichocentrum pfavii
var. zonale Rchb. f., Gard. Chron. (n.s.) 19:44.
1883. Trichocentrum saundersianum Endres &
Rchb. f., name only (w, MS). Trichocentrum
saundersii Endres & Rchb. f., name only (w,
MS). Trichocentrum zonale Rchb. f., name only
(w, MS). Figure 47C.
Plant epiphytic, cespitose, pendent. Rhizome short;
roots slender for the genus, glabrous. Pseudobulbs mi-
nute, rounded, s 5 mm long, covered by a brown sheath,
1 -foliate. Leaf light green, fleshy, sessile from a con-
duplicate base, oblong-elliptic to oblong-ovate, 7.2-12.5
cm long, 1.4-3.8 cm wide, obtuse to retuse. Inflores-
cences 2-4, pendent to suberect, shorter than the leaves,
frequently branched at the base, mostly 2-flowered; pe-
duncle terete, ^ 5.1 cm long, concealed by 2-3 spread-
ing, ovate, concave bracts; ovary linear-clavate, ca. 2 cm
long including the pedicel. Flowers large and showy,
tepals free, spreading, white with large median brown-
rose blotch; lip white marked basally by 1-2 rose-purple
blotches; anther white. Sepals dissimilar, somewhat con-
cave; dorsal elliptic-ovate, to 1.5 cm long, 8 mm wide;
apex rounded to emarginate, sometimes with a dorsal
apicule; lateral sepals obliquely elliptic-lanceolate, 1.5
mm long, 6 mm wide, subcarinate and thickened through
the middle, apex obtuse and minutely apiculate. Petals
spatulate, to 1.4 cm long, 6 mm wide, obtuse. Lip adnate
to the column, cuneate-flabellate, 2.2 cm long, 1.6 cm
wide, with cuneate claw 5 mm long, the base with 2
narrow, erect, falcate, pubescent lateral lobes, expanding
abruptly to the suborbicular blade, producing with the
column base a very short, blunt, saccate spur 3 mm long.
Column short, stout, ca. 5 mm long, with a pair of large,
erect-spreading, subquadrate, rounded wings spotted
brown along the margins; anther hemispheric, papillose;
pollinia 2, pyriform, on a short, triangular stipe; visci-
dium peltate, brown.
Epiphytic on small branches with persistent,
dense foliage and on moss-covered vines close to
water in premontane and lower montane rain for-
est at 800-1500 m. Flowering November to
March. Panama (Chiriqui) to Costa Rica in Valle
de Goto Brus and Valle del General and along the
Pacific slopes of the Cerro de la Muerte and Cerro
Vueltas just to the region of Dota northward.
This species could probably be divided into two
geographic races; one distributed from Panama to
the northern limits of the Valle del General and
the other inhabiting the medium intermountain
valleys in the region of Dota. This last race gen-
erally presents a markedly narrower lip and a
darker, brown color on the tepals.
Trichopilia Lindl.
(D. E. Mora de Retana)
Epiphytic and mostly cespitose herb. Rhizome short;
roots cylindric, white. Pseudobulbs suborbicular, ovoid to
linear-oblong, often much compressed, with nonfoliaceous
sheaths at the base, apex 1 -foliate. Leaves coriaceous, con-
duplicate, elliptic to lanceolate, acute. Inflorescence lateral,
a raceme or flower solitary. Flowers large and often very
showy, usually white to yellow variously marked with pur-
ple or red. Sepals similar, narrow, imbricate in bud, some-
times twisted, the lateral sepals sometimes variously con-
nate to about the middle, margins usually undulate. Petals
similar to the sepals. Lip simple or 3(4)-lobate, shortly ad-
nate to the column, lateral lobes usually inrolled around the
column; midlobe usually emarginate; margins often cris-
pate, dentate, or undulate; callus usually with 1-3 keels.
Column erect, semiterete, the base without a foot, anther
bed dentate to fimbriate; pollinia 2, waxy with stipe with
viscidium. Fruit a capsule.
A Neotropical genus of about 30 species of low
to intermediate elevations. Six species are known
in Costa Rica. Sterile plants or plants in fruit are
usually easily distinguished by the nonfoliaceous,
often spotted sheaths at the base of the apically
1 -foliate pseudobulbs.
According to Chase, Trichopilia is closely re-
lated to Helcia and Neoescobaria, a clade that is
somewhat isolated in the Oncidiinae.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
169
Key to the Species of Trichopilia
la. Pseudobulbs suborbicular to subquadrate, to at most 2 times as long as wide 2
2a. Inflorescence with a single yellowish white flower T. maculata
2b. Inflorescence with 2 or more white flowers with purple spotted lip T. suavis
Ib. Pseudobulbs linear, usually more than 3 times as long as wide 3
3a. Flowers yellowish white, without reddish or brown spots
4a. Flowers 2-5 per inflorescence T. turialbae
4b. Flowers usually 1 per inflorescence T. maculata
3b. Flowers with brown or reddish spots; sepals free or united only at the base 4
5a. Sepals and petals tinged with brown; lip white with brown spots T. tortilis
5b. Sepals and petals white or yellow tinged with red; lip white, lined, tinged, or spotted with
red 5
6a. Lip white tinged variously with red and with white margin T. marginata
6b. Lip yellow, with reddish brown, more or less irregular longitudinal spots
T. galeottiana
Trichopilia galeottiana A. Rich. & Gal., Ann.
Sci. Nat. Bot. ser. 3, 3:26. 1845. TYPE: Mexi-
co. (K, photo seen). ITrichopilia picta Lemaire,
111. Hort. 6:86-87, t. 225. 1859. TYPE: Mexico.
Figure 50C.
Plant a compact, cespitose epiphyte, to ca. 30 cm tall.
Rhizome short; roots 1-2 mm in diameter. Pseudobulbs
subcylindric, ancipitous, 8-12 cm long, 1.2-2 cm wide,
subtended by grayish bracts becoming papery with age,
1 -foliate apically. Leaves elliptic-lanceolate, carinate
abaxially, 16-25 cm long, 3-5 cm wide, apex acute. In-
florescences lateral, pendent, 3-4, each with a solitary
flower; ovary with pedicel 2.7 cm long, subtended by 3
papery, obtuse bracts. Flowers with greenish yellow se-
pals and petals with an interrupted reddish brown stain
near the midvein; lip yellowish white with reddish
brown spots apically and at the base of the lateral lobes,
the lamina with a saffron-yellow spot centrally. Sepals
subequal, elliptic-lanceolate, acute, carinate abaxially,
4.5-5 cm long, 1-1.2 cm wide, acute; dorsal erect, free;
laterals shortly connate and recurved. Petals similar to
the sepals but broader, 4.5-5 cm long, 1-1.5 cm wide.
Lip 3-lobate, trumpet-shaped, 5.5-7 cm long, 2-4 cm
wide in natural position; lateral lobes rounded, overlap-
ping and embracing the column, midlobe broadly emar-
ginate, recurved apically; callus sulcate along the mid-
vein, with 2 hollowed areas laterally. Column subterete,
2-2.2 cm long, with fimbriate anther bed; pollinia 2,
with viscidium and stipe.
Epiphytic, growing on larger branches in pre-
montane rain forests at 1000-1200 m. Flowering
May to July. Mexico to Costa Rica; perhaps also
Panama.
This species is similar to T. marginata but has
yellow flowers with ovary subtended by three
broad bracts, and the sepals and petals are nearly
flat.
Trichopilia maculata Rchb. f., Bonplandia. 3:
215. 1855. TYPE: Port Chagres (Panama?), Ke-
fer stein ex Behr (type not found). Figure 49B.
Plant a cespitose epiphyte 15-20 cm tall. Rhizome
short; roots ^ 2 mm in diameter. Pseudobulbs elliptic-
oblong, ancipitous, 5-7 cm long, 1-1.4 cm wide, the
base enveloped in 3-4 spotted, imbricate bracts, apex 1-
foliate. Leaves subcoriaceous, elliptic-lanceolate, 16-20
cm long, 3.5-4 cm wide, acute. Inflorescence usually
solitary, slender, arcuate or somewhat pendent; scapes
4-6 cm long. Flowers of moderate size, white, lip pale
yellow with orange-red centrally. Sepals subequal, free
or connate near the base, broadly spreading, lanceolate,
often somewhat twisted, 2.5-3 cm long, 3-5 mm wide.
Petals similar to the sepals, lanceolate, 2.5-3 cm long,
3-5 mm wide, acuminate, with undulate margins. Lip
shortly adnate to column base, obscurely 3-lobate, ob-
ovate when spread, 3.5-3.7 cm long, 1.6-2.2 cm wide,
narrowed at the base; lateral margins rounded, convo-
lute, forming a tube; midlobe emarginate, with spreading
or reflexed lobules; callus with a short, inconspicuous,
central keel. Column slender, semiterete; apical margins
distinctly 3-parted and minutely denticulate; pollinia 2,
with stipe and viscidium. Immature capsule 3 cm long.
Epiphytic in premontane rain forests at 100-
400 m. Flowering May to July. Costa Rica and
Panama; also doubtfully reported from Guatemala
and El Salvador.
Trichopilia maculata is similar to T. tortilis but
has yellow-green flowers with a yellow lip, and
the sepals and petals are nearly flat. The pseudo-
bulbs are invested with several heavily spotted
bracts.
Trichopilia marginata Henfr., Gard. Mag. Bot.
3:185, t. 1851. TYPE: New Granada. Tricho-
pilia coccinea Warsc., Paxton's Fl. Gard. 2:79.
1851. TYPE: Costa Rica, Warscewicz (w, photo
170
FIELDIANA: BOTANY
seen). Trichopilia crispa Lindl., Gard. Chron.
342. 1857. TYPE: Probably Costa Rica, based
on Warscewicz collection, Rticker (K, photo
seen). Figure SOB.
Plant epiphytic, cespitose. Rhizome abbreviated;
roots 1-2 mm in diameter. Pseudobulbs subcylindric,
slightly tapering, laterally flattened, 1-foliate, 4-12 cm
long, 1—3 cm wide, covered at the base when young by
several bracts turning papery with age. Leaves coria-
ceous, elliptic-lanceolate, 10-25 cm long, 2.5-5.5 cm
wide, conduplicate at the base, the apex acute and re-
curved. Inflorescences 1-3 borne laterally at the base
of the pseudobulb, each consisting of a single flower
borne on a short scape, pendulous; ovary and pedicel
strongly curved, 2-3 cm long. Flowers 8-14 cm in di-
ameter, variable in color; sepals and petals yellowish
green with a band of reddish brown centrally; lip white
spotted with wine red to entirely red with white margins.
Sepals similar, twisted, linear lanceolate, 7.5-8 cm long,
1-1.3 cm wide, acute and lightly conduplicate at the
apex; lateral sepals very shortly connate at the base. Pet-
als similar to the sepals but somewhat broader, 6.5-7 cm
long, 1.5—1.8 cm wide. Lip tubular, 3-lobate, 6-7 cm
long, 3.5-4 cm wide when spread; lateral lobes clasping
the column, separated from the midlobe by a shallow
sinus; midlobe rounded and emarginate; callus poorly
defined, an intense yellow furrow at the base. Column
semiterete, clavate, greenish at the base, white at the
apex, 2.5-3 cm long, provided apically with a prominent
anther bed with lacerate margins; pollinia 2, waxy, with
stipe and viscidium; stigma entire, cordate.
Epiphytic on the shady trunks of trees in pre-
montane and lower montane rain forests at 1200-
1500 m. Flowering December to July. Reported
from Nicaragua to Colombia.
Trichopilia marginata is distinguished from
similar T. galeottiana A. Rich & Gal. by the sol-
itary white and wine red flowers with crisped se-
pals and petals.
Trichopilia suavis Lindl., Paxton's Fl. Gard. 1:
44. 1850-1851. TYPE: Central America, Lod-
diges (K, photo seen). Trichopilia kienastiana
Rchb. f., Gard. Chron. n.s. 20:166. 1881.
TYPE: Kienast (w, photo seen). Figure 49C.
Plant epiphytic, shortly creeping to cespitose. Rhi-
zome short; roots 1-2 mm in diameter. Pseudobulbs
fleshy, ovoid to suborbicular, laterally flattened, 4-7.2
cm long, 3.5-6.2 cm wide, apically 1-foliate, protected
at the base by several spotted bracts becoming papery
with age. Leaves subcoriaceous, broadly elliptic, form-
ing a conduplicate petiole at the base, the apex acute,
4.5-30 cm long, 3.8-8.2 cm wide. Inflorescence a lat-
eral, short, arcuate or pendulous, 2-5-flowered raceme
supported on a short peduncle; ovary and pedicel 3.5 cm
long. Flowers with a fragrance similar to sweet peas,
white with rose spots. Sepals similar, the laterals shortly
connate, elliptic-lanceolate, the margins undulate, 3.3-
5.2 cm long, 8-10 mm wide, with a prominent keel
abaxially on midvein. Petals similar to the sepals but
broader, 3-5.2 cm long, 1-1.2 cm wide. Lip obscurely
3-lobate, tubular or horn-shaped in natural position,
quadrate when expanded, adnate to the base of the col-
umn, 4.8-7 cm long, 3.5-4.8 cm wide, lateral lobes in-
volute and clasping the column, apex retuse; margins
undulate or crisped, disk of the lip with a prominent
keel. Column elongate, terete, with a conspicuous lac-
erate anther bed at the apex; pollinia 2, waxy, with
prominent stipe and viscidium.
Epiphytic on large branches and trunks in pre-
montane rain forests at 550-1300 m along both
Pacific and Atlantic slopes. Flowering February to
April; rarely to May. Reported from Costa Rica
to Colombia.
Trichopilia suavis has large, white, fragrant
flowers with fragrance of sweet peas, and the lip
is spotted with rose or purple. Occasionally white
forms are seen. Trichophilia suavis is suspected
of hybridizing with T. marginata Henfr. at its low-
est elevations, where the two species are sympat-
ric.
Trichopilia tortilis Lindl., Nat. Syst. Bot. (ed. 2)
446. 1836. Edwards's Bot. Reg. 22: t. 1863.
1836. TYPE: Mexico, Barker (K, photo seen).
Figure 50A.
Plant epiphytic, cespitose. Rhizome abbreviate; roots
2-3 mm in diameter. Pseudobulbs oblong to cylindric,
compressed, 5.5-7 cm long, 1.5-2 cm wide, the base
concealed by one or more bracts spotted with brown,
apically 1-foliate. Leaves subcoriaceous, elliptic-lanceo-
late, abaxially carinate, forming a conduplicate petiole
at the base, apically acute, 8-15 cm long, 3.5-4 cm
wide. Inflorescence a short scape with a single flower
borne laterally at the base of the pseudobulb; ovary and
pedicel curved, 3 cm long. Flowers 10-13 cm across;
sepals and petals greenish white with a brown longitu-
dinal band centrally for its entire length; lip white with
brown spots. Sepals and petals similar, linear-lanceolate,
acute to acuminate, strongly twisted and with undulate
margins, 5.5-7.5 cm long, 8-12 mm wide. Lip adnate
to the base of the column, obscurely 3-lobate, the side
lobes involute and clasping the column forming a tube
at the base, 5.5-6.2 cm long, 4-5.2 cm wide when
spread, elliptic-suborbicular, emarginate; disk with a pair
of excavated areas near the column apex. Column
white, semiterete with fimbriate, tripartite anther bed;
pollinia 2, with prominent stipe and viscidium.
Reportedly uncommon epiphyte of premontane
rain forests of the Atlantic slope at 1000-1200 m,
where it has been collected in Zapotal de Perez
Zeledon. The Costa Rican forms are somewhat
larger than those from farther north. Flowering
April to June. Reported from Costa Rica but seen
only in live collections.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
171
Trichopilia tortilis differs from T. maculata by
the strongly twisted sepals and petals, larger pseu-
dobulbs, and the brown-spotted lip.
Trichopilia turialbae Rchb. f., Hamburger Gar-
ten-Blumenzeitung 19:11-12. 1863. TYPE:
Costa Rica, Wendland (w, photo seen). Figure
49D.
Plant a cespitose epiphyte. Rhizome short; roots
about 2 mm in diameter. Pseudobulbs ovoid-lanceolate,
compressed, 6-9 cm long, 2.5-3.5 cm wide, 1 -foliate.
Leaves coriaceous, elliptic, acute, with prominent keel
abaxially, 1 8-25 cm long, 4-6 cm wide, short-petiolate,
acute. Inflorescence a short, lateral, 2-5-flowered, ra-
ceme 8-1 1 cm long; ovary with pedicel 4-4.5 cm long,
subtended by a triangular floral bract about 3 cm long.
Flowers not spreading widely, membranaceous, short-
lived, 4.4-5 cm long, 1.5-1.8 cm wide in natural posi-
tion. Sepals similar, elliptic-lanceolate, acuminate, 4 cm
long, 4-5 mm wide; dorsal free, recurved; laterals ca. %
connate, the apices somewhat recurved. Petals similar
to the dorsal sepal but somewhat broader and with un-
dulate margins, 4-4.5 cm long, 7-9 mm wide. Lip ad-
nate to the base of the column, 3-lobate, trumpet shaped
with upcurved, tubular base, 4-5 cm long, 3-4 cm wide
when spread; midlobe emarginate and reflexed; lateral
lobes inrolled around the column; callus of 3 elevated
keels centrally. Column subterete, 1.5 cm long, with 3
fimbriate projections apically; pollinia 2, waxy, pyri-
form, compressed, with prominent viscidium and stipe.
Epiphytic in premontane rain forests at 600-
1200 m. Flowering October to January. Nicaragua
to Panama; also reported from Colombia.
Trichopilia turialbae is easily distinguished in
Costa Rica by the rather small creamy white flow-
ers, borne 2-5 per inflorescence. Also, the tepals
are nonspreading. It has been confused by authors
with T. galeottiana A. Rich. & Gal., but that spe-
cies has larger flowers with essentially flat,
spreading, much broader sepals and petals.
Trizeuxis Lindl.
(D. E. Mora de Retana)
Plant a psygmoid epiphyte. Stems short, pseudobulbs
small, suborbicular, compressed, concealed by several
leaves arranged in a fan, apically 1 -foliate. Leaves ar-
ticulate, fleshy, conduplicate, equitant, falcate. Inflores-
cence lateral, paniculate, each branch apex densely flow-
ered. Flowers very small, subglobose. Sepals dissimilar,
concave, shortly united at the base, lateral sepals connate
to near the bifid apex. Petals elliptic-ovate, concave at
the base, obtuse to bluntly acute. Lip simple or obscure-
ly 3-lobate, apex fleshy and recurved, margins erect and
parallel with the column. Column stout, widened at the
apex, with neither foot nor wings; anther terminal; pol-
linia 2, waxy, with simple elongate stipe and viscidium.
Fruit a capsule.
A Neotropical genus of a single widespread
species ranging from Costa Rica to Bolivia, Bra-
zil, and the West Indies.
On the basis of the flowers, Trizeuxis seems
related to lonopsis, a relationship supported by the
molecular data of Chase and Palmer.
Trizeuxis falcata Lindl., Coll. Bot. t. 2. 1823
(neither description nor type seen). Figure SOD.
Plant epiphytic often on twigs, forming small clumps,
to ca. 8 cm tall. Rhizome short; roots large for the plant,
to 1 mm in diameter. Pseudobulbs small, suborbicular,
compressed, 1-1.2 cm long and high, the base covered
by 2-4 leaves, apically 1 -foliate. Leaves articulate,
fleshy, conduplicate, falcate, 5-12 cm long, 5-8 mm
wide (deep), acute. Inflorescence lateral, paniculate, ex-
ceeding the leaves; ovary with pedicel 1.2 mm long.
Flowers minute, subglobose, yellow-green with orange
lip. Sepals dissimilar, shortly united at the base; dorsal
concave, elliptic -ovate, 3-3.2 mm long, 2-2.3 mm wide,
obtuse; lateral sepals connate to near the bifid apex, 3-
3.2 mm long, 1-1.1 mm wide. Petals similar to the dor-
sal sepal, concave at the base, elliptic-ovate, obtuse or
subacute. Lip simple or obscurely 3-lobate, 4-4.2 mm
long, 2-2.2 mm wide, lateral margins erect and parallel
with the column, apex fleshy and recurved; callus sul-
cate, consisting of a pair of fleshy lateral projections to
beyond the middle. Column stout, broadened apically,
to 2 mm long; anther terminal; pollinia with well-de-
veloped stipe and viscidium.
Uncommon epiphyte of twigs and smaller
branches, usually of disturbed evergreen forests at
50-900 m. Often seen growing on citrus, hibiscus,
and coffee. Flowering February to April. Wide-
spread from Costa Rica to Bolivia, Brazil, and the
West Indies.
Trizeuxis falcata is easily recognized by the
densely clustered, tiny, bell-shaped flowers and
the plant with equitant leaves.
Warmingia Rchb. f.
(J. T. Atwood, based on reference below)
REFERENCE — B. Johansen, Warmingia margar-
itacea sp. nov. (Orchidaceae) from Costa Rica and
a review of the genus. Lindleyana 7:194-198.
1992.
Plant a cespitose epiphyte. Pseudobulbs present, sub-
tended by nonfoliaceous bracts; apex 1 -foliate. Inflores-
172
FIELDIANA: BOTANY
cence a lateral few-flowered raceme. Flowers white or
white with yellow. Sepals subequal. Petals usually
rhombic and dentate, broader than the sepals. Lip 3-
lobate, the lateral lobes usually rounded and dentate, the
midlobe elongate. Column elongate, anther lacking a
conspicuous anther bed; pollinia 2, with short or elon-
gate stipe and viscidium. Fruit a capsule.
A Neotropical genus of about four species with
scattered distribution in Bolivia, Brazil, Costa
Rica, and Ecuador.
Warmingia is closely related to Macradenia, a
genus with green to red flowers and usually a
flared, variously dentate anther bed. The DNA
work of Chase and Palmer show a close relation-
ships of these genera with Notylia and Macrocli-
Warmingia margaritacea B. Johans., Lindley-
ana 7:194-196. 1992. TYPE: Costa Rica, Tur-
rialba, C.A.T.I.E., 600 m, Johansen & S0rensen
138 (holotype: c, not seen).
Plant an epiphytic herb to 7 cm high. Roots white,
smooth, 1 mm in diameter. Pseudobulbs small, of a sin-
gle internode, conic to ovoid, 7-8 mm tall, 3-4 mm
wide, covered with 3-4 withering brown sheaths.
Leaves dark green, coriaceous, lanceolate, somewhat
falcate, 3.5-6 cm long including the 0.6- 1.0-cm- long
petiole, 1-1.7 cm wide. Inflorescence a lateral, pendu-
lous raceme, to 3 cm long, with 2-3(4) flowers; bracts
triangular, 6 mm long, 3 mm wide at the base; ovary
with pedicel 8-9 mm long. Flowers ivory white and
semiopaque; apparently autogamous. Sepals similar;
dorsal slightly concave, elliptic-lanceolate, 8 mm long,
3 mm wide with a carinate, acute apex; lateral sepals
slightly falcate, 7 mm long, 3 mm wide. Petals rhombic,
8 mm long, 5 mm wide, apex slightly carinate, margins
denticulate-erose to somewhat crispate. Lip 3-lobate just
above the base, 8 mm long, 9 mm wide across the lateral
lobes when spread, with erose-serrate margins; lateral
lobes erect, transversely elliptic to reniform, 3 mm long,
5 mm wide; midlobe broadly ovate, 8 mm long, 3 mm
wide; callus 2-lobate, deeply sulcate, appearing to orig-
inate from the base of the sidelobes. Column terete to
semiterete, 3 mm long, 1 mm wide, slightly expanded
and with 2 appendages at the apex; anther ovate with a
decurved truncate apex; pollinia with short stipe and vis-
cidium. Capsule ellipsoid to obovoid, 12 mm long, 8
mm wide.
The above description is quoted and para-
phrased directly from the original description be-
cause the species is known only from the holo-
type. Warmingia margaritacea was originally
found growing on a cultivated hibiscus hedge at
Turrialba (600 m) and must have originated from
elsewhere, where it is hopefully more common.
Endemic to Costa Rica, where it is reported to be
autogamous.
Warmingia margaritacea should be easily rec-
ognized by the small plants and white flowers
with dentate petals and lip. They may have been
overlooked because of their small size or, alter-
natively, may be extremely rare.
Addendum to Subtribe M axillariinae
One new species of Maxillaria, published by I.
Bock in Die Orchidee, appeared as this work was
in press. In addition, one other new species, au-
thored by J. T. Atwood, is currently in press in
Selbyana.
The first species, Maxillaria admonens I. Bock
(Orchidee 48:105-107. 1997), should be com-
pared to M. gomeziana J. T. Atwood (p. 59). Bock
claims that M. admonens differs in the broader
egg-shaped pseudobulbs, the more compact plant,
and by the lip forming a sharper angle with the
column foot. At this time we cannot confirm or
reject this as a new species but note that the di-
agnostic drawings resemble M. gomeziana, re-
membering that plant habits are generally variable
in Maxillaria. Flower color patterns of M. ad-
monens evident in the separations are very similar
to those of flowers from which the holotype of M.
gomeziana was prepared. The illustration provid-
ed by Bock shows the same central keel in the
callus that is consistent with both M. bracteata
and M. gomeziana; therefore M. admonens cannot
be excluded from those taxa on this character
alone.
On page 72 a reference is made comparing
specimen Mora 60 (INB) to M. mombachoensis
Heller ex J. T. Atwood. The specimen differs from
that species in the shorter peduncles, shorter ova-
ries, and the shorter and remarkably thick leaves.
This species, apparently a narrow Costa Rican en-
demic, was published shortly after the present
work had gone to press. The description of it is
paraphrased below.
Maxillaria cacaoensis J. T. Atwood, Selbyana 19:
254-255. 1999. TYPE: Costa Rica. Province of
Guanacaste: Canton of Liberia, Cordillera de
Guanacaste, Cerro Cacao, Estacion Cacao,
1100 m, A. Mora 60 (holotype: INB: isotype:
SEL). Figure 51.
Plant cespitose, apparently densely so, to about 10
cm tall; roots to ca. 2 mm in diameter. Pseudobulbs
coarsely rugose in dried specimens, semiorbicular, com-
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
173
1 cm
FIG. 5 1 . Maxillaria cacaoensis.
174
FIELDIANA: BOTANY
pressed, 2-3.5 cm long, 2.5 cm wide, subtended by
sheaths that are apparently bladeless; apex 1 -foliate.
Leaves coriaceous, strongly keeled beneath especially
apically; blades 6-9 cm long, 2-3 cm wide in the type,
obtuse. Inflorescences short; scapes ca. 3 cm long, con-
cealed by inflated bracts; ovary about 1 .5 cm long, con-
cealed by the subtending floral bract. Flowers campan-
ulate to somewhat spreading, yellowish orange with
white at the base of the sepals and petals; lips reddish.
Sepals ovate-lanceolate, 3.5 cm long, 1.5 cm wide,
acute. Petals elliptic-oblanceolate, acute, 2.8 cm, 1 cm
wide. Lip hinged to the column foot, sharply 3-lobate
at about % from the base; lateral lobes rounded in front,
somewhat clasping the column; midlobe fleshy, ovate,
recurved to somewhat reflexed, ca. 1.8 cm long, 1.4 cm
wide in natural position, bluntly acute to obtuse; calli 2,
of 2 keels in front ending just below the base of the
midlobe. Column stout, nearly straight, ca 1 .4 cm long,
including anther; foot 5-6 mm long; anther ca. 4 mm.
Known only by the type collection from a cloud
forest on Cerro Cacao in northern Costa Rica at
1100 m. Flowering at least in January and Feb-
ruary, according to the collection date.
Maxillaria cacaoensis is most similar to Nica-
raguan M. mombachoensis J. T. Atwood in flower
color and in the large floral bracts, but has a much
shorter ovary and noticeably shorter and thicker
leaves. There may be yet another closely related
undescribed species on Volcan Maderas in Lake
Nicaragua with smaller flowers. Apparently, iso-
lation on these mountaintops enables the separate
populations to develop identifiably distinct traits.
Pollinators in this group are unknown, but the
large, inflated floral bracts and fleshy orange to
yellow flowers suggest pollination by humming-
birds.
Maxillaria cacaoensis is a member of the dif-
ficult M. cucullata Lindl. complex, represented in
Costa Rica by four species and in Panama by one.
The following key, modified from page 39, should
distinguish them in this area.
32a. Flowers with trigonous ovaries, sepals and
petals greenish stained with purple
M. obscura
32b. Flowers otherwise X
Xa. Flowers basically cream, lip spotted
with maroon M. cedralensis
Xb. Flowers otherwise Y
Ya. Floral bract > 1.5 X ovary; flow-
ers unspotted . . M. cacaoensis
Yb. Floral bract < 1.25 X ovary,
flowers greenish or orangish,
with small spots arranged in lines
M. punctostriata
Addendum to Subtribe Oncidiinae:
Leucohyle Klotzsch
(Joaquin B. Garcia-Castro and D. E. Mora de
Retana)
Plants generally epiphytic, cespitose and compact;
rhizome short. Pseudobulbs small, especially in the
Costa Rican species, cylindric or conic, truncate, with a
single fleshy, linear or subterete and sulcate leaf. Inflo-
rescence lateral at the pseudobulb base, pendent. Flow-
ers small, attractive, with whitish or translucent ground
color. Sepals and petals similar, slender and undulate,
the lateral sepals free. Lip united to the base of the col-
umn, prominently concave and with irregular border.
Column short, semiterete, apically with a hood with ir-
regularly dentate margin; pollinarium with two pyriform
pollinia, supported on a slender linear-lanceolate stipe
and small viscidium.
A genus of only two accepted species distrib-
uted from Mexico to Brazil and the Antilles; one
species in Costa Rica. Recent work of Norris Wil-
liams and Mark Whitten (unpublished) suggests a
close relationship with Trichopilia Lindl., Helcia
Lindl., and Neoescobaria Garay.
Leucohyle subulata (Sw.) Schltr., Die Orchi-
deen 469. 1914. Epidendrum subulatum Sw.,
Prodr. Ind. Occ. 123. 1788. TYPE: West In-
dies (not seen). Cymbidium subulatum (Sw.)
Sw., Nov. Act. Soc. Ups. 6:73. 1799. Tricho-
pilia subulata (Sw.) Rchb. f. Flora 48:278.
1865.
Plant epiphytic, cespitose; rhizome short; roots 1-
2 mm in diameter. Pseudobulbs fleshy, subconic to
cylindric, 0.8-3 cm long, 2-5 mm broad, concealed
at the base when young by several dark fibrous bracts;
apex 1 -foliate. Leaves fleshy, linear-lanceolate to
semiterete, sulcate above, conduplicate at the base,
apex acuminate, 8-25 cm long, 3-10 mm wide. Inflo-
rescence pendent, a raceme of 3-8 flowers supported
on a slender penduncle; ovary and pedicel slender,
about 1.4 cm long; floral bracts papery, obovate-ellip-
tic, acuminate, 9 mm long, 5 mm wide. Flowers
small, translucent white with the lip spotted with red,
especially near the base. Sepals subequal, linear-lan-
ceolate, the apex mucronate, with undulate margin,
1.6-2.6 cm long, 2-4 mm wide. Petals similar to the
sepals but somewhat shorter and broader, somewhat
twisted, 1.5-2 cm long, 3-6 mm wide. Lip basally
adnate to the column, simple, concave, rounded to
subcordate, with irregularly denticulate margins, 1.5—
2 cm long, 1.2-1.8 cm wide, apex apiculate; disc with
a basal depressed, subquadrate callus continued in
front by two lateral low keels and a somewhat more
prominent central keel. Column semiterete, some-
what recurved; apex with a hood covering the anther,
with irregular border.
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
175
This species is reported from Mexico and Nic-
aragua to Peru and Venezuela and the West Indies.
It is known in Costa Rica from San Carlos, Prov-
ince of Alajuela, and from Turrialba, Province of
Cartago. It appears similar to a small trichopilia
but has subconic-cylindric pseudobulbs in addi-
tion to more slender and more fleshy subterete
leaves. The flowers are somewhat unusual for the
unlobed concave lip and the column apex appear-
ing apically hooded.
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ATWOOD & MORA DE RET ANA: FLORA COSTARICENSIS 177
Index to Subtribes Maxillariinae and Oncidiinae
All taxa accepted for Costa Rica are in boldface Roman type; synonyms or compared species are in
italics. Page numbers for the main text of each taxon are underlined; page numbers of illustrations are
in boldface.
Ada 88, 89, 118, 132, 151
Ada chlorops 89, 90, 114, 134
Aeranthus hispWulus 153
Aeranthus (Cryptoplectri) lehman-
nii 31
Amparoa 89, 1 14
Amparoa costaricensis 90, 114
Anthosiphon 2
Aspasia 88, L14-115, 118, 151
Aspasia bibriana 1 15
Aspasia epidendroides 90, 115,
116
Aspasia epidendroides var. princip-
issa 115
Aspasia barclayi 115
Aspasia fragrans 115
Aspasia principissa 90, 115-116
Aspasia pusilla 119
Aspasia rousseauae 115
Brassia 88, 89, 116, 118, 132, 151
Brassia antherotes 116
Brassia antherotes var. longissima
116
Brassia arachnoidea 1 1 7
Brassia arcuigera 91, 1 16-117
Brassia caudata 91, 117
Brassia caudata var. hieroglyphica
117
Brassia chlorops 89
Brassia gireoudiana 91, 1 17
Brassia lawrenceana var. longissi-
ma 116
Brassia longissima 1 1 6
Brassia parviflora 89
Brassia verrucosa 91, 1 17, 1 18
Broughtonia alba 42
Camaridium 1
Camaridium adolphii 41
Camaridium affine 50
Camaridium album 42
Camaridium amparoanum 77
Camaridium arachnites 54
Camaridium bracteatum 47
Camaridium bradeorum 47, 48
Camaridium brenesii 79
Camaridium costaricense 79
Camaridium dendrobioides 54
Camaridium dichotomum 55
Camaridium exaltatum 57
Camaridium grandiflorum 43
Camaridium imbricatum 76
Camaridium jimenezii 54, 55
Camaridium latifolium 71
Camaridium minus 64
Camaridium nutantiflorum 80
Camaridium ochroleucum 50
Camaridium simile 54, 55
Camaridium squamatum 81
Camaridium tonduzii 41, 42, 79
Camaridium uncatum 80
Camaridium vinosum 80
Camaridium xylobiichilum 56
Camaridium wercklei 81
Camaridium wrightii 68
Camaridium xylobiichilum 56
Campylocentrum lehmannii 31
Campylocentrum hispidulum 153
Centropetalum costaricense 152
Chrysocycnis 32, 79
Chrysocycnis tigrinum 78
Cischweinfia 88, 118
Cischweinfia dasyandra 92, 1 18-
199
Cischweinfia pusilla 92, 119
Comparettia 87, 119, 159
Comparettia falcata 92, 119-120
Comparettia rosea 119
Cryptocentrum 1, 2
Cryptocentrum brenesii 31
Cryptocentrum calcaratum 3, 29,
31
Cryptocentrum flavum 2, 29-30,
32
Cryptocentrum gracillimum 2, 30
Cryptocentrum gracilipes 3 1
Cryptocentrum jamesonii 3 1
Cryptocentrum latifolium 3, 29,
30-31_
Cryptocentrum lehmannii 3, 29,
3J_
Cryptocentrum longiscapum 31
Cryptocentrum standleyi 2, 3, 31-
32
Cryptosanus scriptus 124
Cymbidium ochroleucum 50
Cymbidium pusillum 156
Cymbiglossum 156
Cymbiglossum bictoniense 157
Cymbiglossum hortensiae 157
Cymbiglossum stellatum 158
Cyrtochilum bictoniense 157
Cyrtoglottis 84
Dendrobium album 42
Diadenium 163
Dicrypta baueri 53
Dicrypta crassifolia 53
Dicrypta elatior 56
Dicrypta iridifolia 81
Dicrypta irisphyta 82
Epidendrum carthagenense 140
Epidendrum caudatum 117
Epidendrum cebolleta 140
Epidendrum labiatum 123
Epidendrum pusillum 1 56
Epidendrum satyrioides 122
Epidendrum sessile 53
Epidendrum utricularioides 122
Fernandezia 87, 120
Fernandezia acuta 125
Fernandezia robusta 126
Fernandezia tica 92, 120,
Gomesa planifolia 155
Goniochilus 89, 120-121
Goniochilus leochilinus 93, 121
169
Heterotaxis crassifolia 53
Hybochilus 89, 121
Hybochilus inconspicuus 121-122
lonopsis 87, 88, 119, 122, 127, 172
lonopsis costaricensis 122
lonopsis satyrioides 122
lonopsis utricularioides 93, 122-
123
Kefersteinia bicallosa 46
Lemboglossum 156
Lemboglossum bictoniense 157
Lemboglossum hortensiae 157
Lemboglossum stellatum 158
Leochilus 88, 89, 121, 122, 123
Leochilus labiatus 93, 123
Leochilus parviflorus 121
Leochilus scriptus 1 24
Leochilus tricuspidatus 93, 124
Leucohyle 88, 175
Leucohyle subulata 175-176
Lockhartia 87, 124 "
Lockhartia acuta 95, 125
Lockhartia amoena 94, 125-126.
127
Lockhartia amoena var. triangula-
bia 125
Lockhartia chiriquensis 126
Lockhartia costaricensis 125
Lockhartia dipleura 125
Lockhartia grandibractea 125
Lockhartia hercodonta 94, 125,
126
Lockhartia Integra 127
Lockhartia lamellosa 126
Lockhartia lankesteri 126
Lockhartia micrantha 94, 125,
126
Lockhartia oerstedii 94, 125, 126-
127
178
FIELDIANA: BOTANY
Lockhartia pallida 1 25
Lockhartia pittieri 95, 127
Lockhartia robusta 126
Lockhartia triangulabia 125
Lockhartia variabilis 127
Lockhartia verrucosa 126
Lophiaris 150
Lycaste aciantha 40
Macradenia 88, 127, 173
M iic rod in in m 87, 127, 173
Macroclinium allenorum 128
Macroclinium bicolor 131
Macroclinium confertum 95, 128-
129
Macroclinium cordesii 96, 128,
129
Macroclinium doderoi 128, 129,
Macroclinium generalense 95,
128, 129-130
Macroclinium glicensteinii 128,
130
Macroclinium lineare 128, 130-
131
Macroclinium paniculatum 128,
129, 131
Macroclinium ramonense 96, 128,
131, 132
Macroclinium robustum 128,
131-132
Malaxis caudata 117
Marsupiaria iridifolia 82
Marsupiaria valenzuelana 81
Maxillaria 1, 2, 32-39, 84
Maxillaria acervatal^ 34, 37, 39-
40, 71
MaxuTaria aciantha 11, 34, 40, 58,
76
Maxillaria acostae 27, 39, 40, 73
Maxillaria acuminata 75
Maxillaria acutifolia 23, 38, 41,
44, 60, 65
Maxillaria adendrobium 6, 33, 41
Maxillaria admonens 173
Maxillaria adolphii 7, 33, 41-42,
46, 73
Maxillaria alba 17, 37, 38, 42, 44,
68
Maxillaria albertii 56
Maxillaria alfaroi 4, 32, 42-43, 48
Maxillaria amabilis 9, 33T43T65
Maxillaria amparoana 74, 77"
Maxillaria ampliflora 16, 36, 43,
48, 50
Maxillaria anceps 18, 37, 42, 43-
44, 68, 71
Maxillaria angustisegmenta 22,
38, 44, 57
Maxillaria angustissima 20, 37,
44-^5
MaxnTaria appendiculoides 5, 33,
45, 62
MaxHTaria arachnitiflora 26, 39,
45, 68
MaxHTaria atrata var. brachyantha
67
Maxillaria attenuate 25, 39, 45-
46, 51
MaxUTaria aurantiacum 61
Maxillaria auyantepuiensis 62
Maxillaria bicallosa 20, 37, 46, 5 1
Maxillaria biolleyi 7, 33, 427 46,
73
Maxillaria brachybulbon 27, 39,
46-^7, 73
MaxITTaria bracteata 18, 36, 47,
59
Maxillaria bradeorum 16, 36, 43,
47^48, 50
MaxiUaria brenesii 48, 49
Maxillaria brenesii var. longiloba
62
Maxillaria brevilabia 11, 34, 48,
66
Maxillaria brevipedunculata 66
Maxillaria brevipes 73
Maxillaria brunnea 23, 38, 48-49,
54, 62
Maxillaria burgeri 14, 35, 49
Maxillaria cacaoensis 173,T74
Maxillaria caespitifica 27, 35, 38,
49-50, 83
MaxHTana camaridii 34, 36, 50
Maxillaria campanulata 17,~~36,
50
Maxillaria cedralensis 24, 39, 50-
51, 72
Maxillaria chartacifolia 20, 37,
46, 5J
Maxillaria chionantha 26, 39, 45,
51, 68
MaxUTaria chiriquensis 53, 82
Maxillaria cobanensis 83-84
Maxillaria concavilabia 11, 34, 52
Maxillaria conduplicata 7, 33, 3~37
52
MaxiHaria conferta 69
Maxillaria confusa 25, 39, 52-53,
73
Maxillaria costaricensis 13, 35,
38, 53, 57, 83
Maxillaria crassifolia 19, 37, 53-
54
MaxlITaria cryptobulbon 22, 38,
39, 54
Maxillaria ctenostachya 10, 34, 54
Maxillaria cucullata 67, 72
Maxillaria curtipes 82, 84
Maxillaria densifolia 43
Maxillaria dendrobioides 5, 33,
54, 82
MaxnTaria dichotoma 16, 36, 55,
69
Maxillaria discolor 63
Maxillaria diuturna 18, 37, 5_5
Maxillaria dressleriana 23, 38756,
65
Maxillaria elatior 9, 34, 36, 56, 80
Maxillaria endresii 22, 38, 44, 57
Maxillaria endresii var. angustis-
egmenta 44
Maxillaria exaltata 8, 33, 34, 57,
63
MaxiUaria exigua 69
Maxillaria falcata 9, 33, 46, 57-58
Maxillaria flava 14, 35, 49, 5S, 53
MaxiUaria foliosa 40, 7 1
Maxillaria friedrichsthalii 10, 34,
40, 58-59, 76
Maxillaria Tulgens 8, 33, 35, 37,
59, 70
MaxUTaria fuerstenbergiana 74
Maxillaria gatunensis 54
Maxillaria gomeziana 18, 36, 47,
59
MaxUTaria gracilifolia 78
Maxillaria graminifolia 62
Maxillaria grandiflora 43
MaxiUaria guadalupensis 46
Maxillaria haberi 13, 35, 59-60
Maxillaria hedwigiae 23,~3~8,~~39,
41, 60, 65
Maxillaria hedyosma 42
MaxiUaria heidelbergensis 79
Maxillaria horichii 12, 34, 60
Maxillaria houtteana 56, 84
MaxiUaria imbricata 47, 76
Maxillaria inaudita 4, 32, 6_U 79
Maxillaria iridifolia 81
Maxillaria lactea 74
Maxillaria lankesteri 15, 35, 37,
61, 83
MaxUTaria lexarzana 67
Maxillaria linearifolia 5, 33, 45,
61-62
MaxHTaria linearis 83
Maxillaria longiloba 21, 38, 62
Maxillaria longipetiolata 21~38,
62
MaxnTaria lueri 4, 32, 62-63
MaxiUaria luteoalba 44,~~57
MaxiUaria luteorubra 45
Maxillaria macleei 80
Maxillaria maleolens 19, 37, 63
Maxillaria meridensis 8, 33,~~34,
57, 63
Maxillaria microphyton 14, 35,
49, 58, 63-64, 66
Maxillaria minor 15, 36, 55, 64,
69
MaxiUaria mombachoensis 72
Maxillaria monteverdensis 9, 34,
64-65
Maxillaria moralesii 24, 38, 60, 65
Maxillaria muscoides 14, 35, 65^
66
MaxUTaria nagelii 42, 44, 56
MaxiUaria nana 80
MaxiUaria nasalis 66
Maxillaria nasuta 19, 37, 66
Maxillaria neglecta 11, 34~4"8, 52,
66, 69, 72, 78
MaxUTaria nicaraguensis 6, 33,
34, 67
Maxillaria nutantiflora 80
Maxillaria obscura 24, 39, 51, 67
MaxiUaria ochroleuca 50
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
179
Maxillaria oreocharis 17, 36, 37,
38, 42, 67-68
Maxillaria pacfiyacron 26, 39, 45,
51, 68, 74
Maxillaria paleata 15, 36, 55, 64,
68-69, 76
MaxUTaria pallidiflava 64
Maxillaria parviflora 11, 34, 52,
69
MaxUTaria parvilabia 6, 33, 69-70
Maxillaria picta 32
Maxillaria piestopus 21, 38, 70, 84
Maxillaria pittieri 8, 33, 35,~TO
Maxillaria planicola 13, 35, 607 71
Maxillaria ponerantha 9, 34, 5T.
40, 71
Maxillaria porrecta 48, 49
Maxillaria powellii 49
Maxillaria pseudoneglecta 12, 34,
60, 66, 69, 71-72
Maxillaria pterocarpa 47
Maxillaria pubilabia 74
Maxillaria punctostriata 24, 39,
51, 67, 72
Maxillaria quadrata 7, 33, 42, 46,
72-73
MaxflTaria ramonensis 25, 39, 53,
58, 73
Maxillaria reichenheimiana 26,
39, 45, 68, 73-74
Maxillaria rhodosticta 76
Maxillaria ringens 22, 38, 49, 74
Maxillaria rodrigueziana 25, 39,
74-75
MaxiUafia rouseauae 74
Maxillaria rubrilabia 72
Maxillaria rufescens 32, 41, 56, 65,
84
Maxillaria sanguinea 12, 35, 38,
75, 78
MaxUTaria schistostele 35, 75-76
Maxillaria schlechteriana 16. 36,
55, 64, 68, 76, 77
Maxillaria scorpioidea 10, 34, 40,
58, 76-77
Maxillaria semiorbicularis 47, 48
Maxillaria serrulata 19, 36, 77
Maxillaria sessilis 53
Maxillaria sigmoidea 15, 36, 55,
64, 77, 81
Maxillaria simulans 69
Maxillaria speciosa 75
Maxillaria squamata 81
Maxillaria stenophylla 52
Maxillaria stenostele 81
Maxillaria striatella 81
Maxillaria strumata 21, 38, 77-78
Maxillaria suaveolens 16, 3^7 5T£
78
MaxUTaria surinamensis 69
Maxillaria tenuifolia 12, 35, 75,
78
MaxHTaria tigrina 13, 32, 35, 78-
79
MaxUTaria tonduzii 6, 33, 41, 79
Maxillaria triangularis 56
37,
38,
Maxillaria trilobata 5, 33, 34, 61,
79
MaxWaria trinitatis 48
Maxillaria tubercularis 10, 35, 36,
80
MaxWaria tuerckheimii 74
Maxillaria turialbae 58
Maxillaria umbratilis 17, 50, 80
Maxillaria uncata 12, 35, 80-ST
Maxillaria vagans 47
Maxillaria vaginalis 15, 36, 55, 64,
65, 77, 8_[
Maxillaria valenzuelana 20,
81-82
MaxUTaria valerioi 4, 33, 82
Maxillaria variabilis 13,35,
50, 53, 82-83, 84
Maxillaria villosa 63
Maxillaria vinosa 80
Maxillaria vittariifolia 27, 39, 83
Maxillaria wercklei 15, 35, 37, (TT,
64, 83
MaxiUaria wrightii 68
Maxillaria wrightii var. imbricata
69, 76
Maxillaria xylobiiflora 47
Maxillaria yzabalana 74
Maxillariinae 1-2
Mesospinidium §9, 132
Mesospinidium endresii 133
Mesospinidium horichii 96, 132
Mesospinidium leochilinum iJI
Mesospinidium warscewiczii 96,
133
Miltonia 118
Miltonia endresii 133, 134
Miltonia schroederiana 147
Miltonia stenoglossa 147, 148
Miltonia superba 133
Miltonioides confusa 147
Miltonioides stenoglossa 147
Miltoniopsis 88, 133
Miltoniopsis vexillaria 134
Miltoniopsis warscewiczii 97,
133-134
Mormolyca 2, 70, 84
Mormolyca fuchsii 7T)
Mormolyca lineolata 84
Mormolyca ringens 28, 32, 84
Neoescobaria 169
Neokoehleria 160
Neourbania adendrobium 41
Neourbania nicaraguensis 67
Notylia 89, 127, 134, 173
Notylia barkeri 135
Notylia brenesii 134
Notylia cordesii 129
Notylia lankesteri 134-135,
Notylia linearis 130
Notylia paniculata 131
Notylia pittieri 97, 135
Notylia ramonensis 131
Notylia trisepala 97, 135
Notylia turialbae 135
Odontoglossum 133, 140
Odontoglossum anceps 151
Odontoglossum aspasia 115
Odontoglossum biberianum 115
Odontoglossum bictoniense 157
Odontoglossum cariniferum 139
Odontoglossum chiriquense 152
Odontoglossum confusum 147
Odontoglossum convallarioides 150
Odontoglossum coronarium var.
chiriquense 152
Odontoglossum egertonii 151
Odontoglossum hortensiae 157
Odontoglossum krameri 165
Odontoglossum krameri var. album
165
Odontoglossum krameri var. smi-
thianum 165
Odontoglossum leucomelas 148
Odontoglossum maculatum 158
Odontoglossum oerstedii 165
Odontoglossum pauciflorum 148
Odontoglossum principissa 115
Odontoglossum schlieperianum 159
Odontoglossum schroederianum
147
Odontoglossum section Myanthium
151
Odontoglossum stellatum 158
Odontoglossum stenoglossum 147
Odontoglossum Subgenus Osmo-
glossum 150
Odontoglossum warscewiczianum
133
Odontoglossum warscewiczii 133
Oliveriana 163
Oncidiinae 86-89
Oncidium S8T S9~, 124, 133, 135-
137, 151, 154
Oncidium advena 149
Oncidium altissimum 133
Oncidium ampliatum 101, 136,
137
Oncidium angustisepalum 143
Oncidium ansiferum 103, 137,
138, 147
Oncidium ascendens 98, 136, 138
Oncidium asparagoides 139
Oncidium baueri 146
Oncidium bernoullianum 137
Oncidium bifrons 149
Oncidium bracteatum 101, 136,
_138-139, 149
Oncidium brenesii 144
Oncidium bryolophotum 100,
136, 139
Oncidium cabagrae 142
Oncidium calyptostalix 144
Oncidium cardiochilum 145
Oncidium cariniferum 99, 136,
139-140
Oncidium carthagenense 98, 136,
140
Oncidium castaneum 142
Oncidium caudatum 117
180
aa/ai
FIELDIANA: BOTANY
Oncidium cebolleta 98, 136, 138,
140
Oncidium cheirophoroides 144
Oncidium cheirophorum 101,
136, 140-141
Oncidium cheirophorum var. exaur-
iculatum 141
Oncidium chelidon 145
Oncidium chelidonizon 145
Oncidium costaricense 142
Oncidium crispum 155
Oncidium cristagalli 100, 136,
141-142
Oncidium decipiens 141
Oncidium dichromaticum 102,
137, 142
Oncidium dichromaticum var. rosea
142
Oncidium dielsianum 141
Oncidium ensatum 103, 137, 142
Oncidium exauriculatum 141
Oncidium fulgens 143, 145
Oncidium globuliferum 100, 136,
142-143
Oncidium globuliferum var. costar-
icense 142
Oncidium gnoma 155
Oncidium graciliforme 143, 144
Oncidium guttulatum 146, 147
Oncidium henricigustavi 138
Oncidium heteranthum 139
Oncidium hondurense 156
Oncidium isthmi 104, 137, 143,
146-147
Oncidium klotzschianum 102,
137, 143-144, 145
Oncidium kramenanum 154
Oncidium kramenanum var. res-
plendens 155
Oncidium lankesteri 138
Oncidium lintriculus 149
Oncidium luridum 135, 136, 149-
150
Oncidium luteum 101, 136, 144
Oncidium macrorhynchum 141
Oncidium megalous 139
Oncidium microphyton 139
Oncidium naranjense 147
Oncidium oberonia 156
Oncidium obryzatoides 102, 137,
144, 145
Oncidium obryzatum 143, 144, 145
Oncidium ochmatochilum 136,
145
Oncidium ornithorhychum 150
Oncidium paleatum 138, 139
Oncidium panduriforme 99, 136,
145-146
Oncidium papilio var. kramerianum
155
Oncidium parviflorum 102, 137,
146
Oncidium peliograma 146
Oncidium pittieri 144
Oncidium planilabre 136, 150
Oncidium polycladium 104, 137,
143, 146-147
Oncidium poweWi 148
Oncidium pubes 155
Oncidium pumilio 155
Oncidium pumilum 150
Oncidium pusillum 156
Oncidium rechingerianum 142
Oncidium schroederianum 98,
136, 147
Oncidium sclerophyllum 143, 144
Oncidium stenobulbon 103, 136,
137, 138, 147
Oncidium stenoglossum 99, 147-
148
OIK ulmm stenotis 103, 137, 148
Oncidium storkii 99, 136, T3~9,
148-149
Oncidium teres 98, 136, 138, 140,
149
Oncidium tetraskelidion 144
Oncidium titania 155
Oncidium tonduzii 146
Oncidium tricuspidatum 124
Oncidium turialbae 148
Oncidium varians 143, 145
Oncidium warscewiczii 100,
149
Oncidium wercklei 142
Orchidotypus hispidulus 153
Orchidotypus muscoides 153
Ornithidium 1
Ornithidium alfaroi 48
Ornithidium anceps 44, 66, 71
Ornithidium aurantiacum 61
Ornithidium biolleyi 46
Ornithidium bracteatum 47
Ornithidium chloroleucum 69
Ornithidium conduplicatum 52
Ornithidium confertum 69
Ornithidium costaricense 57, 58
Ornithidium fulgens 59
Ornithidium lankesteri 72
Ornithidium neglectum 66
Ornithidium paleatum 68
Ornithidium pallidiflavum 64
Ornithidium parvulum 63
Ornithidium pittieri 70
Ornithidium ramonense 58
Ornithidium rhomboglossum 69
Ornithidium schlechterianum 76
Ornithidium squamatum 81
Ornithidium sigmoideum 77
Ornithidium stenophyllum 52
Ornithidium strumatum 77
Ornithidium tonduzii 41
Ornithidium wercklei 83
Osmoglossum 88, 150
Osmoglossum acuminatum 151
Osmoglossum anceps 151
Osmoglossum convallarioides
104, 150-151
Osmoglossum egertonii 104, 150,
151
Otoglossum 88, 151
136,
Otoglossum chiriquense 104, 151-
152
Pachyphyllum 87, 120, _152
Pachyphyllum costaricense 152-
153
Pachyphyllum crystallinum 105,
152, 153
Pachyphyllum hispidulum 105,
152, 153-154
Pachyphyllum muscoides 153
Pachyphyllum pastii 1 53
Pachyphyllum stuebelii 153
Palumbina 150, 159
Papiliopsis krameriana 155
Pittierella calcarata 29
Plectrophora 87, 154
Plectrophora alatsFTDS, 154
Pleuranthium adendrobium 41
Pleurothallis valenzuelana 81
Ponera adendrobium 41
Pseudomaxillaria 1
Pseudomaxillaria chloroleuca 69
Pseudomaxillaria parviflora 69
Psychopsis 88, 154
Psychopsis krameriana 105, 154-
155
Psygmorchis 87, 124, 155
Psygmorchis glossomystax 156
Psygmorchis gnoma 156
Psygmorchis pumilio 106, 141,
155-156
Psygmorchis pusilla 155, 156
Rhynchostele 88, 156-157, 159
Rhynchostele cordata 158
Rhynchostele bictoniensis 106,
157
Rhynchostele hortensiae 106,
157-158
Rhynchostele maculata 158
Rhynchostele stellata 106, 158
Rhynchostele cervantesii 157
Rodriguezia 87, 119, 127, 159
Rodriguezia candelariae 12T
Rodriguezia compacta 107, 159
Rodriguezia dressleriana 159
Rodriguezia inconspicua 121
Rodriguezia lanceolata 159
Rodriguezia leochilina 121
Rossioglossum 88, 159
Rossioglossum schlieperianum
107, 159-160
Rossioglossum williamsianum 1 60
Scaphyglottis parviflora 69
Scaphyglottis pendula 66
Scelochilus 87, 119, 159, 160
Scelochilus aureus 107, 16TP161
Scelochilus tuerckheimii 160-161
Sepalosaccus 1
Sepalosaccus horichii 60
Sepalosaccus humilis 77
Sepalosaccus strumatum 77
Sigmatostalix 89, 161
Sigmatostalix ad a in si i 109, 161
ATWOOD & MORA DE RETANA: FLORA COSTARICENSIS
181
Sigmatostalix brownii 108, 161,
162
Sigmatostalix costaricensis 162,
163
Sigmatostalix guatemalensis 1 63
Sigmatostalix hymenantha 108,
161, 162
Sigmatostalix macrobulbon 108,
161, 162
SigmatostalTx picta 108, 161, 162-
163
Sigmatostalix poikilostalix 1 62
Sigmatostalix unguiculata 107,
161. 163
Solenidium endresii 133
Symphyglossum 151
Systeloglossum 87, 163
Systeloglossum acuminatum 109,
163, 164
Systeloglossum costaricense 109,
163, 164
Ticoglossum 88, 164-165
Ticoglossum krameri 109, 165
Ticoglossum krameri var. album
165
Ticoglossum krameri var. smi-
th ianum 165
Ticoglossum oerstedii 110, 165
Tolumnia pumilio 1 56
Tolumnia pusilla 1 56
Trichocentrum 87, 154, 165-166
Trichocentrum alatum 154
Trichocentrum brenesii 111, 166
Trichocentrum caloceras 111,
166, 167, 168
Trichocentrum candidum 168
Trichocentrum capistratum 111,
166, 167, 168
Trichocentrum costaricense 111,
166, 167-168
Trichocentrum cymbiglossum
112, 166, 168
Trichocentrum dianthum 110,
166, 168
Trichocentrum estrellense 110,
166, 168-169
Trichocentrum panamense 167
Trichocentrum pusillum 1 67
Trichocentrum pfavii 110, 166,
169
Trichocentrum pfavii var. zonale
169
Trichocentrum saundersianum 169
Trichocentrum saundersii 169
Trichocentrum zonale 169
Trichopilia 88, 118, 169-170
Trichopilia coccinea 170
Trichopilia crispa 171
Trichopilia dasyandra 118
Trichopilia galeottiana 113, 170,
171, 172
Trichopilia kienastiana 171
Trichopilia maculata 112, 170,
172
Trichopilia marginata 113, 170-
171
Trichopilia picta 170
Trichopilia pusilla 1 19
Trichopilia suavis 112, 171
Trichopilia tortilis 113,T70, 171-
172
TricHopilia turialbae 112, 172
Trigonidium 1, 32, 84, 85 ~
Trigonidium amparoanum 86
Trigonidium egertonianum 28,
85, 86
Trigonidium insigne 86
Trigonidium lankesteri 28, 85, 86
Trigonidium ringens 84
Trigonidium riopalenquense 28,
85, 86
Trigonidium seemannii 85
Trizeuxis 87, 127, 172
Trizeuxis fa 1 cat a 1T37172
Warmingia 87, 127, 172-173
Warmingia margaritacea 173
Zygopetalum bicallosum 46
182
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