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Full text of "Flora Costaricensis"

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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The paper used in this publication meets the requirements of ANSI/NfSO Z39.48-1992 (Permanence of Pap 



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 -V 5 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 ^ 1 A 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 1 A 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 1 A 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' 3 r 9 IK cm ^Pf"- , Roots wh f' to l 

,...-., o /-i mm in diameter, rseudobulbs 2(l)-iohate, ovoid, com- 

totype: K). Maxillaria fohosa Ames & C. presse d, 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 1 A 
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 1 A 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 57 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 V 5 , 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 l h. 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 1 A 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 1 A- 1 A, 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 1 A 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 1 A 
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 1 A 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, 11.4 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 1 A 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 1 A, 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 l h 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 1 A, 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 1 A 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, 1.54 
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, 34.5(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 1 A, 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, 68 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, 1.41.7 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 1.31.5 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, 13 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, 1.51.8 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 



FIELDIANA: BOTANY 



UNIVERSITY OF ILLINOIS-URBANA