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MISSOURI BOTANICAL GARDEN
ERRATA
Page 4, line 26—for “35. Совма” substitute “35. BorHRIOCHILUS"; line 28
— for "column" read “pollinia.”
Page 145, caption of fig. 2, second line—-for “var. ioense" read “var. ioensis.”
Page 146, Explanation of Plate 1—for "var. ioense” read “var. ioensis.”
Annals
of the
Missouri Botanical
Garden
Volume XXXIII
1946
With 19 Plates and 276 Figures
Published quarterly at Galesburg, Illinois, by the Board of Trustees of
the Missouri Botanical Garden, St. Louis, Mo.
Entered as second-class matter at the post-office at Galesburg, Illinois
nder the Act of March 3, 1879.
.
.
ы .
. • Фо го P
* . . '.. . o
ME C . . H .
ы • ee of o
* LIS .
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CROTO
Pu Pu Их ) fe
ОСА, scrot
Annals
of the
Missouri Botanical Garden
A Quarterly Journal containing Scientific Contributions from the
Missouri Botanical Garden and the Henry Shaw School of Botany of
Washington University in affiliation with the Missouri Botanical
Garden.
Information
The ANNALS OF THE MissouRI BOTANICAL GARDEN appears four times
during the calendar year: February, April, September, and November. Four
numbers constitute a volume.
Subscription iy $10.00 per volume
Single Numbers |... 2.50 each
Contents of previous issues of the ANNALS OF THE Missouni BOTANICAL
GARDEN are listed in the Agricultural Index, published by the H. W. Wilson
Company.
STAFF
OF THE MISSOURI BOTANICAL GARDEN
Director
СЕОКСЕ Т. Moore
Assistant to the Director
Henry N. ANDREWS
HERMANN VON SCHRENK, Ковект E. Woopson, Jr.
Pathologist Assistant Curator of
the Herbarium
ЈЕЗЗЕ M. GREE Henry М. ANDREWS,
Curator of $$ Herbarium Paleobotanist
CARROLL У. Dopcr, Ковект №. 5сн
Mycologist Research Munt"
EDGAR ANDERSON, Мета. C. Hor
Geneticist кұс ion Editor
Publications
BOARD OF TRUSTEES
OF THE MISSOURI BOTANICAL GARDEN
President
СЕОКСЕ C. Нітснсоск
Vice-President
DANIEL К. CATLIN
Second Vice-President
EUGENE PETTUS
L. Ray CARTER Кіснакр J. Lockwoop
Поргеу FRENCH СЕОКСЕ T. Moore
Jonn S. LEHMANN А. WESSEL SHAPLEIGH
ETHAN А. Н. SHEPLEY
EX-OFFICIO MEMBERS
ARTHUR Н. COMPTON, STRATFORD L. Mor
Chancellor of Washington арзй of the Academy of
iversity Science of St. Lou
A. P. KAUFFMANN, WILLIAM SCARL
Mayor of the City of Bishop p. ei Bs. of
St. Louis Miss
ELMORE M. PUTNEY
President of the Board of Education of St. Louis
GERALD Urmici, Secretary
TABLE ОЕ CONTENTS
Flora of Panama. Part ІП. Fascicle 2 (Orchidaceae)
4 Robert E. Woodson, Jr.,
“Robes W. Schery, and Collaborators
Contributions to Our Knowledge of American Car-
boniferous Floras. VIII. Another Medullosa from
WA. Henry N. Andrews and Jules A. Kernen
Maize in Mexico—A Preliminary Survey. _
ой И TAM төге
A Cytological Study of Yeast (Saccharomyces cere-
WEN) S Lillian Nagel
Contributions to Our Knowledge of American Car-
boniferous Floras. ІХ. Some Petrified Seeds from
Iowa Ellen M. Kern and Henry N. Andrews
The Gallatin Fossil Forest... 27758
— Henry N. Andrews and Lee W. Lenz
Flora of Panama. Part III. Fascicle 3 (Orchidaceae,
cH IE). 1. 3 Robert E. Woodson, Jr.,
Robert W. Schery, and Collaborators
General Index to Volume XXXIII | | .
PAGE
1-140
141-146
147-247
249-288
291-306
309-313
315-404
405-408
ЕГОКА ОЕ РАМАМА
E BY
ROBERT E. WOODSON, Jr.
/ AND
ROBERT W. SCHERY
AND COLLABORATORS
PART III
Fascicle 2
ORCHIDACEAE (Williams)
(in part)
ANNALS
OF THE
Missouni BOTANICAL GARDEN
Vol. XXXIII FEBRUARY, 1946
No.
-
FLORA OF PANAMA
Part III. Fascicle 2
ORCHIDACEAE
Bv LOUIS O. WILLIAMS
Epiphytic, terrestrial, rarely semi-aquatic or saprophytic, perennial herbs. Veg-
etative growth of two main types: (1) Termed MoNoropiArzs, in which the
main axis or stem grows steadily upward year after year, the annual growths at
length being indistinguishable and a monopodium being formed; the inflorescences
rne on lateral shoots. (2) Termed ЗумроглАТЕ$, in which a new growth de-
velops laterally from the base of the previous year's growth, which is completed
and matures in a few months. In the ЗУМРОГАТЕ$ if the annual growth is
terminated by an inflorescence the plant is termed acranthous (terminal flower-
ing) ; if the inflorescence originates on lateral branches (usually at the base of or
laterally on a pseudobulb) and the annual growth is terminated only by leaves
the plant is termed pleurantbous (lateral flowering). Stems (secondary) of many
orchids developed into thickened or swollen structures (pseudobulbs) which act
as storage organs for food and moisture. Flowers hermaphroditic and gynandrous,
occasionally monogamous or polygamous, the male and female ones being very
different; zygomorphic; superior. Perianth of two tripartite whorls, alternating,
often variously united; in the same whorl the even pair similar, the odd one usually
different in form. Sepals 3, similar, or the dorsal differing from the laterals, free
or variously united. Petals 3, two similar and termed petals, the other one usually
strikingly different and termed lip or labellum. Stamens in two tripartite whorls,
only one (MoNANDRAE) ог two (DIANDRAE) fertile, the others, along with the
styles and sterile stigmas, forming the column by consolidation; in the MONANDRAE
the odd stamen of the outer whorl fertile; in the Dianprar the paired stamens
of the inner whorl fertile. Pollen (except in the СүркІРЕрПОІрЕАЕ) usually
consolidated into pollinia. Stigmas 3, usually only one or two fertile (if two,
often confluent and appearing as one). The column, the central structure іп the
flowers, which is made up of the consolidated styles and filaments along with the
suppressed anthers and stigmas, is diagnostic of the family. Fertile anthers and
stigmas usually borne toward the apex of the column. Ovary 1-celled or rarely
3-celled. Seeds numerous, minute, lacking endosperm.
Issued March 8, 1946.
(107) (1)
[Vor. 33
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN
A cosmopolitan family of about 600 genera and 15,000—20,000 species. Most
abundant in tropical regions of the world.
The Orchidaceae is an extremely complex family, the morphology of which is
often not too well understood. The genera, in some groups, are technical and per-
haps not too well differentiated; in other groups they are fairly well differentiated.
With the exception of a few terrestrial genera and even fewer of the epiphytic ones,
the respective genera are limited to either the eastern or to the western hemis-
phere. A few genera contain a very large number of species (Dendrobium, eastern
hemisphere, probably more than 1000; Epidendrum, western hemisphere, possibly
nearly 1000). The species of orchids, as a whole, are distinctive, and extreme
variation within species is not too common. Terrestrial species, especially those of
temperate climates, are inclined to be more variable than are epiphytic ones. Species
are inclined to be restricted in distribution although terrestrials and epiphytes
growing at low elevations often range widely.
The greatest concentrations of orchids occur within twenty degrees of the
equator. While some are found at low elevations, the great majority of them
grow on mountains within the tropics. Malaysia and tropical America are richest
in species and genera, with Africa a poor third. New Guinea probably contains
more species than any comparable land area, although it is still not well known
botanically. Colombia is perhaps the richest in species of any comparable area in
the western hemisphere.
KEY TO THE SUBFAMILIES, TRIBES AND GENERA
a. Stamens 2, a third usually MONIO E m a large рен. роПеп
Ға
гапшаг, not united into masses or mily DIAN
тті а
аа. Stamen 1, the — abortive or forming staminodes; pollen c
soli a)
solidated into masses d bodies (pollinia) ...................... Su bfamily MONANDRAE
b. Caudicle and dud feno from the base of the pollinia; anthers
rec e less resupinate, very closely adnate to t roa
e
ased column, never deciduous after flowering; pollinia always gran-
ular (sectile ribe II OPHRYDOIDEAE
bb. Caudicle and gland arising from the apex of the pollinia; —
о npn ла nt, the filaments short and slender, generally ar-
rowly joined to the column, usually deciduous but if persistent soon
wither
e. Pollinia gn soft; anthers commonly persistent but wither-
ing; inflorescence (normally) always terminal................ Tribe ІП PotycHONDREAE
. Pollinia waxy or cartilaginous; anther commonly soon ез
inflorescence terminal or lateral e ТҮ KEROSPHAEREAE
o
о
I. Subfamily Пллмрклв, Tribe CvPRiPEDILOIDEAE. Two genera іп
1. SELENIPEDIUM
anama.
a. Stems sig leafy; leaves thin
aa. Stems very short or — none, leaves coriaceous 2. PHRAGMIPEDIUM
т Subfamily. MOoNANDRAE, Division Biserowax, Tribe OPHRYDOIDEAE.
e genus in Panam, 3. HABENARIA
ш. = ај aig А mas, Divis sion ACROTONAE, Tribe PoLyCHONDREAE.
ma.
nag Ае “distribu ted in five subtribes in Panam
a. Prin r more or less prone, incumbent.
(108)
19461
FLORA OF PANAMA (Orchidaceae)
b. Leaves not articulated, persistent; stems not cane-like.
c. Terrestrial d with fleshy roots; small or large rather showy
urplish flow 4. PocoNiA
сс Scandent herbs am epiphytic, without fleshy roots; flowers
large, y whit 5. VANILLA
bb. Leaves articulated, лера stems cane-like.
c. Pollinia
а. "s ase > the lip strongly gibbous or m ornamented with
o prominent calli; flowers ipe ied mall 6. ELLEANTHUS
dd. ~ of the lip not strongly gib or же fos with or
without calli at the base; денді үлек деге very large.............. 7. SOBRALIA
сс. Pollinia 4
aa. Anther more or less ea: nec erect or suberect.
b. Leav t plicate-nerved, usually soft.
c. Roots fasciculat iri
d. Lip uppermost, adaxial; flowers not resupin
e. Pe tals, and sometimes the lip, insert de on the column............
c
f. Lip inserted at the
5. Is not united into а basal 11.
£g. Sepals united at the base into a slender tu 9.
f. Lip united at the I to a cup formed of the united
epals, helmet-shape 10.
dd. Lip lowermost, abaxial; ^ ers resupina 13
cc. Roots not А сол ичейин Ди а from the si on the lower part
of the 14.
Leaves оиа peut 22) or subcoriaceous 15;
IV. Subfamily МОМАМРКАЕ, Division Acro AE, Tribe KEROSPHAEREAE.
Seventy-two era, distributed in twenty-six subtribes, in Panama.
a. Series CRANTHAE. Inflorescence normally terminal or by abor-
tion of terminal inflorescence axillary in uppermost leaves. (See also
LocKHARTIA)
b. Viscid disc, when present, Б. ак babi the apex of the pollinia,
commonly gr rudim none
c. Ovary articulated to the иегі pedicel persistent; stems slender,
rigid, sometimes reduced, usually unif
d. Sepals all distinctly connate at the bas
e. Sepals also connate at den jeas open bétwéen tips and bases... 18.
ee. Sepals not con heir
Sepals forming a narrow or "айкай tube at the base.
2. Inflorescence 173
gg. c yg a ‘ingle ‘flo ower, or rarely 2"Howers;.........2 19.
ff. Sepals more or less rotate, not forming a narrow tube at
the base 6.
dd. -— not E — connate at the base, at least the dorsal
pal free arly
e. “Blade of the pads diu i.e. strongly bilobed (except
tal not transverse, not strongly bilobed.
ee. Blade
column-foot; lip with a hood-shaped
Т; ta
f. Petals inserted on the
ca gule at the b
N
N
ase
ff. Petals n not жас rted оп the column-foot nor with а hood-
shaped callus or ты at the base
8. Inflorescence terminal or at Мей subterminal оп the
secondary stems
N
ә
gg -— қалы about the middle of the sec-
8. PALMORCHIS
. PONTHIEVA
CRANICHIS
STENOPTERA
PRESCOTTIA
SPIRANTHES
ERYTHRODES
CORYMBORCHIS
CRYPTOPHORANTHUS
PHYSOSIPHON
MASDEVALLIA
STELIS
21. LEPANTHES
ACOSTAEA
PLEUROTHALLIS
SCAPHOSEPALUM
ма 1 a with the
red.
се. 4. not "articulated to the e cel;
hy, ver - flow
wer; stems slender or fles - to
d. "Pollinia Sot appendages, » e. d disc or caudicle.
e. Column very short; We ad өсері іп үс лердің erect..
Column elongated; anther ји > incumben
dd. Роба nin ed, i. e. with at least a аа: viscid disc
audicle with a viscid apex
(109)
. MALAXIS
. Lrparis
[Vor. 33
ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
. Column лет lip more ог less connate with the base of
"а colum
f. Pollinia 4, two in each cell of the anther.
. Lip geniculate at its junction with the
gg. Lip not өзекте ulate at its junction with the colum
h. Flowers large and showy; stems always ебут 29. CATTLEYA
hh. Flowers usually пос pae and showy; stems either
атару ог ebu
with two large hollow horn- like processes; lip
column 26. HEXISEA
28. DiACRIUM
or nearly so :
ii. Lip without hollow horn-like processes; lip com-
pasci connate with the column at base but not
27. EPIDENDRUM
alw
ff. бє 8, du ur in each cell of = anther,
1al 31. npe
30.
ze, 1al
into a krenen foot at the base; lip hardly
P
connate to the column or at most connate at the base of the
33. PLATYGLOTTIS
a
"те dstichous and scattered along an elongated stem.
6 -
34. IsocuiLus
h. энем
Ki ТА ТЕТЕ 4. еле ы шын кыы E
gg. Leaves пос “шш һоив оп ап een и anm terminal
nm eim or short indurated s
32. SCAPHYGLOTTIS
35,
ff. Pollin
bb. Viscid disc voa о with the margins well defined, arising
from the apex of the colum
with a foot; ч with pseudobulbs; pollinia 4, ог 4
irs.
a
„©
ша
=
37. GALEANDRA
36. POLYSTACHYA
38. EPIDANTHUS
16
. Column footless; plants without pseudobulbs; pollinia E
aa. Series B. PLEURANTH Inflorescence lateral, arising near the base
of the pseudobulb or in m axils of the lower leaves or sheaths. (This
eries.
A. Subseries a. Sympopraces. Plants forming sympodium,
stems approxi imate or superimposed and the apical growth Pirna
termin
B. Pollinia without a stipe; viscid disc commonly rudimentary or the
La of i caudicle glutinous or none
. Rhizome short; terrestrial plants with the stems bulbose, usually
Fer гэн buried in the edi leaves plicate.
59. CALANTHE
D. Lip with a spur
DD. Lip without a 40. BLETIA
сс. a — ог је eic plants Raa Ay pseudo-
bulbose s ved; leaves r plica
D. га sdobs is eme (i. e. of se d ps^n = "ВР term-
nal on which bears c == т, many-leaved; leaves
eet po кафа relativ 41. Cysts
DD. Pseud т heteroblastic (ы е. single node with one or
more term nal leaves), usually M. ee coriaceous or fleshy,
€ НР elatively small 42. BULBOPHYLLUM
BB. Pollinia бен а prominent ы sometimes short; viscid disc distinct.
ollinia of ire exture, easily mashed
Lip spurred or wit saccate bas 43. EULOPHIA
=
DD. Lip not spurred nor saccate ас the bas
E. Lateral iniri nd column Шын а prominent mentum;
base of the han and peduncle not enclosed in а submem-
branaceous sheath
EE. Lateral sepals and column-foot forming an inconspicuous
ment Жы; a ase of the leaves and peduncle enclosed in a sub-
44. WARREA
ous sheath 45. GOVENIA
mbra
СС. Pollinia buon in texture, not easily mashed.
(110)
19461
FLORA ОЕ PANAMA (Orchidaceae)
D. Leaves convolute in verna
E. Pseudobulbs large, ACH homoblastic, many-leaved.
. Flowers perfect, monomorphic; column twiste 46. MORMODES
pe
FF. Flowers dimorphic or trimorphic (rarely perfect); column
not twiste
G. Ee “thik, imet in male flowers usually with
wo retrorse antenn 47. CATASETUM
GG. Ба slender, ud or arcuate, without retrorse
antennae
EE. crie са heteroblastic, 1- to few-leaved.
tinuous with the base of е и or solidly at-
кар to ди pe colum re 79% ot articulated, more or
less оо divided into ап Ем and а hypochile
or rarely
G. Бе "s much narrower than the sepals, usually in-
column.
serted on the base of the
48. CYCNOCHES
©; Е saccate ed nF like 57. CORYANTHES
H. Epichile sacca .56. GONGORA
Е. fab not dis em narrower than the sepals, from a
little n wer to broader
H. Epichile saccate and tuac: like
HH. Epichile not saccate.
I. с е connate and forming a distinct men-
tum base; flowers about 1.5 cm. long,
П. Me sepals free or if connate not forming a
ct mentum at the base.
57. CORYANTHES
49. COELIOPSIS
ip entir 50. SIEVEKINGIA
JJ. Lip not entire, usually divided into a hypochile
pichile.
R lip
shallowly 3-lobed, the «неде E ES ud 55. STANHOPEA
KK. Hypochile of lip usually no ncave-saccate
or if so then the lip иы divided and
lobe
‚ ырда ог p lobe of lip narrowly
lan ate, minate
LL. Epichile or terminal e of lip not narrow-
ly lanceolate, acum
M. Lateral lobes 21 the E p (pleuridia, meso-
chile ) erect or at least not parallel to the
epic
N. Rachis of Mc son and sepals dor-
sally brown pilose-pubescen
NN. а of ўн ўч bua sepals not
ubescent.
O. to жас of the hed joined by a
large central callus; hypochile of the
lip бх о. at least as bid as the lateral
lobes
OO. Lateral lobes of the lip not joined
y a ede callus; hypochile of the
54. NEOMOOREA
1. KEGELIELLA
52. ACINETA
and inconspicuous 53. PERISTERIA
MM. Lateral Mo d the lip (pleuri dia) more
or less parallel to the ne ensiform.... 55. STANHOPEA
FF. Lip articulated to the apex of the colu
. Inflorescence basal, i.e. arising Кы e he ы of the
pseudobulb; lip ueaslly provided with a long low callus.
H. Inflorescence consisting of a single flower
HH. Inflorescence a few- to several-flowered raceme.
I. Flower with a conspicuous spur-like mentum............ 58
60. LYCASTE
. XYLOBIUM
II. Flowers without a conspicuous spur-like mentum.... 59. BIFRENARIA
(111)
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
GG. Inflorescence suprabasal, i.e. in the axils of the lower
sheaths; lip usually provided with a transverse callus
crest
or
DD. Leaves MEN in vernation.
E. Colum
61. ZYGOPETALUM
uced i a foot and рест а mentum with
the њи sepals; а Һа de duced, emarginate.
F. ж llus of the lip transverse, sd fabellate at еше base of
e lip Paga inan suprabasal, n the axils of the
hath or sheath-like pain риа Mod or rudi-
ary.
G. “Callas at the гені of the lip pectinate ог frin
GG. Bo ый the base of the lip not B un
ire or bet obscurely lob
oie 24 obed, or — into a narrow basal and а
pical pa
FF. Callus “of = lip ІЗ often puc or
rarely none; inflor ce borne from the base of a pseudo-
bulb; ми germ well. pope or Pd elon-
BOO canc 64. HUNTLEYA
inate га fringed.
2. CHONDRORHYNCHA
3, WARSCZEWICZELLA
gated s
С. Lateral S forming a spur at pe base about as long
or longer than the blades of the sep
7. CRYPTOCENTRUM
on
: nnate and forming a pei Aud at the base;
lip not half as long as a sepals 6
a tube at я base;
the sepals.............. 65. MAXILLARIA
EE. Column on rostellum commonly и ди sometimes
subulate or
Е. An ther incumbent; rostellum porrect or deflexed, never
ascen
© Base a the lip spurred or saccate or appendaged at the
bas
. TRIGONIDIUM
H. Base of lip forming a spur or deeply saccate.
I. Lateral sepals free
II. Lateral sepals con
HH. Base of the lip eim rns to long appendages; јен
"e wed connate and commonly spurred or sac-
{ 8. TRICHOCENTRUM
. RODRIGUESIA
3
ca the 69. Іоморѕ15
GG. Base p^ the D genres spurred nor saccate nor with
appendages
H. „Ропа 2
· Clinandeium low, slightly excavated, margin entire
and not hyaline.
J. Column winged near the stigma 71. TRIZEUXIS
. Column wingless near the stigma 76. BR
ng the an-
early so, or pe olumn m spreading
wings near b stigma.
J.
ТІ Clinandrium with a high margin enfo
ther or
K. = of the lip. bed the column and
hortly adnate to it at its base
KK. Base of the lip not a column.
jen d nserted on the ear the midak 75. ASPASIA
LL. Lip not inserted on “a co узыйм near the
mid le.
Я
2. TRICHOPILIA
М. Lateral sepals ог е sepals and spoon
Eis ен ; column not winged
the 76. BRASSIA
MM. Lateral sepals or all sepals and petals
audate, or if so iih eligen winged
near yon sti
N. Base of un lip forming a shallow sac.... 73.
MESOSPINIDIUM
NN. Base of the lip not forming a місе
sac.
(112)
1946]
FLORA OF PANAMA (Orchidaceae) 7
O. Column with a pair of nar
stelidia at the т lip ит un-
lobed, pandur use 79. LEOCHILUS
OO. Column la ae не narrow stelidia at
the middle; lip v
P. Lip long- и: lionis ѕтај.. 80. SIGMATOSTALIX
PP. Lip not TN unguiculate; flowers
usually not small.
. Lip erect др me. or con-
tiguous to the column at the
base; calli on the lip s Bush Z
parallel rae not joined; column
usually ged 74. ODONTOGLOSSUM
QQ. Lip pie а; approach-
ing a right angle) from the col-
umn; calli on the lip various
but if 2 then usually joined;
M doji usually winged at the
a
R. Lip usually анау large
and not sharply lobed, us-
ћу lacking calli except at
the base; flowers white.......... 77. MILTONIA
RR. Lip usually relatively small
а а tly 3-lobed, usual-
ly provided У rich cali above
the base (as well as often at
the base); flowers commonly
yellow to mar 78. ONcIDIUM*
JJ. Leaves not articulated, i.e. marcescent; plants
with densely equit md cun elongated stems,
without died obulbs 81. LoCKHARTIA
. Pollinia 4; lip with a retrorse callus near the base...... 82. ORNITHOCEPHALUS
FF. бее erect on the back of the column or erect оп the
apex of the column under the clinandrium; rostellum erect
or ascendin
G. Column пен with stiff hairs 85. TELIPOGON
GG. Column not provided with stiff hai
12% булоон margins small, not prem surrounding
the a 83. NOTYLIA
HH. RUE margins large, almost forming a calyptra
over the anther 84. MACRADENIA
AA. е » MONOPODIALES. E ee forming a monopodium, i.e. the
$ having infinite apical growt
в. Co sha ма ап infrastigmatie Бай stems with equitant leaves.... 86. DICHAEA
BB. Column without а ТАН ligule; stems not having equitant
leaves, priate sometimes leafles 4 87. CAMPYLOCENTRUM
ж Oncidium, Miltonia, Оба Brassia, Aspasia, Leochilus and Mesos pinidium are 95%
one genus, naturally, for they merge comp letely. However for purposes of convenience they
best left apart. The first three menti we are и well known си: except for borderline els
even the amateur can soon learn to distinguish them easily by their aspect
(113)
[Vor. 33
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1. SELENIPEDIUM Reichb. f.
SELENIPEDIUM Reichb. f. Xenia Orch. 1:3, 2. 2. 1854; emend. Pfitz. in Engl.
Pflanzenr. IV. 50 (Heft 12): 27. 1903.
Solenipedium Beer, Prakt. Stud. Orch. 310. 1854.
Tall terrestrial herbs with leafy, often branched stems. Leaves narrow, many-
nerved, inarticulate. Inflorescence terminal, racemose, many-flowered. Flowers
small to medium-sized, resupinate. Sepals spreading, the dorsal free, the laterals
connate nearly to their apices. Petals smaller than the sepals, free. Lip calceiform,
saccate. Column short, terete. Fertile stamens 2, the third stamen (staminodium)
sterile. Stigma wider than the staminodium, papillose, trisulcate. Capsule fusi-
form, trilocular, crowned by the marcescent perianth.
One species in Panama.
1. SELENIPEDIUM CHICA Reichb. f. Xenia Orch, 1:3, Ё. 2. 1854; Pfitz. in Engl.
Pflanzenr. IV. 50 (Heft 12): 27. 1903; Ames, Sched. Orch. 2:1, Ё. І. 1923.
Stems leafy, erect, slender, up to about 5 га. tall. Leaves when mature 15-30
cm. long and 1.5-5 cm. broad, elliptic-lanceolate to lanceolate, acuminate, sub-
membranaceous, many-nerved, inarticulate, sparsely pubescent dorsally, especially
along the nerves, or becoming glabrate, contracted at the base into a cylindric
sheath covering the stem. Inflorescence terminal, many-flowered, densely pubes-
cent; flowers small to medium-sized, opening in succession, soon deciduous if
not fertilized; bracts normally 10-15 mm. long, lanceolate, acute, puberulent.
Dorsal sepal about 2 cm. long and 1 cm. broad, elliptic-oval, acute, puberulent
dorsally toward the base. Lateral sepals together about 2 cm. long and 1 cm.
broad, elliptic-oval, acute, connate nearly to their apices. Petals up to 2 cm.
long and about 2 mm. broad, linear, puberulent dorsally along the mid-nerve and
within near the base. Labellum 2-3 cm. long and 1-1.8 cm. broad in natural
position, calceiform, margins strongly involute.
Endemic in Panama.
“PANAMA”: without data, eo PANAMA: in foothills 20 miles northeast of
Panama cn alt. 60-90 m., Powell 258; northeast of Panama City, Powell 3007. CANAL
ZONE: Ancon Hill, Killip pi Cañon of Río Chagres, alt. 70-80 m., Sfeyermark &
Allen 17506.
This is undoubtedly the largest terrestrial orchid in the Americas.
2. PHRAGMIPEDIUM (Pfitz.) Rolfe
PHRAGMIPEDIUM (Pfitz.) Rolfe іп Orch. Rev. 4:331. 1896; Pfitz. in Engl.
7 Pflanzenr. IV. 50 (Heft 12): 42. 1903, as Phragmopedilum.
Papbiopedilum 8 Phragmopedilum Pfitz. in Engl. Bot. Jahrb. 25:527. 1898.
(114)
Fig. 64. Selenipedium chica
(115)
[Vor. 33
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Terrestrial or epiphytic herbs with very short stems. Leaves duplicative,
coriaceous, sulcate above and carinate below, inarticulate. Inflorescence terminal,
racemose or paniculate, several-flowered; flowers medium to large-sized, resupinate.
Sepals spreading, the dorsal free, the laterals connate almost to their apices. Petals
narrower than the sepals but often much elongated and caudate. Lip calceiform,
saccate. Column short. Fertile anthers 2, the third stamen (staminodium)
sterile, stigma deflexed from the apex of the column, subparallel to the ovary or
subrotund, papillose. Capsule long-fusiform, trilocular, the perianth deciduous.
Two species in Panama and about a dozen species in South America. Often
in cultivation.
a. Petals mostly more than 30 cm. long when mature; leaves lorate.......... 1. P. CAUDATUM
. Petals mostly less than 10 cm. long when mature; leaves linear or
linear-ligulate 2. P. LONGIFOLIUM
ы
a
PHRAGMIPEDIUM CAUDATUM (Lindl.) Rolfe in Orch. Rev. 4:332. 1896; Pfitz.
in Engl. Pflanzenr. IV. 50 (Heft 12): 52. 1903, in synon; L. Wms. in Ann:
Missouri Bot. Gard. 26:279. 1939.
Cypripedium caudatum Lindl. Gen . & Sp. Orch. РІ. 531.
—
.
b. f. 1852.
уе caudatum Pfitz. in Engl. Bot. Jahrb. 19: 41. 1894; Kerch. Orch. 454.
ndn caudatum var. Warscewiczii Hort. in Kerch. loc. cit. 1894.
Cypripedium caudatum var. roseum Hort. in Kerch. loc. cit. 1894.
Pbragmopedilum оаа Schltr. in Fedde Rep. Sp. Nov. Beih. 17:9. 1922.
Stems very much reduced, leafy. Leaves 15-20 cm. long, 3-5 cm. broad,
lorate, obtuse, apex shallowly and unequally bilobed, coriaceous, inarticulate,
glabrous. Inflorescence terminal, few-flowered, exceeding the leaves; rachis
velutinous; flowers large; bracts 3-6 cm. long, obtuse, subrotund, clasping the
rachis. Dorsal sepal 10-15 cm. long and 1.5-2 cm. broad at the base, lanceolate,
minutely pubescent dorsally. Lateral sepals together about 8-11 cm. long and
2.5-4 cm. broad, ovate-lanceolate to lanceolate, connate to their apices. Petals
30-70 cm. long when mature, 0.5—1 cm. broad at the base, long-caudate, puberu-
lent. Lip 5-6 cm. long and about 2.5-3 cm. broad in natural position, calceiform,
margins involute, lateral edges of the opening pilose. Ovary velutinous.
Panama, Venezuela (?), Colombia, Ecuador and Peru.
CHIRIQUÍ: probably Volcán de Chiriqui, Warscewicz; vicinity of Casita Alta, Volcan
de Chiriquí, alt. 1500—2000 m., Woodson, Allen & Seibert 062.
The description is based on the specimens collected by Woodson, Allen and
Seibert. Мо living material was available, and the plant is difficult to describe
from dried material. The form of the species in Panama is the superior horti-
cultural form and for this reason has been almost exterminated there. The species
is dificult to grow at sea-level.
2. PHRAGMIPEDIUM LONGIFOLIUM (Warsz. & Reichb. f.) Rolfe in Orch. Rev.
4:33. 1896.
(116)
1946] А
FLORA ОЕ PANAMA (Orchidaceae) 11
рне и» Warsz. & Reichb. f. in Bot. Zeit. 10:690. 1852; Hook. f. in
. Mag 5070, 18
бзен VAR dh Reichb. +. Xenia Orch, 1:3. 1054.
Cypripedium Hincksianum Reichb. f. in Gard. Chron. n. s. 9:202.
Paphiopedilum longifolium Pfitz. in Engl. & Prantl, Nat. Pflanzenfam. ^ 6:84. 1888.
Papbiopedium longifolium Kerch. Orch. 455.
Papbiopedilum Hincksianum Pfitz. in Engl. Bot. тік, 19:41. 1894.
Stems much reduced, leafy. Leaves up to 80 cm. long and 1-4 cm. broad,
linear or linear-ligulate, canaliculate, acute, inconspicuously and unequally bilobed
at the apex, inarticulate, distichous. Inflorescence terminal, several-flowered,
glabrous; flowers medium-sized; bracts 4-9 cm. long and 1-2 cm. broad, acute,
distichous, glabrous. Dorsal вера! 3-5.5 cm. long and 1–1.8 cm. broad, lanceolate
or lanceolate-acuminate. Lateral sepals 2.5-5 cm. long and 1.7-2.5 cm. broad,
ovate-lanceolate. Petals 5-10 cm. long and 0.4-0.8 cm. broad, linear-lanceolate,
caudate, pubescent within, especially at the base. Lip 2.5-4 cm. long and about
half as broad, calceiform, pubescent within, at least at the base, margins involute.
Costa Rica, Panama and Colombia.
CHIRIQUÍ: in Jom west of Caldera River, alt. 3800 feet, Powell 308; without
locality, Wars cocLÉ: valley of the upper Río Mata Ahogado, alt. 350 m., Allen
145; Las залдың | Bouché 4. PANAMA: without locality, Seemann 1114.
3. HABENARIA Willd.
Hapenaria Willd. Sp. Pl. 4:44. 1805; Kranzl. Orch. Gen. & Sp. 1:174. 1901;
Ames, Orch. 4:1-288. 1910.
Gymnadenia R. Br. in = Hort. Kew. ed. 2, 5:191. 1813.
Hina na L. C. Rich. in Mém. Mus. Par. 4:48. 1818.
Limnorchis Rydb. in Mem. N. Y. Bot. Gard. 1:104. 1900.
Terrestrial or palustrine herbs with cauline or basal leaves or rarely the leaves
reduced to bracts. Leaves thin to fleshy, usually prominent. Inflorescence few-
to many-flowered, spicate or racemose; flowers various; bracts often large and
conspicuous. Sepals subequal, free or connate at the bases, the dorsal erect,
usually concave, the laterals usually spreading. Petals similar to the sepals but
usually smaller, or more or less deeply bifid (rarely trifid or polyfid). Lip simple or
3 or more-lobed, often adnate to the column, spreading or pendulous, spurred at
the base. Column short, footless. Stigmas 2 or 1 stigma 2-lobed, often elongated
into short or comparatively long processes. Anther erect, rarely more or less
resupinate, persistent; pollinia (or pollen) granular, with short or sometimes long
caudicles from the base.
A polymorphic genus of world-wide distribution, probably one of the oldest
genera in the family. There are many more generic synonyms than those cited
but the ones given are the essential ones for the Panama flora. А good account
of the North American species known to 1910 will be found in Ames, Orchidaceae,
Vol. 4.
(117)
12
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
a. Petals entire or merely toothed; lip entire, toothed at the base or 3-
lobe
b. Petals lanceolate
bb. Petals not lanceolate
aa. Petals bifid; lip 3-lobed.
Palustrine or aquatic plants
1. H. ALATA
H. PETALODES
..8. H. REPENS
bb. Not palustrine nor aquatic plants.
c. Sepals less than 5 mm
cc. Sepals much more than
d.
etals
5 mm. long
=. Dn of the lip very ман shorter than the mid-lobe;
4
. long (mostly 3 mm.)
7. H. HEPTADACTYLA
m
. AVICULA
dd иш pem of the lip about as long as the mid-lobe or longer;
etals mal.
Fig. 65.
Habenaria alata
e. Bracts of the inflorescence large and
leaf-like; flowers mostly less than 5... 6. H. PAUCIFLORA
ee. Bracts of the inflorescence not large
and leaf-like; flowers mostly more
than 5
2 "n 40—50 ram. long; leaves en-
siform to linear-lanc . H. BICORNIS
ff. Spur m. long; leaves
те to ovate-lanceolate. 2. H. MONORRHIZA
1. HABENARIA АГАТА Hook. Exot. Fl. 3:7. 160.
1827; Ames, Orch. 4:273. 1910; Bot. Mus. Leafl.
Harv. Univ. 3:20, fig. 1934.
Erect or ascending terrestrial herbs with leafy
stems, 2-7 dm. tall Leaves 6-14 cm. long and
0.8-2 cm. broad, linear-lanceolate to lanceolate,
acute or acuminate, reduced to bracts below and
above. Inflorescence subracemose, up to 2 dm. long,
densely flowered; flowers pale green. Dorsal sepal
6-10 mm. long and 5-7 mm. broad, ovate to sub-
orbicular, concave. Lateral sepals 7-10 mm. long
and 3.5—4.5 mm. broad, lanceolate to triangular-
lanceolate, acute or acuminate. Petals 5.5-10 mm.
long and 2-2.5 mm. broad, lanceolate, acute or
obtuse, auriculate at the base on the anterior margin.
Lip 5.5-8 mm. long and about 2 mm. broad, linear-
lanceolate to lanceolate, entire, auriculate or dentate
on either side at the base; spur up to 13 mm. long,
recurved, subclavate. Ovary prominently winged
when mature.
Neotropical.
CANAL ZONE: Barro Colorado Island, Kenoyer 240;
Ana Lago and Corozal, Powell 316, 317, 318, 321, 3403.
The Panamanian material is somewhat atypical.
(118)
19461
FLORA ОЕ PANAMA (Orchidaceae) 13
2, HABENARIA MONORRHIZA (Sw.) Reichb. f. in Ber. Deut. Bot. Ges. 3:274. 1885.
Orchis monorrhiza Sw. Nov. Gen. & Sp. Prodr. 118. 1788.
Erect, slender, terrestrial herbs up to about 12 dm. tall. Leaves 3-13 cm. long
and 1.5—4 cm. broad, elliptic-oblong to lanceolate to ovate-lanceolate, acute or
acuminate, margins chartaceous, reduced to bracts upward. Inflorescence a dense-
ly flowered raceme up to about 20 cm. long; flowers white. Dorsal вера! 5-8 mm.
long and 4-5.5 mm. broad, ovate or suborbicular-ovate, obtuse, cucullate. Lateral
sepals 4-8 mm. long and 3-4 mm. broad, ovate, oblique, obtuse. Petals 5-8 mm.
long, bipartite, the lobes subequal; anterior lobe filiform; posterior lobe elliptic or
elliptic-oblong, obtuse or acutish. Lip about 7-10 mm. long, trilobate; lateral
lobes filiform, spreading; mid-lobe linear-ligulate; spur 15-25 mm. long, pendu-
lous, slender. Ovary somewhat winged.
Guatemala to Panama, the West Indies and South America.
Quí: Boquete, alt. 1150 m., Terry & Terry 1641. сосі.Е: hills south of El Valle
de Antón, alt. 600-800 m., Allen 2
3. HABENARIA PETALODES Lindl. Gen. & Sp. Orch. Pl. 316. 1835.
Habenaria petalodes var. micrantha Reichb. f. Beitr. Orch. Centr. ES 5. 1866.
Habenaria Warszewiczii Schltr. in Апех. Mem. Inst. Butantan 17:25. 1921.
Erect terrestrial herbs up to about 6 dm. tall. Leaves 4-17 cm. long and 1-5.5
cm. broad, lanceolate to elliptic or elliptic-oblong, acute or somewhat acuminate,
crowded near the middle of the stem, reduced to bracts above. Inflorescence up
to about 25 cm. long, strict, densely flowered; bracts up to 3 cm. long, lanceolate
to lanceolate-ovate, acute or acuminate. Dorsal вера! 6-10 mm. long and 4-6
mm. broad, ovate to broadly ovate, obtuse, cucullate. Lateral sepals 7-10 mm.
long and 3.5—5 mm. broad, semi-lunate or oblong-arcuate, obtuse. Petals 5-9
mm. long and 3.5—6.5 mm. broad, oblong to subquadrate to triangular-oblong
and cuneate at the base, dilated toward the truncate, bi- or tridentate apex. Lip
7-12 mm. long and about 1 mm. broad, linear, acute or obtuse, sometimes dilated
at the apex and sometimes with obscure teeth on either side at the base; spur up
to 30 mm. long (mostly about 20 mm.), slender, straight.
os ей Brazil and ES
Nata, alt. about 50 m., Allen 820. PANAMA:
Tubos ped ЕТІ pde py pe. same y Dee on alt. 0-350 m., Allen 1280. cANAL
ZONE and PANAMA: Chorerra, Tapia, Sieba Terra, Sosa Hill, ообо Hill, Juan Diaz
Range, Powell 212, 26о, 261, 262, 264, 265, 266, 326, 327, 328, 330, 331, 332, 336,
337» 339» 346, 3417, 3419, 3421.
I have reported a specimen of this, in error, as H. strictissima var. odontopetala
(Reichb. f.) L. Wms., in Ann. Missouri Bot. Gard. 27:271. 1940.
4. HABENARIA AVICULA Schltr. in Fedde Rep. Sp. Nov. Beih. 17:138. 1921.
Habenaria patentiloba Ames in Proc. Biol. Soc. Wash. 34:151. 1921.
Slender, erect, terrestrial herbs up to 4.5 dm. tall, with leafy stems. Leaves
7-15 cm. long and 1—3.5 cm. broad, lanceolate, elliptic to elliptic-oblong, obtuse,
(119)
[ VoL. 33
14 ANNALS OF THE MISSOURI BOTANICAL GARDEN
acute or acuminate, reduced to bracts above and below. Inflorescence laxly to
densely flowered; bracts 1-2.5 cm. long and 0.4-0.8 cm. broad, lanceolate to
ovate-lanceolate, acute or acuminate. Dorsal вера! about 4-6 mm. long and as
broad, suborbicular, concave. Lateral sepals 6-10 mm. long and 4-7.5 mm.
broad, broadly sub-lunate, spreading. Petals 4-6 mm. long and about 3 mm.
broad, bifurcate, the lobules or teeth acute, the anterior tooth shortest. Lip 12-16
mm. long and 1-2 mm. broad, linear, provided with a pair of narrow basal teeth
about 1-3.5 mm. long; spur 15-30 mm. long, slightly recurved.
Endemic in Panama.
"PANAMA": Joseph; Purdom. PANAMA: Orange River Valley, Killip 3124. CANAL
ZONE: near feit Kobe, Allen 2023; hills near Summit, Allen 2251; foothill savannahs
east of city, Powell 162.
5. HABENARIA BICORNIS Lindl. Gen. & Sp. Orch. Pl. 309. 1835.
Erect terrestrial herbs up to 6.5 dm. tall, with leafy stems. Leaves 5-25 cm.
long and 0.5-1.5 cm. broad, ensiform to linear-lanceolate, acute or acuminate.
Inflorescence a many-flowered raceme; bracts 1-3 cm. long, lanceolate, acuminate.
Dorsal sepal 6-8 mm. long and 4-6 mm. broad, ovate-orbicular, cucullate, obtuse
or acute, apiculate. Lateral sepals 8-10 mm. long and 4-5 mm. broad, semi-lunate,
acute, apiculate. Petals bifid; posterior lobe 5-6 mm. long and 1.5—2 mm. broad,
acute, arcuate; anterior lobe 6-9 mm. long and about 1 mm. broad, linear-
filiform, acute, arcuate. Lip tripartite; the mid-lobe 7-10 mm. long and 1-2
mm. broad; lateral lobes 9-12 mm. long and 1-1.5 mm. broad, arcuate-spreading,
acute; spur 40-50 mm. long, slender, subclavate.
Panama, the West Indies.
PANAMA: near Pacora, Powell 315. CANAL ZONE: Las Sabanas, Pittier 6792.
6. HABENARIA PAUCIFLORA (Lindl.) Reichb. f. in Bonplandia 2:10. 1854; L.
W ms. in Ann. Missouri Bot. Gard. 26:280. 1939.
Bonatea pauciflora Lindl. Gen. & Sp. Orch. Pl. 329. 1835.
Habenaria setifera Lindl. in Ann. Nat. Hist. 4:381. E 252 Orch. 4:209, 1. бо.
1910; in Bot. Mus. Leafl. Harv: Univ. 3:35, fig. 1, p. 2 934.
Habenaria паа Rich. & Gal. in Ann. Sci. Nat. III, 3: A pin
Erect terrestrial herbs up to 5.5 dm. tall. Leaves 4-12 cm. long and 0.3—1.8
cm. broad, linear-lanceolate to elliptic-lanceolate, acute or acuminate, sometimes
much reduced, usually partly sheathing the stem, reduced to bracts above.
Inflorescence few-flowered (rarely more than 5-flowered); bracts similar to the
leaves but smaller. Dorsal вера! 9-12 mm. long and 5-8 mm. broad, ovate-
lanceolate to ovate, apiculate, cucullate. Lateral sepals 8-12 mm. long and 3.5-5
mm. broad, triangular-lanceolate, subfalcate, acute, acuminate or apiculate. Petals
bipartite; anterior lobe 6—10 mm. long and up to 1 mm. broad, subfiliform, some-
what arcuate; posterior lobe 8—10 mm. long and 3—4 mm. broad, lanceolate or
elliptic-lanceolate, acute, somewhat arcuate. Lip tripartite; the lateral lobes
divergent and subequal to the mid-lobe in length; mid-lobe 10-14 mm. long and
up to 2 mm. broad; spur 2.5—6 cm. long, subclavellate.
(120)
19461
FLORA OF PANAMA (Orchidaceae)
Fig. 66. Habenaria repens
(121)
15
[Vor. 33
16 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico to Argentina.
CHIRIQUÍ: Llanos del Volcán, Seibert 326. PANAMA: Campana to Chica, Allen 2653;
Pacora to Chepo, Woodson, Allen & Seibert 1665.
An extremely variable species which has twenty or more synonyms.
7. HABENARIA HEPTADACTYLA Reichb. f. іг Linnaea 22:812. 1849.
Slender, erect, terrestrial herbs up to 3.5 dm. tall. Leaves up to 7 cm. long,
linear-filiform, aristate, reduced to bracts above. Inflorescence short, several-
flowered, more or less secund; bracts up to 1.5 cm. long, ovate-lanceolate, long-
acuminate; flowers small for the genus. Dorsal sepal 3—3.5 mm. long and 2-2.5
mm. broad, ovate, obtuse, cochleate, 3-nerved. Lateral sepals 3.5-4 mm. long
and 1.2-1.5 mm. broad, lanceolate, acute, 3-nerved. Petals 2.5-3 mm. long,
bipartite; the segments subequal or the outer shorter, lanceolate, acute, somewhat
arcuate. Lip 3-lobed; the mid-lobe linear, 3-4.5 mm. long and about 0.5 mm.
broad, linear, obtuse; lateral lobes 2.5—3.5 mm. long and 0.4—0.6 mm. broad
linear, obtuse, spreading; spur 10—12 mm. long, recurved, slightly saccate at the
tip.
Panama, Colombia, Venezuela, British Guiana and Brazil.
опт LOCALITY: Bouché 7. PANAMA: near Arraiján, alt. about 15 m., Woodson,
Allen 4 Seibert 1406.
8. HABENARIA REPENS Nutt. Gen. М. Am. Pl. 2:190. 1818; Ames, Orch. 4:216.
1910; in Bot. Mus. Leafl. Harv. Univ. 3:35, fig. p. 23. 1924.
Erect, slender, palustrine or aquatic herbs up to about 1 m. tall. Leaves up to
20 cm. long and 0.5-2.3 cm. broad, linear-elliptic to elliptic-lanceolate, acute or
acuminate, reduced to bracts upward. Inflorescence a densely flowered raceme;
flowers green. Dorsal sepals 4-7.5 mm. long and 3-5 mm. broad, oval to sub-
orbicular-ovate, cochleate, apiculate. Lateral sepals 4-8 mm. long and 2-3 mm.
broad, ovate to lanceolate-oval, oblique, apiculate. Petals 4-9 mm. long, bipartite,
the lobes subequal; anterior lobe filiform; posterior lobe 1-2 mm. broad, linear-
lanceolate to lanceolate, arcuate. Lip 5-10 mm. long and 1-2.5 mm. broad across
the base, trilobate, the lobes subparallel; lateral lobes filiform; mid-lobe linear-
lanceolate, usually a little shorter than the laterals; spur 8-15 mm. long, slender.
Southeastern United States, Guatemala, British Honduras, Nicaragua, Costa
Rica, Panama, the West Indies and South America.
CANAL ZONE: Barro Colorado Island, Bailey & Bailey 663; Dodge 3483; Woodworth
8 Vestal 55A, 65A.
Often found in floating debris and in very wet places. The illustration, fig. 66,
is redrawn from Ames, Orch. 1:51, 7. 16. 1905.
(122)
19461
FLORA OF PANAMA (Orchidaceae) 17
4. POGONIA Juss.
Росомтл Juss. Gen. 65. 1789.
Triphora Nutt. Gen. N. Am. Pl. 2:192. 1818.
Cleistes L. С. Rich. in Мет. Mus. Par. 4:31. 1818.
Rhizome or tuber-bearing, leafy or leafless terrestrial herbs. Sepals subequal,
free, erect or rarely spreading. Petals similar to the sepals or shorter and broader,
usually erect. Lip erect, simple or 3-lobed, sessile or unguiculate, plane or with
various lamellate or thickened nerves, spurless. Column elongated, wingless, foot-
less; stigma oblong or broader; clinandrium more or less raised, entire or denticu-
late; anther subincumbent or apparently erect, substipitate (at least in some
species) ; pollinia 2 masses, granular.
Ames (Orch. 7:7. 1922), in his account of this group of allied genera, sep-
arates Cleistes (Pogonia rosea) from Pogonia (typified by P. ophioglossoides) on
the basis of simple pollen grains for Pogonia and compound pollen grains for
Cleistes. However the so-called Cleistes seem to have simple pollen grains as
often as not. The case for maintaining Triphora as a distinct genus is much better
and is based on better characters. It is a matter of choice whether or not it is
maintained.
a. Lip with two stipitate processes at the base; lamina simp 1. P. ROSEA
aa. Lip without stipitate processes at n base; lamina 3- E.
b. Apex of the lip, at least, verruc 4. P. CUBENSIS
bb. Apex of the lip, nor any part, verrucose
. Cauline leaves well develope 2. P. MEXICANA
cc. Cauline leaves none or very much reduced 3. P. WAGNERI
1. Росоміл rosea (Lindl) Hemsl. in Godm. & Salvin, Biol. Centr.-Am. Bot.
3:304.
Cleistes rosea Lindl. Gen. & Sp. Orch. Pl. 410. 1840.
Tall, slender, erect, terrestrial herbs up to 1.5 m. tall (Panamanian material
up to 0.7 m. tall). Leaves 3-12 cm. long, 0.8—2.5 cm. broad, lanceolate, obtuse
or acute, sessile and clasping the stem, apparently rather thick when fresh.
Inflorescence of 1 or 2 rather large and showy flowers in the axils of the upper
leaves. Dorsal sepal 5-6.5 cm. long and 0.6-0.8 cm. broad, linear-elliptic to
lanceolate-elliptic, acute. Lateral sepals 5.5-6.5 cm. long and 0.8—1.1 cm. broad,
linear-lanceolate to lanceolate, acute. Petals 5—6.5 cm. long and 1—1.6 cm. broad,
elliptic to elliptic-oval, acuminate. Lip 4.5-5.5 cm. long and 1.6—3 cm. broad,
roadly oblanceolate to narrowly obovate, obtuse, the median line of the lip
callus-thickened or lamellate, lacerate toward the apex, with two stipitate pro-
cesses at the base.
Panama, Colombia, Venezuela, British Guiana, possibly also in Costa Rica, and
doubtless in other of the South American countries,—the “West Indies" according
to Hemsley.
COCLÉ: Seibert 504. CANAL ZONE: Powell 113, 3305, 3400. "Panama," Duchas-
saing.
(1239
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Fig. 67. Pogonia rosea
(124)
[Vor. 33
19461
FLORA OF PANAMA (Orchidaceae) 19
Mr. Powell reported a white-flowered form of the species.
2. PoGonta MEXICANA S. Wats. in Proc. Am. Acad. 26:154. 1891.
Tripbora mexicana ie fso Fedde Rep. Spec. Nov. Beih. 17:139. 1921; Ames, Orch.
7:40, 1. 100, f. 5-6.
Weak terrestrial herbs up to 27 cm. tall, with leaf-bearing stems. Leaves
8-20 mm. long and 5-17 mm. broad, narrowly to broadly ovate, obtuse or acute.
Inflorescence of 1-3 single flowers borne іп the axils of the upper leaves (or leaf-
like bracts?). Dorsal sepals 10-17 mm. long and about 2 mm. broad, linear to
linear-oblanceolate, obtuse or acute. Lateral sepals 10-17 mm. long and 1.5-3
mm. broad, linear to narrowly elliptic, subfalcate, acute. Petals 9-17 mm. long
and 1-3 mm. broad, linear to linear-elliptic, subfalcate, acute or obtuse. Lip
10-16 mm. long and 5-9 mm. broad, narrowly oval to obovate in outline, con-
spicuously narrowed to the base, 3-lobed; lateral lobes obtuse, subtriangular,
terminal lobe suborbicular to deltoid; disc trilamellate.
Mexico, Guatemala and Panama.
CANAL ZONE: Bevins.
Very close if not the same as Pogonia trianthophora (Sw.) BSP.
3. Poconta Wagneri (Schltr.) L. Wms., comb. nov.
Tripbora Wagneri Schltr. in Fedde Rep. Sp. Nov. 17:139. 1921.
Small, leafless, erect, terrestrial herbs up to 7.5 cm. tall. Inflorescence 1-
flowered, the flower erect. Dorsal вера! about 12 mm. long, ligulate, obtuse.
Lateral sepals about 12 mm. long, ligulate, obtuse, falcate. Petals a little shorter
and a little broader than the dorsal sepal, ligulate, obtuse, subfalcate-oblique. Lip
about 10 mm. long and 5 mm. broad, narrowly obovate-cuneate in outline, promi-
nently 3-lobed near the apex; lateral lobes obliquely ovate, obtuse; mid-lobe semi-
ovate, obtuse, somewhat undulate; disc with 3 thickened, subverrucose nerves
toward the apex.
Known only from Panama.
CHIRIQUÍ and vERAGUAs: Wagner 1778.
The description is taken from the original and a manuscript analysis.
PoGONIA CUBENSIS Reichb. f. іп Nederl. Kruidk. Arch. 4:322. 1850, as Р.
тибепзіз by error.
Tripbora cubensis Ames, Sched. Orch. 7:35. 1924.
>
Small terrestrial herbs up to 15 cm. tall. Leaves few, reduced to sheathing
bracts, often much reduced above, 0.8-2 cm. long and 0.4-1 cm. broad, lanceolate
to suborbicular. Inflorescence several-flowered, the pedicels of the lower flowers
often elongated and hence the raceme often appearing subcorymbose. Dorsal
вера! 7-8 mm. long and 1-1.5 mm. broad, linear to linear-lanceolate, obtuse.
Lateral sepals similar to the dorsal вера! except more or less arcuate. Petals 7-7.5
(125)
[Vor. 33
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mm. long and 1-1.2 mm. broad, narrowly elliptic to narrowly elliptic-oblanceo-
late, acute or obtuse. Lip 7-8 mm. long and about 3 mm. broad, lanceolate-
obovate, unguiculate, 3-lobed, mid-nerve more or less lamellate and often dentate;
mid-lobe subrotund; lateral lobes usually short, lanceolate to lanceolate-ovate.
Florida, Mexico, Panama and Cuba.
CANAL ZONE: Barro Colorado Island, Kenoyer 250.
This species has the appearance of a saprophyte but probably is not. It has
been collected but once in Panama.
5. VANILLA Sw.
VANILLA Sw. in Nova Acta Soc. Sci. Upsal. 6:66. 1799; Rolfe in Jour. Linn.
Soc. Bot. 32:439-478. 1896.
Usually epiphytic, scandent, leafy (in Panama) or leafless herbs, often with
branched stems. Leaves coriaceous or fleshy, sessile or short-petiolate. Inflorescence
a (short) raceme from the axils of the leaves. Sepals subequal, spreading, free.
Petals similar to the sepals. Lip with its claw adnate to the column, often en-
folding the column. Column elongated, footless, wingless; anther incumbent
(rarely appearing erect); pollinia granular. Capsule (“bean”) elongated, fleshy,
often fragrant.
The only truly commercial members of the Orchidaceae belong to the genus
Vanilla. The extract of the Vanilla "bean" was perhaps the first flavoring used
by the American Indians. Natural Vanilla flavoring is now largely displaced
by the synthetic.
The taxonomy of Vanilla is most difficult because of the fugaceous flowers.
a. Lip with verrucose lines or papillae 1. V. PLANIFOLIA
aa. Lip without verrucose lines or papillae 2. V. POMPONA
1. VANILLA PLANIFOLIA Andr. Bot. Repos. 8: ¢. 538. 1808; Dillon іп Am. Orch.
Soc. Bull. 10:339, 7. тт. 1942.
Vanilla fragrans of authors.
Scandent branched leafy herbs. Leaves 8—23 cm. long and 2-8 cm. broad,
narrowly lanceolate to elliptic-oblong, acute or acuminate, fleshy. Raceme short,
5-7 cm. long, axillary; bracts 5-10 mm. long, oblong, obtuse or subacute. Sepals
4-7 cm. long and 1-1.5 cm. broad, linear to oblanceolate, obtuse or acute. Petals
similar to the sepals but smaller. Lip 4-5 cm. long and 1.5-3 cm. broad, with
longitudinal, verrucose lines or papillae and a tuft of pubescence on the disc.
Column about 3 cm. long, pubescent dorsally. Capsule (“bean”) up to about
20 cm. long and 1 cm. in diameter.
Mexico to Panama, a also in the West Indies. Cultivated throughout
the tropics of the wor
SAN BLAS: Perme, Cooper 287. CANAL ZONE: Fort Clayton and Corozal, Standley
20103. PANAMA: Río Tecümen, Standley 20353; near Panama, Standley 20731.
(126)
57-26-%;
65793 А
1946]
FLORA OF PANAMA (Orchidaceae)
Fig. 68. Vanilla planifolia
21
[Vor. 33
22 ANNALS OF THE MISSOURI BOTANICAL GARDEN
We think that Vanilla rubra (Lam.) Urban will probably prove to be the
correct name for this species.
This is the most commonly cultivated of the Vanillas. There are apparently
several forms of the species and considerable difference in the flavor of the Vanilla
extract in the various forms.
2. VANILLA POMPONA Scheide in Linnaea 4:573. 1829; Ames, Sched. Orch.
9%6,> 1925.
Vanilla Pompona Lindl. Gen. & Sp. Orch. Pl. 437. 1840. |
Vanilla Pittieri Schltr. in Fedde Rep. Sp. Nov. 3: 106. 1906.
Scandent, branched, leafy herbs. Leaves 10-30 cm. long and 3.5-9 cm.
broad, variable in shape, narrowly elliptic-oblong or elliptic-lanceolate to lanceo-
late-ovate, rarely obovate-lanceolate, often oblique, acute or obtuse, coriaceous.
Inflorescence up to about 16 cm. long, several-flowered, axillary; bracts about 1
cm. long, lanceolate to ovate, obtuse or acute, cucullate. Sepals 7-8.5 cm. long,
linear-elliptic to narrowly oblanceolate, obtuse or acute. Petals similar to the
sepals or a little smaller. Lip 7—8.5 cm. long and 2.5—4 cm. broad, obovate-
lanceolate to obovate, unguiculate, crenulate at least toward the apex, with a
tuft of hair in the center of the disc. Column up to about 6 cm. long. Capsule
("bean") about 15 cm. long, fusiform.
Mexico to Panama, the West Indies, Venezuela, Ecuador, Bolivia, British and
Dutch Guiana; possibly as a cultigen in some of these countries and possibly
elsewhere.
CANAL ZONE: Hunter & Allen 852; Maxon 6770; Powell 127, 412, 3507; Piper.
Vanilla Pompona is cultivated in some places. It is probably inferior to V.
planifolia for making extract of Vanilla
6. ELLEANTHUS Presl
ELLEANTHUus Presl, Rel. Haenk. 1:97. 1827; Benth. & Hook. Gen. Pl. 3:522.
1883.
Evelyna Poepp. & Endl. Nov. Gen. & Sp. Pl. 1:32. 1835.
Terrestrial or epiphytic herbs ordinarily with strongly nerved, plicate, cauline
leaves. Sepals subequal, free or the laterals somewhat connate at the base and
forming a mentum. Petals subequal to the sepals, usually narrower. Lip adnate
to the base of the column and usually enfolding it, subequal to or exceeding the
sepals, usually gibbous-concave at the base and constricted above the base, the
basal cavity containing two large, usually subovoid calluses. Column erect, foot-
less; anther operculate, more or less incumbent or erect, bilocular; pollinia 8,
subceraceous.
A rather natural genus which is highly developed in the Andes.
a. Inflorescence distichou
. Stems fasciculately nched 4. Е. AURANTIACUS
bb. Stems not "here и pow]
(128)
$
~
1946]
FLORA OF PANAMA (Orchidaceae) 23
c. Leaves 1.5 cm. or less long, oblong-lanceolate or broader, per-
sistent 7. Е. MUSCICOLA
сс. Leaves 1.5 cm. ог more long, elliptic-lanceolate or narrower,
eciduous.
d. Bracts of the inflorescence imbricated; leaves subfiliform to
inear 6. E. LINIFOLIUS
dd. € = the inflorescence not imbricated; leaves elliptic-
5. E. LAxus
aa. ieee not Ea. tic :
b. Stems fasciculately "um 4. E. AURANTIACUS
bb. Stems not fasciculately branc
c. Inflorescence capitate or subcapitate 1. E. CAPITATUS
cc. Inflorescence spicate or racemose.
. Lip trilobulate 3. E. TRILOBATUS
dd. Lip not trilobulate 2. E. HYMENOPHORUS
1. ELLEANTHUS CAPITATUS (P. & E.) Reichb. f. in E Ann. 6:475. 1862.
Evelyna capitata Poepp. & Endl. Nov. Gen. & Sp. Pl. 1:3
Bletia = R. Br. in Ait. Hort. Kew. ed. 2, 5:206. TR bas not basis of Elleanthus
capitatus
Epiphytic or terrestrial, reed-like herbs up to about 12 dm. tall. Stems slender,
erect, covered with leaf-sheaths or naked below. Leaves 5—25 cm. long (mostly
15-20 cm.) and 1-5.5 cm. broad, linear-lanceolate to ovate-lanceolate, acuminate,
plicate, reduced upward and becoming bract-like. Inflorescence a dense, many-
flowered, capitate or subcapitate raceme; outer bracts sterile, triangular-ovate,
acuminate, up to about 4 cm. long, inner bracts fertile, ovate-lanceolate, acute or
acuminate, shorter than the outer bracts; flowers covered with a glutinous material.
Dorsal sepal 8-12 mm. long and 2.5-4 mm. broad, oblong-lanceolate, acute,
slightly cucullate. Lateral sepals 8-12 mm. long and 3-4 mm. broad, elliptic-
oblong, acute, somewhat oblique. Petals 11-12 mm. long and 1-2 mm. broad,
linear, obtuse or acute. Lip 8-13 mm. long and 8-11 mm. broad, oval to sub-
orbicular-flabellate, anterior margins more or less lacerate-dentate, base gibbous
and with two approximate, subglobose calluses; disc thickened in the middle.
Mexico to Panama, the West Indies and in South America to Peru.
CHIRIQUÍ: Monte Lirio, alt. 1300-1900 m., Seibert 140, 268.
A distinctive and widespread species. The name is based on Evelyna capitata
Poepp. & Endl., and not on Bletia capitata В. Br., although the two names ap-
parently represent the same species.
2. ELLEANTHUS HYMENOPHORUS Reichb. f. in Walp. Ann. 6:480. 1862.
Evelyna hymenophora Reichb. f. in Bot. Zeit. 10:710. 1852.
Caespitose, epiphytic herbs up to about 6.5 dm. tall (usually much less).
Stems slender, covered with leaf-sheaths. Leaves 4-17 cm. long and 1.8-8 cm.
broad, elliptic, elliptic-ovate, ovate or oval, acuminate, pergameneous. Inflorescence
a compact many-flowered spike up to about 9 cm. long; bracts up to about 1.5
cm. long, lanceolate-ovate or ovate, acuminate. Dorsal sepal 5.5-7 mm. long and
abaut 2.5 mm. broad, lanceolate to oval, acute. Lateral sepals 5-7 mm. long and
2-3 mm. broad, lanceolate to oblong-lanceolate, acute, strongly keeled dorsally.
(129)
[Vor -35
24 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Petals 3.5-7 mm. long and 1-1.5 mm. broad, linear-spatulate to spatulate, obtuse
or acutish, more or less crenulate above. Lip 7-9 mm. long and 6.5-9 mm.
broad, broadly ovate to orbicular, lacerate-dentate, the gibbous-saccate base en-
closing 2 ovoid calluses and the sac with a bidentate lamellate callus in front.
Costa Rica, Panama, Colombia and Peru.
CHIRIQUÍ: "Chiriquí," Warscewicz. cocLÉ: summit of Valle Chiquita, alt. 1360 m.,
Bouché 3x; gcn north te El Valle de Antón, alt. 1000 m., Allen 2172; near Cerro Turega,
alt. 650-70 , Woodson & Schery 168. PANAMA: hills above Campana, alt. 600-800
m., Allen 1885.
3. ELLEANTHUS TRILOBATUS Ames & Schweinf. Sched. Orch. 8:53. 1925.
Slender epiphytic herbs up to about 8 dm. tall. Leaves 5-18 cm. long and
0.5-2.5 cm. broad, linear-lanceolate to lanceolate, acuminate, plicate, apex tri-
lobulate, reduced upwards, basal part of the stem in mature plants naked. In-
florescence up to 8 cm. long, many-flowered; bracts up to 2.5 cm. long, ovate-
lanceolate to ovate, acuminate, cucullate, chartaceous, reduced upward. Sepals
more or less pubescent dorsally. Dorsal sepal 6-8 mm. long and 2-3.5 mm.
broad, oblong-lanceolate to narrowly ovate-lanceolate, acute, apiculate. Lateral
sepals 6-8 mm. long and 2.5-3 mm. broad, ovate-lanceolate to oblong-lanceolate,
somewhat oblique, acute, cucullate. Petals 5-7 mm. long and 1-1.5 mm. broad,
linear-oblong, obtuse. Lip trilobulate, 6-7.5 mm. broad, obovate-orbicular, flabel-
late, fimbriate-lacerate on the terminal half, base gibbous-saccate and with 2 oval
calluses, disc with 2 transverse, sublamellate calluses about opposite each sinus.
Ovary puberulent.
Costa Rica, Panama and Colombia.
COLON: near Porto Bello, Maxon 5808. E vicinity of Campana, alt. about
1000 m., Allen 2405; ime east of Disi a] C an Juan, үлі 97, 3266, 3267,
3208, 3273, 3529. CANAL ZONE: Barro Colored id. Shattuck 20
The Powell specimens, regardless of numbers, are Ng all from one col-
lection. Schlechter has determined Maxon 5808 and Powell 97 as his Elleanthus
Brenesii and says of the latter collection. (Fedde, Rep. Sp. Nov. Beih. 17:13.
1922): "Die Pflanze stimmt recht gut mit meinem Originalexemplar aus Costa
Rica überein." The description and analysis of E. Brenesii are not entirely satis-
factory but when the type can be studied E. trilobatus will probably prove to be
а synonym.
4. ELLEANTHUS AURANTIACUS (Lindl) Reichb. f. in Walp. Ann. 6:482. 1863.
velyna aurantiaca Lindl. in Benth. РІ. age i 149. 1844.
dcinde Tonduzii Schltr. in Fedde Rep. Sp. Nov. 8:567. 1910.
Erect or ascending, branched or usually fasciculately branched herbs up to
about 1 m. or more tall. Stems slender, covered with sheaths but becoming naked
below, the lateral stems usually smaller than the main stem. Leaves 2-12 cm.
long and 0.2-1.4 cm. broad, linear-lanceolate to lanceolate, acuminate, apex 2-3-
lobulate and short-aristate, plicate. Inflorescence up to about 4 cm. long, bilateral;
1946]
ek га
FLORA OF PANAMA (Orchidaceae)
“Бір. 69. Elleanthus trilobatus
(131)
25
[Vor. 33
26 ANNALS OF THE MISSOURI BOTANICAL GARDEN
bracts up to about 1.5 cm. long, ovate-lanceolate or ovate, acuminate, cucullate,
scarious; rachis somewhat fractiflex. Sepals often puberulous dorsally. Dorsal
sepal 4.5-7 mm. long and 2-3.5 mm. broad, lanceolate to ovate-lanceolate, acute,
cucullate. Lateral sepals 6-8 mm. long and 3—4 mm. broad, lanceolate to lanceo-
late-triangular, acute, somewhat arcuate. Petals 5-8 mm. long and 2-2.5 mm.
broad, linear-oblong or oblanceolate, obtuse or acute, often somewhat undulate.
Lip 7-10 mm. long and 7-9 mm. broad, obovate-flabellate to suborbicular, lacerate-
dentate at least toward the apex, saccate-gibbous at the base and with 2 approxi-
mate calluses.
Costa Rica, Panama, and in South America to Peru.
CHIRIQUÍ: Bajo Chorro, alt. 2100 m., Davidson 123. PANAMA: hills above Campana,
July 1, 1939, Allen 1884.
Elleantbus aurantiacus is а widespread species with a distinctive mode of
branching. The description is based on the Central American material and typical
material from South America. Some specimens from South America are much
more robust and have larger leaves than typical.
5. ELLEANTHUS Laxus Schltr. in Fedde Rep. Sp. Nov. 12:213. 1913; in Fedde
Rep. Sp. Nov. Већ. 59: 7. 4, fig. 16. 1931.
Slender erect caespitose herbs up to about 4 dm. tall. Leaves 4-12 cm. long
and 0.4—1.2 cm. broad, elliptic-lanceolate, acuminate, 2—3-dentate at the apex,
plicate, mostly near the summit of the stems. Inflorescence up to 5 cm. long,
few-several-flowered, distichous, strongly fractiflex; bracts up to 4.5 cm. long,
reduced upward, lanceolate to ovate-lanceolate, acuminate, chartaceous, strongly
cucullate, longer than the flowers. Sepals ligulate, acuminate, 7 mm. long, the
laterals oblique. Petals oblique, linear, subacute, about equal to the sepals. Lip
concave-subcucullate at the base, dilated above and deeply emarginate, the margins
crenate-incised and undulate, 6 mm. long and 4 mm. broad above the middle,
base with 2 oblong calluses, the disc with a few papillae.
Costa Rica (?) and Panama.
DARIEN: Sambi Basin, alt. 500—974 m., Pittier 5640.
The description of the flowers is taken from the original publication. No
flowers from typical material have been seen but there is an analysis (by
Schlechter) in the Ames Herbarium. If the analysis is correct the Costa Rican
material, of which flowers are available, belongs to a different species. The Costa
Rican material called E. laxus has a lip which is essentially oblong, emarginate,
lacks surface papillae of any sort (certainly lacks the tooth-like “рарШае” of
Schlechter's analysis and the figure cited) ; the petals are differently shaped as are
the lateral sepals.
6. ELLEANTHUS LINIFOLIUS Presl, Rel. Haenk. 1:97. 1827.
Isochilus linifolium Lindl. Gen. & Sp. Orch. Pl. 113. 1831.
Densely caespitose epiphytic herbs up to about 3.5 dm. tall, but mostly up to
(132)
19461
FLORA OF PANAMA (Orchidaceae) 27
about 1.5 dm. tall in Panama. Stems slender, covered with the leaf-sheaths or
partially naked. Leaves 1.5-9 cm. long and 0.1-0.4 cm. broad, subfiliform to
linear, acute, obtuse or denticulate at the apex. Inflorescence up to 2.5 cm. long,
a short, distichous, obscurely fractiflex raceme; bracts up to about 1.5 cm. long,
lanceolate or ovate-lanceolate, acute or acuminate, cucullate, scarious, imbricated.
Sepals glabrous or sparsely brown-pubescent dorsally. Dorsal sepal about 3 mm.
long and 1 mm. broad, elliptic-lanceolate, cucullate. Lateral sepals 3-3.5 mm.
long and 1.2-1.5 mm. broad, ovate-lanceolate, short-acuminate, somewhat arcu-
ate. Petals about 3 mm. long and 1-1.2 mm. broad, linear-oblong to oblong-
spatulate, obtuse, truncate, obscurely crenulate, arcuate. Lip about 3-3.5 mm.
long and 4-5 mm. broad, subquadrate-flabellate to flabellate, truncate, denticulate,
gibbous at the base and with 2 lightly attached ovoid calluses, with a short trans-
verse callus above the base.
Mexico, British Honduras, Guatemala, Costa Rica, Panama, the West Indies
and south to Peru.
ON: Cativa-Porto Bello trail, Powell 353. состЕ: El Valle de Antón, alt. 600 m.,
Ай. pa hills north of El Valle de Antón, alt. 1000 m., Allen 2860. CANAL ZONE:
Gatun Lake, Purdom
7. ELLEANTHUS MuscIcoLa Schltr. in Fedde Rep. Sp. Nov. Вей. 19:10. 1923.
Small caespitose epiphytic herbs up to about 12 cm. tall. Stems slender,
covered with leaf-sheaths. Leaves 10-15 mm. long and 2.5-5 mm. broad, oblong-
lanceolate to oblong-ovate, obtuse, coriaceous, persistent, distichous. Inflorescence
up to about 1.7 cm. long, a short distichous, obscurely fractiflex raceme; bracts
up to about 1 cm. long, lanceolate-ovate or ovate, compressed, cucullate, scarious,
sphacelate, imbricated. Sepals substellate-furfuraceous dorsally; dorsal sepal about
3 mm. long and 1 mm. broad, elliptic, acute, cucullate; lateral sepals about 3
mm. long and 1.5 mm. broad, ovate-lanceolate, acuminate, cucullate. Petals
about 3.5 mm. long and 1.5 mm. broad, oblanceolate-subflabellate, obtuse, crenu-
late-undulate above. Lip about 3-3.5 mm. long and 3.5—4 mm. broad, flabellate-
triangular, truncate and denticulate.
Costa Rica and Panama.
COCLÉ: hills north of El Valle de Antón, alt. 1000 m., Allen 2150.
Elleanthus muscicola is unusual in the genus because of the habit and small
persistent leaves.
7. SOBRALIA Ruiz & Pavon
SoBRALIA Ruiz & Pavon, Fl. Peruv. & МЕ Ргодг. 120, #. 26. 1794.
Fregea Reichb. f. in Вог. Zeit
Lindsayella Ames & Тозығы іп A [^i e Harv. Univ. 5:33. 1935.
Large or small, reed-like, terrestrial or epiphytic herbs. Leaves pergameneous,
plicate-veined, sessile, usually few. Inflorescence a terminal raceme or óften re-
duced to 1 flower; flowers fugaceous in Panamanian species. Sepals equal or nearly
so, erect or somewhat spreading, connate at their bases. Petals similar to the sepals
(133)
[Vor. 33
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN
but usually broader. Lip erect, from the base of the column, usually covering
or enfolding the column, simple, retuse or 3-lobed, usually concave, a little longer
than the sepals and petals, often undulate or fimbriate; the disc smooth, lamellate
or lamellate-cristate, usually with inconspicuous or conspicuous callus thickenings
at the base. Column elongated, semi-terete, more or less arcuate, with narrow
wings (auricles) or acute angles at the apex, footless; anther incumbent; pollinia
8, granular or subceraceous.
A small tropical American genus, possibly with its center of distribution in
Panama. Difficult to work because of the fugaceous flowers.
Fregea, a genus described by Reichenbach, has been maintained as distinct but
has no characters which will separate it from Sobralia.—The original specimen was
collected in Chiriqui.
Lindsayella, a genus based on specimens from Panama, seems to have no char-
acters of generic value. The main segregation character is the callus at the base
of the lip. Rather than a generic character these calluses seem to be the culmina-
tion of a tendency within the genus Sobralia.
Our knowledge of this genus in Panama is entirely inadequate. A large num-
ber of the species presumed to grow there were described by Reichenbach from
material collected by Warscewicz, and of many of them nothing further is known.
Paul H. Allen, long a resident of Panama and the keenest collector of orchids who
has been in the country, is inclined to believe that many of the Warscewicz
Sobralias which were ascribed to Panama were actually collected elsewhere. Mr.
Allen writes that when Sobralias are present in the Panama flora they are apt to
be quite abundant; yet he has not been able to discover many of the species that
Warscewicz is said to have collected in Chiriquí Province although he has collected
the same area many times.
We have not been able to make a satisfactory key to this genus because many
of the species are practically unknown and require more study, as a whole, than is
possible now.
1. SoBRALIA ALLENI L. Wms. in Ann. Missouri Bot. Gard. 29:336, pl. 30, figs.
1-3. А
Caespitose, epiphytic plants up to about 5 dm. tall. Stems about 1-2 mm. іп
diameter, slender, bearing 1-3 leaves toward the apex, leafless below or the leaves
reduced to sheaths. Leaves 13-18.5 cm. long and 1.8-2.5 cm. broad, elliptic-
lanceolate to narrowly elliptic, acuminate, plicate, with seven principal nerves,
lepidote on the lower surface especially along the nerves at the base, glabrous above
or essentially so; leaf-sheaths closely appressed to the stem, lepidote. Inflorescence
terminal, 1-flowered; flowers small, white with a pale yellow lip. Dorsal sepal
about 3.5 cm. long and 7 mm. broad, oblanceolate, apiculate, 7-nerved. Lateral
sepals about 3.5 cm. long and 7 mm. broad, linear-oblong, apiculate, 7-nerved.
Petals about 3.5 cm. long and 6-7 mm. broad, oblanceolate, acute, terminal half
serrulate, 7-nerved. Lip about 3.5 cm. long and 1.8 cm. broad, oblong-oval,
truncate or shallowly retuse, terminal half lacerate-dentate; disc with several
(134)
19461
Fig. 70. Sobralia Allenii
(135)
[Vor. 33
30 ANNALS OF THE MISSOURI BOTANICAL GARDEN
inconspicuous longitudinal carinae and with a small bipartite callus thickening at
the base, pubescent longitudinally along the middle, especially toward the apex.
Panama.
cocLÉ: hills north of El Valle de Antón, alt. 1000 m., Allen 2686.
2. SoBRALIA amabilis (Reichb. f.) L. Wms., comb. nov.
Fregea amabilis Reichb. f. in Bot. Zeit. 10:712. 1852; Beitr. Orch. Centr.-Am. 10,
f. 2. 1866.
Epiphytic or terrestrial herbs up to 6 dm. tall. Leaves 3-10.5 cm. long and
1-3.5 cm. broad, elliptic to broadly lanceolate-elliptic, acuminate; leaf-sheaths
prominently verrucose. Inflorescence a single terminal flower; flower small for
the genus, purple. Sepals 23-27 mm. long and 7-8 mm. broad, elliptic to elliptic-
oblanceolate, acute, apiculate. Petals 20-25 mm. long and 8-10 mm. broad,
broadly elliptic to elliptic-obovate, obtuse ос acute. Lip 30-35 mm. long and
20—25 mm. broad, quadrate-obovate, emarginate, somewhat narrowed at the base
and enfolding the column; column 10-15 mm. long.
Costa Rica and Panama.
CHIRIQUÍ: An einer nassen und kalten Stelle, Cordilleren von Chiriqui, Warscewicz.
The original species of the genus Fregea, which we do not consider distinct
from Sobralia.
3. ЗбовклІлА Ві ЕТІЛЕ Reichb. f. in Bot. Zeit. 10:713. 1852.
Plants of unknown but probably large size. Leaves 10-17 cm. long and 3-4.5
cm. broad, elliptic to broadly lanceolate, acute. Lip about 4 cm. long and about
2 cm. broad, cuneate or narrowly obovate in outline, 3-lobed; lateral lobes
lanceolate-triangular, falcate; mid-lobe obovate-orbicular, emarginate, slightly
exceeding the lateral lobes; disc with five crisped lamellae.
Panama.
CHIRIQUÍ: Warscewicz.
We have seen no material of this species. The description is taken from the
original and from a photograph of the very poor type specimen.
4. SoBRALIA Boucuri Ames & Schweinf., Sched. Orch. 10:4. 1930.
Slender terrestrial or epiphytic herbs up to 7.5 dm. tall. Stems about 3-4
mm. in diameter. Leaves 8-17 cm. long and 3—4.5 mm. broad, elliptic or broadly
elliptic-lanceolate, acute, marginate. Sepals 5.5—6.5 cm. long and 1.5-1.7 cm.
broad, elliptic to lanceolate, the laterals somewhat arcuate. Petals about 6 cm.
long and 2.2 cm. broad, elliptic-oblanceolate, acute, somewhat oblique. Lip about
5.5 cm. long and 5 cm. broad, obovate-orbicular, trilobulate, margins crenulate-
dentate anteriorally; mid-lobe small, suborbicular; disc with 7 longitudinal,
crispate carinae. Column about 2.5 cm. long.
Panama.
сост: El Valle de Antón, alt. about 1000 m., Allen 2454. “PANAMA”: Bouché s. n.
Probably the original came from Coclé or Chiriqui. It is very closely allied
to S. Bletiae Reichb. f.
(136)
1946)
FLORA OF PANAMA (Orchidaceae) 31
5. Зовклла callosa L. Wms., nom. nov.
Lindsayella amabilis Ames & Yn in pe: Mus. Leafl. Harv. Univ. 5:33. 1937, non
Sobralia amabilis (Reichb. f.) L.
Slender, caespitose (?), epiphytic or terrestrial herbs up to about 4 dm. tall.
Leaves 4.5-10.5 cm. long and 3-8 mm. broad, narrowly elliptic to linear-
lanceolate, acute or obtuse, apex 2—3-dentate, prominently plicate; leaf-sheaths
concealing the stems, striate, glabrous. Inflorescence usually 1-flowered, from
the axis of the uppermost leaf. Sepals 20-25 mm. long and 7-8 mm. broad,
elliptic, acute, the laterals slightly arcuate, slightly connate at the base. Petals
2, 20-23 mm. long and 10-12 mm. broad, lanceolate to lanceolate-ovate, acute.
Lip about 30 mm. long and about 20 mm. broad toward the apex, triangular-
obovate in outline, emarginate, somewhat crenate, the lamina with 1 prominent,
crispate lamella and with a prominent oblong-obovoid callus at the base. Column
about 10-12 mm. long.
Panama.
coc ecimen originally collected in Coclé, cultivated in the Canal Zone Ex-
Me Gd. Lindsay 449.
6. SoBRALIA FENZLIANA Reichb. f. in Bot. Zeit. 10:714. 1852.
Slender terrestrial or epiphytic herbs. Leaves 13-17 cm. long and 4-6 cm.
broad, elliptic-oblong to lanceolate-ovate, acuminate. Sepals about 4 cm. long,
elliptic or oblong-elliptic, acute. Petals a little shorter than the sepals, narrowly
obovate, acute. Lip (4-6 cm. long?) obovate-orbicular, crenate-dentate anterior-
ally, ecarinate.
Panama.
CHIRIQUÍ: Warscewicz 48.
The description is taken from the original and an pow of the type. Seems
to differ from S. Bouchei only in lacking carinae on the lip
7. SoBRALIA FRAGRANS Lindl. in Gard. Chron. 598. 1853.
Sobralia eublepharis Reichb. f. ex Kránzl. іп Fedde Rep. Sp. Nov. 26:255, f. 78. 1929.
Relatively small, terrestrial or epiphytic herbs up to 35 cm. tall Stems
ancipitous, with the principal leaf near the middle. Leaves 6.5—23.5 cm. long and
1.5—5 cm. broad, narrowly elliptic to broadly lanceolate, acute or acuminate.
Sepals 3-4 cm. long and 0.4-0.7 cm. broad, linear to oblong-lanceolate, acute or
apiculate, the laterals somewhat falcate. Petals 3-4 cm. long and about 0.5 cm.
broad, lanceolate to oblanceolate-linear, obtuse or acute, sometimes apiculate. Lip
28-35 mm. long and 16-20 mm. broad, oval to obovate, the anterior margin
dentate-fimbriate, the posterior margins entire; disc with several thickened nerves
and 2 callus thickenings at the base, pubescent especially toward the apex. Column
about 16-18 mm. long.
British Honduras, Guatemala, Honduras, Costa Rica and Panama.
HiRIQUÍ: San Felix to Cerro Flor, alt. 100-800 m., Allen 1041. УЕКАСУА8: head-
waters of Rio Cafiazas, alt. 300—600 m., Allen 167. сост: between Las Margaritas and
(137)
[Vor. 33
32 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ЕІ Valle, Woodson, Allen & Seibert 1314. PANAMA: drowned forest near Vigia and San
Juan, Dodge, Steyermark & Allen 16521. CANAL ZONE: Powell 114, 3404, 3424.
Perhaps the most distinctive of the Panamanian species of the genus.
8. SOBRALIA LABIATA Warsz. & Reichb. f. in Bot. Zeit. 10:714. 1852.
A plant of unknown size. Leaves up to about 27 cm. long and 2 cm. broad,
linear-lanceolate, acuminate. Sepals about 6 cm. long, oblong-lanceolate, acute.
Petals about as long as the sepals and a little broader. Lip about 7-8 cm. long
and 5—6 cm. broad, flabeilate, anteriorally crenulate, retuse.
Panama.
CHIRIQUÍ: Warscewicz.
The description is taken from the original and a poor analysis of the type.
9. SOBRALIA LEUCOXANTHA Reichb. f. Beitr. Orch. Centr.-Am. 68. 1866; Hook.
f. in Bot. Mag. 115: /. 7058. 1889.
Sobralia Powellii Schltr. in Fedde Rep. Sp. Nov. Beih. 17:11. 1922.
Tall, slender, terrestrial or epiphytic herbs up to 1.5 m. tall. Leaves 8—22
cm. long and 3—7 cm. broad, elliptic to lanceolate, acuminate, glabrous above,
more or less scurfy below; leaf-sheaths verruculose or obscurely punctate. In-
florescence 1-flowered; the flowers large. Sepals and petals white, the lip white
with the throat yellow to orange. Sepals 5-8.5 cm. long and 1.5—2.5 cm. broad,
linear-lanceolate to oblong. Petals 5-7 cm. long and 2.5—3.5 cm. broad, oblong
to obovate. Lip 5—7 cm. long and up to 5 cm. broad, obovate, enfolding the
column, somewhat retuse, margin undulate, with short calluses at the base and
1-3 thickened nerves above. Column 3-4.6 cm. long.
Costa Rica and Panama.
COCLÉ: south of El Valle, alt. 600 m., Allen 2839, 2840; mountains beyond La
Pintada, alt. 400—600 m., Hunter & Allen 620. PANAMA: Peluca Hydrographic Station,
alt. about 90 m., Hunter 9 Allen 648; Gatün Lake, Powell 2, 3329, 3330, 3331.
There seems to be little or no difference between Sobralia leucoxantha and S.
Powellii.
10. SoBRALIA LINDLEYANA Reichb. f. in Bot. Zeit. 10:713. 1852; Xenia Orch.
2:156, Ё. 163. 1868.
Slender epiphytic or terrestrial herbs up to 6 dm. tall. Leaves 5.5—13 cm.
long and 2-5 cm. broad, elliptic-lanceolate to ovate-lanceolate, acuminate, often
vernicose above; sheaths neither pubescent nor verruculose. Inflorescence 1-
flowered; the flowers white or (with age?) yellowish, the lip with red maculations.
Sepals 4-5 cm. long and 0.8-1.2 cm. broad, elliptic-lanceolate to elliptic-oblong,
acute. Petals 3.5—4.5 cm. long and 1.2-1.4 cm. broad, similar to the sepals or
broader. Lip 4—5 cm. long and 3—4 cm. broad, cuneate-flabellate or obovate, sub-
trilobate, the lobes rounded, apical part of the lip dentate-lacerate or crisped;
disc with a tricornute callus at the base and pubescent longitudinally within.
Column about 2 cm. long, with 2 acinaciform, terminal stelidia or auricles.
(138)
1946)
FLORA OF PANAMA (Orchidaceae)
Fig. 71. Sobralia panamensis
(139)
33
[Vor. 33
34 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Costa Rica and Panama.
CHIRIQUÍ: Llanos del тш, 3i 1120-1200 m., Seibert 325; Chiriqui Viejo Valley,
W bite 35; "Chiriquí," Warsce
11. SOBRALIA PANAMENSIS Schltr. іп Fedde Rep. Sp. Nov. Beih. 17:11. 1922.
Sobralia panamensis var. albiflos Schltr. loc. cit.
Erect or ascending, epiphytic or terrestrial herbs; the stems up to about 15 dm.
long, simple, branching or producing new plants at the nodes, lepidote or fur-
furescent. Leaves 5-20 cm. long and 2-6.5 cm. broad, elliptic to broadly elliptic-
lanceolate, acuminate, glabrous or slightly furfurescent at the base dorsally;
leaf-sheaths densely pubescent, lepidote or furfurescent, older sheaths appearing
pustulate. Flowers purple to white. Sepals 4.5-6 cm. long and about 1.2-1.6
cm. broad, ligulate to lanceolate, acute, the laterals somewhat oblique. Petals a
little shorter than the sepals and up to about 2 cm. broad, obtuse or acute. Lip
4-6 cm. long and 3-4 cm. broad, obovate or oblong-obovate, obtuse, glabrous,
apex undulate, with 2 small calluses at the base. Column up to about 3 cm. long.
Panama.
26 e
CANAL в. Santa Rita nu Cowell 146; Quebrada Fata Dodge 8 Steyermark
106a, 1065; near Vigía and San Juan, Dodge et al. 16557; hills east of canal, Powell 21,
31, 3082, 3086, 3105; Barro Colorado Island, Woodworth & Vestal 388.
Sobralia panamensis is closely allied to S. decora Batem. and to S. sessilis Lindl.
but seems to be distinct.
12. SOBRALIA DECORA Batem. var. AERATA Allen & Williams in Ann. Missouri
Bot. Gard. 29:337, Ё. 31. 1942.
Slender, erect, epiphytic or terrestrial herbs up to about 5 or 6 dm. tall. Stems
covered with the amplexicaul leaf-sheaths, simple or rarely branched. Leaves
8-16 cm. long and 1.3-5 cm. broad, lanceolate or lanceolate-ovate, acuminate,
coriaceous. Inflorescence sessile, 1-flowered. Sepals 5.2—5.5 cm. long and 1.2-1.5
cm. broad, lanceolate, acute, greenish on the outer surface, very near "brick red"
on the inner surfaces. Petals about 4.7 cm. long and 1-1.2 cm. broad, elliptic-
lanceolate, acute, reflexed, washed "brick red" with lighter margins and dorsal
median line, the inner surface "brick red" with a white border. Lip about 4 cm.
long, more or less tubular but flaring at the apex, the disc "brick red" with a
median white stripe and the lateral margins white toward the base, base of the lip
“Acajou red" within.—Description from the type specimen and ample field notes.
The species is found from Mexico to Honduras. Тһе variety only in Panama.
cocLÉ: bad lands south of El Valle de Antón, alt. 500—600 m., Allen 2755, 2846.
13. SonRALIA ROLFEANA Schltr. іп Fedde Rep. Sp. Nov. Већ. 17:12. 1922.
Erect, slender, probably terrestrial herbs up to about 1 m. tall Leaves
6.5—21 cm. long and 2-7.5 cm. broad, oblong-elliptic to elliptic-oblanceolate or
elliptic-lanceolate, acute to acuminate, glabrous and smooth, mostly near the
(140)
1946)
35
FLORA OF PANAMA (Orchidaceae)
Sobralia decora var. aerata
Pig. 72.
Inflorescence probably consisting
reduced upward.
М
middle of the stem and above
yellow with a
-1.6 cm. broad
flowers creamy
of a single flower in the axis of an upper leaf;
ligulate to
Petals 5.5-6.5
,
Sepals 5.5—7.5 cm. long and 0.8
lemon-yellow throat.
elliptic or elliptic-lanceolate, acute, the laterals somewhat oblique.
(141)
[Vor. 33
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN
cm. long and 0.8—1.4 cm. broad, ligulate to oblong-ligulate, obtuse or acute. Lip
5,5-7 cm. long and 2.5—4.7 cm. broad, obovate, retuse, anterior margin lacerate-
dentate, obscurely bilamellate at the base, glabrous. Column 2-3 cm. long,
enfolding the lip.
Panama.
NAMA: Cerro Campana, alt. 800 m., Allen 2663. CANAL ZONE: without definite
Бену, Powell 3, 3327, 3357, Er 3374. BOCAS DEL Tomo: epiphyte with yellow
flowers, von Wedel 409.
Sobralia Rolfeana is very closely allied to S. leucoxantba Reichb. f. and to S.
Lindleyana Reichb. f.
14. SOBRALIA SUAVEOLENS Reichb. f. in Gard. Chron. n. s. 9:622. 1878.
Sobralia epiphytica Schltr. in Fedde Rep. Sp. Nov. 12:213. 1913.
Strict terrestrial or epiphytic
herbs up to about 4 dm. tall. Leaves
5-19 cm. long and 2-4.5 cm.
broad, elliptic to elliptic-lanceolate,
acute; leaf-sheaths glabrous, not
verrucose. Inflorescence usually 2-
3-flowered, from the axils of the
uppermost leaf; flowers small for
the genus, yellow. Sepals and petals
20-25 mm. long and 4-5 mm.
broad, linear-lanceolate, acute. Lip
20-25 mm. long and about 10-15
mm. broad, oval to ovate in out-
line, trilobulate toward the apex,
with several (about 7) longitudinal
lamellae which become lacerated
toward the apex and are branched
into two groups at the base; mid-
Fig. 73. Sobralia suaveolens
lobe about 8 mm. long and broad,
subquadrate. Column about 12 mm. long.
Panama.
PANAMÁ: Río La m alt. 0-25 m., Allen 50. CANAL ZONE: near Vígia and San
Juan on R. Pequeni, alt. m., Dodge et ‘al. 16623; Rio Indio de Gatun, alt. sea-level,
Maxon 4829; around Culebra, alt. 50-150 m., Pittier 4780
The type was a specimen collected near Colén and grown in England.
15. SOBRALIA VALIDA Rolfe in Kew Bull. 1909:65. 1909.
Stems robust, glabrous, 20-25 cm. tall, 3-4-leaved, base covered with 2 tubular
sheaths. Leaves broadly elliptic, subacute, subplicate, glabrous, subcoriaceous,
12-20 cm. long, 6-8.5 cm. broad, amplexicaul; sheaths tubular, subcompressed,
ancipitous, 3-5 cm. long. Spathe abbreviated, subfoliaceous, apex subacute.
(142)
19461
FLORA OF PANAMA (Orchidaceae) 37
Bracts small. Flowers few, appearing at intervals, ochroleucous, fragrant. Pedi-
cels short, subincluded. Sepals subconnivent, lanceolate-oblong, acute, equal,
more or less concave. Lip entire, oblong, obtuse, minutely crenulate, concave,
about 4 cm. long; disc tricarinate, veins pilose at the apex. Column clavate, 2
cm. long; teeth linear-obleng, incurved, about 4 mm. long.
Panama.
DARIEN: near the Darién Gold Fields, Dr. Hodgkinson.
No specimens were seen. The description is taken from the original Latin.
16. SoBRALIA Warszewiczn Reichb. f. in Bot. Zeit. 10:714. 1852; Beitr. Orch.
Centr.-Am. 9, 7. 1, figs. І, 1-2. 1866.
Terrestrial herbs of unknown height. Stems robust, the younger ones hispidu-
lous-pubescent, becoming glabrous and more or less verrucose with age. Leaves
13-18 cm. long and 4-6 cm. broad, elliptic-oval, acute or acuminate. Inflorescence
a single terminal or subterminal flower; flowers average for the genus, purple.
Sepals up to about 5 cm. long and 2 cm. broad, broadly oblanceolate. Petals
about as long as and broader than the sepals, cuneate-obovate, acute. Lip about
5 cm. long and 3 cm. broad, flabellate, emarginate, apical part crisped, with 2
basal lamellae, somewhat verrucose at the base. Column 2—3 cm. long.
Panama.
CHIRIQUÍ: “Vulcanes von Chiriquí,” alt. 1850 m., Warscewicz 8.
We have seen only a photograph and a record of the type.
OBSCURE SPECIES
SOBRALIA MACROPHYLLA Reichb. f. in Bot. Zeit. 10:713. 1852; Xenia Orch.
1:218, #. 00. 1856.
The type of this species is apparently a sterile specimen which was accompanied
by the collector's sketch and color notes. Reichenbach later published a plate of
the species and at the same time reduced the Brazilian S. chlorantha Hook. to
synonymy. Inasmuch as the type is not satisfactory for diagnosis in this difficult
genus we think it best to place 5. macrophylla among the obscure species. It is
possible that it and 5. Rolfeana Schltr. are the same.
8. PALMORCHIS Barb. Rodr.
Patmorcuis Barb. Rodr. Сеп. & Sp. Orch. Nov. 1:169. 1877; Schweinf. &
Correll in Bot. Mus. Leafl. Harv. Univ. 8:109. 1940.
Jenmania Rolfe i in Kew Bull. 198. 1898, "i Wachter, 1897.
Rolfea Zahlbr. n Jour. Bot. 36:493. 189
Neobartlettia Schltr in Fedde Rep. Sp. Nov. 16:440. 1920.
Large or small, terrestrial, reed-like herbs. Leaves pergameneous, plicate-
veined, petiolate, usually few. Inflorescence a terminal or axillary raceme or
(143)
[Vor. 33
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sepals equal or nearly so, connivent or somewhat spreading. Petals
panicle.
Lip erect from the base of
similar to the sepals but narrower and shorter.
the column and adnate with it toward the base along the median line, broader than
the sepals and petals, enfolding the column, retuse or 3-lobed; the disc pubescent
or glabrous, callused. Column elongated, slender, more or less arcuate, footless;
anther incumbent, operculate, the terminal part fleshy and sterile; pollinia 4,
pyriform, granular or subceraceous.
A small genus allied to Sobralia. There are eight or nine species with the
center of distribution in northern South America. Specimens have not been often
collected.
a. Disc of the lip without parallel lamellae; flowers about 13 mm. long... 1. Р. TRILOBULATA
aa. Disc of the lip with 5 parallel lamellae; flowers about 17 mm. long...... 2. P. PowELLII
1. PALMORCHIS TRILOBULATA L. Wms. in Ann. Missouri Bot. Gard. 28:415, f. 20.
1941.
Caespitose, terrestrial herbs up to 35 cm. tall. Stems up to about 15 cm. long
and 4 mm. in diameter, bearing 3—4 leaves near the apex. Leaves subchartaceous,
plicate; lamina of the mature leaves 10-22 cm. long and 3-6 cm. broad, elliptic,
acute or acuminate, with 7 prominent nerves; petiole up to 7 cm. long, plicate,
nervose. Inflorescence up to 7 cm. long, terminal, much exceeded bv the leaves,
few-flowered; rachis about 3-4 cm. long; bracts 1-1.5 cm. long and 1.5-5 mm.
broad, elliptic to lanceolate-ovate, acute or acuminate. Flowers about 13 mm.
long, white. Dorsal sepal about 12 mm. long and 3 mm. broad, spatulate-ob-
lanceolate, obtuse, 3—5-nerved. Lateral sepals 10-12 mm. long апа 3-4 mm.
broad, arcuate, spatulate-oblanceolate, obtuse, 3—5-nerved, somewhat carinate
along the mid-nerve dorsally, Petals 11-12 mm. long and about 2 mm. broad,
linear-oblanceolate, obtuse, somewhat arcuate, 3-nerved. Lip about 10-12 mm.
long and about 5-6 mm. broad near the apex, oblanceolate-flabellate, trilobulate at
the apex, the base or claw adnate to the column along its median line for about
2-3 mm., more or less enfolding the column; disc of the lip pubescent, especially
so toward the base, less pubescent and fleshy-thickened toward the apex; lateral
lobes about 1 mm. long (free portion), rounded, obtuse; mid-lobe about 1.5 mm.
long, transversely oblong, blunt, slightly exceeding the lateral lobes. Column
slender, arcuate, about 10 mm. long; anther operculate, incumbent; pollinia 4,
exappendiculate, ceraceous or appearing so, pyriform.
Panama.
cocLÉ: El Valle de Antón, alt. 1000 m., Allen 2463; alt. 500-700 m., Seibert 459.
Рагмокснів Роувли (Ames) Schweinf. & Correll in Bot. Mus. Leafl. Harv.
Univ. 8:119. 1940.
Rolfea Powellii Ames, Sched. Orch. 7:32, fig. б. 1924.
~
(144)
1946]
FLORA OF PANAMA (Orchidaceae) 39
Caespitose, terrestrial herbs up to
about 60 cm. tall. Stems about 30
cm. tall and 3-4 mm. in diameter.
Leaves subchartaceous, plicate; lam-
ina of the mature leaves 15-28 cm.
long and 3-5 cm. broad, elliptic-
lanceolate, acuminate, with 5-7
prominent nerves; petiole up to 8
cm. long, plicate, nervose. Inflores-
cence terminal or axillary, up to 5
cm. long, simple or branched; bracts
5-6 mm. long, ovate, acute. Flowers
about 17 mm. long, white. Dorsal
Fig. 74. Palmorchis Powellii
broad, spatulate, obtuse. Lateral
sepals similar to the dorsal sepal but slightly arcuate. Petals about 15 mm. long
and 3 mm. broad, rotundate-flabellate, trilobulate in front; disc pilose, with 5
parallel lamellae. Column about 12 mm. long, slender, arcuate.
Panama.
CANAL ZONE: Barro Colorado Island, Starry 323; “Juan Diaz,” near Panama City,
Powell 325. |
9. STENOPTERA Presl
STENOPTERA Presl, Rel. Haenk. 95, £ 14, 1827; L. Wms. in Caldasia 5:11. 1942.
Gom phichis Lindl. Gen. & Sp. Orch. Pl. 446. 1840.
Terrestrial leafy herbs. Leaves radical or cauline, contracted or petiolate at
the base. Inflorescence a terminal, compact or loose spike. Flowers mostly small,
non-resupinate. Sepals subequal or the dorsal narrower, free or joined into a short
tube at the base, subconnivent to recurved. Petals linear to nearly as broad as
the sepals. Lip uppermost, erect or parallel to the column, contracted or unguicu-
late at the base, simple or 3-lobed lamina plane or concave or subgaleate. Column
subterete, erect, wingless, glabrous or villous; anther erect in the clinandrium;
pollen granular or dust-like. Capsule ovoid, erect.
The single species known in Panama has been referred to Gom phichis, but there
are several species, unknown to Lindley, which seem absolutely to close the gap
between the two presumed genera.
1. SrENOPTERA COSTARICENSIS Schltr. in Вей. Bot. Centralbl. 36, Abt. 2:375.
1918.
Gom phichis costaricensis Ames, Hub. & Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:37.
1934.
Terrestrial herbs up to about 6 dm. tall. Leaves basal or those on the stem ге-
duced to bracts, 7-30 cm. long and 2.5-3.5 cm. broad, mostly elliptic-lanceolate,
(145)
[Vor. 33
40 ANNALS OF THE MISSOURI BOTANICAL GARDEN
acute or acuminate, contracted at the base into a broad petiole. Scape erect, with
several bracts which are reduced upward, villous above but glabrescent below.
Inflorescence a spike 3-16 cm. long. Dorsal sepal about 8 mm. long and 2.5 mm.
broad, broadly elliptic, acute or acuminate, cucullate, 1-nerved, pubescent dorsally.
Lateral sepals similar to the dorsal but more oblong and oblique. Petals 3.5-5 mm.
long and 2.5-3.5 mm. broad, elliptic-ovate, acute or obtuse, ciliate or subfimbriate,
slightly cucullate, 3-nerved. Lip 4-5 mm. long and 3-4 mm. broad, 3-lobed,
slightly unguiculate; lateral lobes rounded; terminal lobe ovate, about 2 mm. long
and 1.5 mm. broad, fleshy, villous; disc fleshy, villous; with a small marginal
callus on either side at the base. Column about 3 mm. long, terete, straight or
nearly so. Capsule ovoid.
Costa Rica, Panama, and possibly South America.
CHIRIQUÍ: Bajo Chorro, alt. 2000 m., Davidson 163, 309.
This species is an Andean type and has several close allies, if not synonyms, in
the Andes.
10. PRESCOTTIA Lindl.
Prescottia Lindl. in Hook. Exot. Fl. 2: £. 115. 1824.
Small terrestrial herbs with basal or semi-basal leaves. Leaves usually radical
(Panama), sessile or petiolate, membranaceovs, large or small. Inflorescence a
terminal spike. Flowers small, non-resupinate. Sepals connate at their bases into
a tube, spreading or revolute. Petals narrow, adnate to the base of the column,
often revolute. Lip uppermost, unguiculate and with the claw adnate to the
sepals, lamina plane to concave, cucullate or galeate, biauriculate at the base, often
enclosing the column. Column short; stigmata 2; clinandrium erect, acuminate
or subfilamentous, the margins adnate to the rostellum; pollen granular or
powdery.
A small genus with but two distinctive species known in Panama.
a. Lamina of the leaf less than 7 cm. long; petiole much shorter than
the lamina 1. P. OLIGANTHA
aa. Lamina of the leaf more dom 7 cm. long (usually 10-20 ст.); sinc
about as long as the lam . P. STACHYODES
1. PRESCOTTIA OLIGANTHA (Sw.) Lindl. Gen. & Sp. Orch. Pl. 454. 1840; С.
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 7:20. 1938.
Cranichis oligantha Sw. Prodr. Veg. Ind. Occ. 120. 178
Prescottia panamensis Schltr. in Fedde Rep. Sp. Nov. 16: ae 1920.
Small erect terrestrial herbs up to about 32 cm. tall. Leaves 1.5-7 cm. long
and 0.8—3 cm. broad, basal, membranaceous, suberect or spreading, elliptic to
cordate or obovate, acute or obtuse; petiole shorter than the lamina, usually much
shorter. Scape erect, slender, glabrous, with several bracts which are reduced
upward. Inflorescence up to 10 cm. long, a densely flowered spike. Sepals 1.5-3
mm. long and 0.6-1 mm. broad, oblong-ovate, obtuse, the laterals oblique, connate
(146)
19461
FLORA OF PANAMA (Orchidaceae) 41
Fig. 75. Prescottia oligantha
into a short tube at the base and forming an inconspicuous mentum. Petals
1-1.5 mm. long and about 0.5 mm. broad, linear-oblong, obtuse, somewhat
oblique. Lip 1.5-2 mm. long and as broad, suborbicular, strongly cucullate, with
a small auriculate callus on either side at the base; the disc puberulent. Column
about 0.5 mm. long.
Florida, Mexico, Costa Rica, Panama, the West Indies, Venezuela, and
Colombia.
осі: Cerro Vaca, alt. 900-1000 m., Pittier 5358. coc Lt: valley of upper Rio
M ma 350 m., Allen 130; vicinity of El Valle, alt. 600-1000 m., Allen 1150.
2. PRESCOTTIA sTACHYODES (Sw.) Lindl. in Bot. Reg. 22: sub 1, 1010. 1836;
L. Wms. in Ann. Missouri Bot. Gard. 27:272. 1940.
Cranichis stachyodes Sw. Fl. Ind. Occ. 3:1427. 1799.
(147)
[ VoL. 33
42 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Terrestrial herbs, up to about 1 m. tall. Leaves basal, long-petiolate, large,
membranaceous, suberect; petiole up to about 25 cm. long, slender; lamina 8-22
cm. long and 3-16 cm. broad (those of Panamanian specimens large), elliptic or
elliptic-oval to broadly oval, acute, acuminate or obtuse, margin often sphacelate.
Inflorescence a densely flowered spike up to about 30 cm. long. Sepals 3-5 mm.
long and 0.75-1.25 mm. broad, linear-oblong, obtuse, 1-nerved, strongly coiled,
connate ас their bases into a short tube. Petals 3-5 mm. long and 0.2-0.5 mm.
broad, linear, obtuse, 1-пегуе4. Lip 4-6 mm. long, strongly concave ог calcei-
form, short-unguiculate, with a lateral callus on either side at the base, glabrous.
Column about 2 mm. long.
Mexico, British Honduras, Guatemala, Honduras, Panama, the West Indies,
Colombia, Brazil, and perhaps elsewhere in South America.
LE: vicinity of El Valle de Antón, alt. 600-1000 m., Allen 1183; same ns
Allen. per hills north of El Valle de Antón, vicinity of > Mesa, alt. about 1
Allen 2326; mountains beyond La Pintada, Hunter & Allen 623.
11. CRANICHIS Sw.
CRANICHIS Sw. Nov. Gen. & Sp. Prodr. 8, 120. 1788.
Small terrestrial herbs with basal leaves or rarely with reduced cauline leaves.
Leaves sessile or petiolate, membranaceous. Scape slender, simple, with several
bracts. Inflorescence a terminal spike. Sepals free, similar, or the laterals often
broader. Petals free or obscurely adnate to the base of the column. Lip upper-
most (i. е. flowers non-resupinate), sessile or rarely unguiculate, adnate to the
base or up to the middle of the column, erect, plane, concave or saccate. Column
short; rostellum erect, often as long as the column; pollen granular.
A small genus with but one species known in Panama.
1. CRANICHIS MUSCOsA Sw. Nov. Gen. & Sp. Prodr. 120. 1788.
Small terrestrial herbs 10-38 cm. tall. Leaves petiolate, basal or semi-basal,
usually with leaf-like bracts at the base of the scape, which are reduced upward;
lamina 3-8 cm. long and 1-4 cm. broad, elliptic, elliptic-oval or oval, acute or
obtuse, membranaceous; petiole 1-8 cm. long, usually a little shorter than the
lamina. Dorsal sepal about 2.5 mm. long and 1 mm. broad, elliptic-lanceolate,
cucullate, obtuse or acute. Lateral sepals about 2.5 mm. long and 1 mm. broad,
ovate-lanceolate, oblique, obtuse or acute. Petals 2-3 mm. long and 0.3—0.5 mm.
broad, linear or linear-oblanceolate. Lip 2.5-3 mm. long and 1.5-2 mm. broad,
simple, strongly cucullate, elliptic-oval to oblong-oval, acute or acutish. Column
1-1.5 mm. long.
Florida, Mexico, Costa Rica, Panama, and the West Indies.
COCLE: vicinity of El Valle, alt. 800-1000 m., Allen 74, alt. 600-1000 m., Allen 1235.
12. PONTHIEVA R. Br.
Pontuieva К. Br. in Ait. Hort. Kew. ed. 2, 5:199. 1813; Benth. & Hook. Gen.
Pl. 3:593. 1883.
(148)
1946]
43
FLORA OF PANAMA (Orchidaceae)
Cranichis muscosa
Fig. 76.
(149)
[Vor. 33
44 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Small glabrous or pilose terrestrial herbs with basal leaves and naked or
bracteate scapes. Sepals free, spreading, subequal or dorsal smallest. Petals at-
tached above the middle of the column, spreading. Lip uppermost (non-
resupinate), adnate to the column by its unguiculate base, abruptly dilated from
the claw. Pollinia granular.
A small genus limited to the warmer regions of the western hemisphere.
a. Leaves a escen t 3. Р. MACULATA
aa. Leaves glabr
b. Lip with М M calluses at the base of the blade 1. P. EPHIPPIUM
bb. Lip without calluses at the base but the claw with a single callus... 2. P. RACEMOSA
1. PoNTHiEvA Ернтррюм Reichb. f. in Linnaea 28:382. 1856; L. Wms. in Ann.
Missouri Bot. Gard. 26:280. 1939.
Small terrestrial herbs up to about 3.5 dm. tall. Leaves basal, 5-12 cm. long
and 1.2-3.5 cm. broad, petiolate, variable, elliptic to oblong to oblanceolate to
oval or subcordate, acute or obtuse, membranaceous. Inflorescence a few-several-
flowered raceme on a bracteate peduncle, pubescent. Flowers small, white. Dorsal
sepal 4,5-7 mm. long and 1.5-2.5 mm. broad, elliptic to broadly oblanceolate,
acute, pubescent dorsally. Lateral sepals 4.5-7 mm. long and 2-5 mm. broad,
obliquely ovate, acute or obtuse, pubescent dorsally. Petals 4—6.5 mm. long and
1-3 mm. broad, oblique and with the anterior margin developed into a large sub-
basal auricle or angle, unguiculate, obtuse. Lip 2-5 mm. long, trilobulate, un-
guiculate, attached near the apex of the column; the lamina from obovate in
outline to transversely oblong, with 2 calluses near the base, mid-lobe small, about
0.5 mm. long; claw thickened.
Mexico, Guatemala, Honduras, and Panama.
CHIRIQUÍ: Finca Lérida to Boquete, alt. about 1300-1700 m., Woodson, Allen t$
Seibert 1116.
2. PoNTHIEVA КАСЕМОЗА (Мај) Mohr in Contr. О. S. Nat. Herb. 6:460. 1901;
Ames & Schweinf. in Sched. Orch. 10:14. 1930.
Arethusa racemosa Walt. Fl. Carol. 222. 1788.
Neottia glandulosa Sims in Bot. Mag. 21: /. 842. 1805.
Ponthieva glandulosa R. Br. in Ait. Hort. Kew. ed. 2, 5:200. 1813.
Pontbieva rostrata Lindl. in Ann. & Mag. Nat. Hist. 15:385. 1845.
Pont bieva guatemalensis Reichb. f. in Beitr. Orch. Centr.-Am. 63. 1866.
Mostly small terrestrial herbs up to about 6 dm. tall, but usually about 2-3
dm. tall. Leaves basal, 4-30 cm. long and 1.5-6 cm. broad, variable, elliptic to
oblanceolate to ovate to obovate, acute, petiolate, glabrous. Inflorescence pubes-
cent, а few-many-flowered, loose raceme; bracts prominent. Dorsal вера! 4-7
mm. long and 1.5-3 mm. broad, lanceolate, acute or acuminate, pubescent dorsally.
Lateral sepals 4.5-7 mm. long and 2-4 mm. broad, oblique, lanceolate to ovate-
(150)
19461
45
FLORA OF PANAMA (Orchidaceae)
Ponthieva racemosa
Fig. 77.
(151)
[VoL. 33
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN
lanceolate, acute, pubescent dorsally. Petals 4.5-6 mm. long and 2-4 mm.
broad, obliquely subovate or subtriangular, unguiculate, obtuse. Lip 4-7 mm.
long and 2-5.5 mm. broad, trilobulate, unguiculate, suborbicular to subquadrate
in outline; lamina fleshy, cucullate at the base; claw fleshy with a conspicuous
callus at its base.
Virginia to Florida and Louisiana (U.S.A.), Mexico, Guatemala, Honduras,
Costa E Panama, the West Indies, and in South America to Peru.
RiQUÍ: Volcán de Chiriquí, alt. 3500-4000 m., Woodson & Schery 460. cock:
бегу of El Valle, alt. 600-1000 m., Allen 1161. PANAMA: San Juan ыға alt. near
sea- аа, Powell 3 56.
3. PoNTHIEVA MACULATA Lindl. in Ann. & Mag. Nat. Hist. 15:385. 1845;
Hook. f. іп Bot. Mag. 108: f. 0637. 1882.
Pontbieva Brenesii Schltr. in Fedde Rep. Sp. Nov. Beih. 19:165. 1923.
Small pubescent terrestrial herbs up to about 3 dm. tall. Leaves basal, 5-20
cm. long and 0.8—5.2 cm. broad, variable, linear-lanceolate or oblanceolate to
ovate, acute or acuminate, petiolate, strigose-pubescent on both surfaces, mem-
branaceous. Inflorescence а few-—several-flowered raceme on a slender bracteate
peduncle. Dorsal вера! 9-14 mm. long and 3--4.5 mm. broad, elliptic to lanceolate,
acute or acuminate, pubescent dorsally. Lateral sepals 9-14 mm. long and 5.5-8
mm. broad, broadly oval to obovate, obtuse, pubescent dorsally. Petals 6-9 mm.
long and 2-3 mm. broad, oblique, obtuse, angled toward the base anteriorally,
short-unguiculate, inserted above the middle of the column. Lip 3-4 mm. long
and 2—3 mm. broad, oblong to suborbicular, trilobulate, cucullate, with a hood-
shaped callus which forms a cavity at the base, inserted near the apex of the
column.
Mexico, Costa Rica, Panama, Venezuela, Colombia and Ecuador.
CHIRIQUÍ: Cerro Punto to headwaters of Río Caldera, alt. 2250-2500 m., Allen 1430.
One of the most pubescent orchids in Panama.
13. SPIRANTHES L. C. Rich.
SPIRANTHES L. C. Rich. in Мет. Mus. Par. 4:50. 1818, пот. conserv.; Schltr.
in Beih. Bot. Centralbl. 37, Abt. 2:317-454. 1920.
Gyrostachys Pers. 2 РІ. 2:511. 1807
Stenorrbyncbus. С ich. in ae Mu Par. 4:59. 1818.
Pelexia L. C. Rich. I cit
Sarcoglottis Presl, Rel. Henk 1:95, t. I5. 1827.
Cyclopogon Presl, loc. cit
Belogloth is is Schltr. га Већ. ~ Centralbl. 37, Abt. 2:364. 1920.
Brachystele Schltr. loc. cit. 370
Small, or rarely large, terrestrial (rarely epiphytic) herbs with basal or cauline
leaves or both, or leafless. Roots often fleshy, fasciculated or tuberous. Leaves
various, produced before, with, or after, the flowers. Sepals free; dorsal sepal
usually erect and forming a galea with the petals; lateral ones erect or spreading,
(152)
1946]
FLORA OF PANAMA (Orchidaceae) 47
affixed to the summit of the ovary, decurrent and forming a free or adnate
mentum. Petals usually narrow and usually coherent to the dorsal sepal. Lip
sessile or clawed, plane, concave or gibbous, simple or lobed, in some species bi-
caudate at the base, adherent to the column in almost all the species, ecallose or
callose. Column terete; clinandrium often membranaceous and conspicuous, often
continued into the rostellum; rostellum various, inconspicuous or conspicuous,
truncate and retuse to lobed to aristate; anther dorsal, erect, sessile or stipitate;
pollinia 2, powdery or granular, usually attenuated at one end.
A large and technical genus with its main center of distribution in Mexico.
a. Lip less than 8 mm. long.
b. Lip less than 3 mm. long . S. GUYANENSIS
bb. Ка т. ore lon
ral mare al ligulate to pun petals oblanceolate-ligulate....2. S. COSTARICENSIS
cc. ` Lateral sepals linear-lanceolat to arrowly lanceolate-triangular;
Lans s linear to narr ovii allip 3. S. PRASOPHYLLUM
aa. Lip
le iouis.
b. dede ыы p arsa a produced, acute or obtuse mentum at the
base.
c. Lip bicaudate at the base 9. S. FUNCKIANA
. S. ORCHIOIDES
oo
cc. Lip not bicaudate at the base
bb. Lateral sepals not forming a produced, obtuse or acute mentum at
their base.
c. Lip not bicaudate at the base; rostellum terete 6. S. SPECIOSA
. Lip bicaudate at the base; rostellum flattened or tere
“2 Terminal part of the lip not = nor јаје nor with
pubescent рақ ridges; rostellum t
dd. тана part of the lip when or Mes or with pubescent
ened.
ч
. 5. NAVARRENSIS
callus ges; rostellum flat
e. Base т й stem leafy; рақ? from rhizomes 4. S. Моорзоми
ee. Base of the stem not leafy; plants from fascicled roots.......... 5. S. ACAULIS
. SPIRANTHES GUYANENSIS (Lindl.) Cogn. in Mart. Fl. Bras. 34:209, /. 47, f. П.
1895.
Good yera guyanensis Lindl. Gen. & Sp. Orch. Pl. 494. 1840.
Brachystele guyanensis Schltr. in Beih. Bot. Centralbl. 37, Abt. 2:373. 1920.
рта 3 aguacatensis Reichb. f. in Bonplandia 3:214. 1855; Ames, Sched. Orch. 2:7.
92
Sees aguacatensis O. Ktze. Rev. Gen. 2:664. 1891.
Brachystele aguacatensis Schltr. lod cit, ЗАК
Erect terrestrial herbs up to 20 cm. tall. Stems with several loose, lanceolate
or ovate-lanceolate, acuminate bracts. Leaves unknown. Inflorescence up to 6
cm. long, densely flowered. Dorsal sepal 2-3 mm. long and about 0.7-1.5 mm.
broad, lanceolate, acute or obtuse. Lateral sepals 2-3 mm. long and 0.7-1.5 mm.
broad, lanceolate-triangular, acute or obtuse, somewhat oblique. Petals 2-2,5 mm.
long and 0.6—0.9 mm. broad, oblanceolate-spatulate, truncate, or obtuse, coherent
to the dorsal sepal, unguiculate. Lip 2-2.8 mm. long and 1.7-2.2 mm. broad,
oblong-pandurate to ovate or suborbicular and constricted toward the apex, the
terminal lobe small, oblong to suborbicular, with two retrorse auricles or caudae
at the base.
(153)
[Vor. 33
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico, Guatemala, Honduras, Costa Rica, Panama, Trinidad, the Guianas,
Colombia, and Bolivia.
CANAL ZONE: open fields near railway station, Monte Lirio, Maxon 6862.
The smallest-flowered species of Spiranthes in Panama.
N
SPIRANTHES COSTARICENSIS Reichb. f. in Bonplandia 3:214. 1855; Beitr. Orch.
Centr.-Am. 46. 1866; Xenia Orch. 2:185, #. 179, f. I, 1-30. 1868; Ames,
Sched. Orch. 9:7. 1925.
Gyrostachys costaricensis О. Ktze. Rev. Gen. 2:664. 1891
Beloglottis costaricensis Schltr. in Beih. Bot. Centralbl. 37, Abt. 2: a. 1920.
Spirantbes subpandurata Ames & Schweinf. Sched. Orch. 8:4, f. 1
Erect terrestrial or epiphytic herbs with basal leaves, or leafless at flowering
time, up to 4.5 dm. tall. Leaves 3-18 cm. long and 1-5 cm. broad, elliptic to
oval or oblanceolate, acute or acuminate, membranaceous, petiolate; petiole 2—8
cm. long. Inflorescence a few-many-flowered raceme, up to about 20 cm. long.
Dorsal вера! 4—6.5 mm. long and 1-1.5 mm. broad, lanceolate, acute, pubescent
dorsally. Lateral sepals 4—6.5 mm. long and 0.7-1.3 mm. broad, linear-ligulate
to ligulate, acute or obtuse, pubescent dorsally. Petals 3.5-6 mm. long and 0.7-1
mm. broad, oblanceolate-ligulate, acute or obtuse, oblique, coherent to the dorsal
sepal. Lip 5—6.5 mm. long and 1.5-2 mm. broad, lanceolate-pandurate, short-
unguiculate, retrorsely bicaudate at the base.
Mexico, Guatemala, Honduras, Costa Rica and Panama.
CANAL ZONE: upper Chagres River Range, Powell 377.
А species easy to distinguish because of the unusual lip shape.
3. SPIRANTHES PRAsoPHYLLUM Reichb. f. Beitr. Orch. Centr.-Am. 65. 1866;
Ames, Sched. Orch. 9:8. 1925.
Gyrostachys Prasophyllum O. Ktze. Rev. € 2:664. 1891.
Spiranthes epiphytica Schltr. in Fedde Rep. Sp. Хоу. 2:130. 1906
Cyclopogon Prasophyllum Schltr. in Beh. P» Centralbl. 37, Abt. 2:393. 1920.
Small epiphytic herbs 8-25 cm. tall, with basal leaves. Leaves short-petiolate;
lamina 3-13 cm. long and 0.8-2.8 cm. broad, elliptic, acute; petiole 1-4 cm. long.
Inflorescence 2—8 cm. long, the flowers often unilateral. Dorsal вера! 5-6 mm.
long and 1-1.5 mm. broad, lanceolate, acute or acuminate, pubescent at the base
dorsally. Lateral sepals 5—6.5 mm. long and 1—1.8 mm. broad, linear-lanceolate to
narrowly lanceolate-triangular, acute, pubescent at the base dorsally. Petals 4-5
mm. long and 0.5—1 mm. broad, linear to narrowly elliptic, acute or obtuse, co-
herent to the dorsal sepal. Lip 5-7 mm. long and 2-3 mm. broad, narrowly ob-
long-pandurate to lanceolate-ovate and constricted toward the apex; apical lobe
oblong to transversely oval or reniform, bicaudate at the base, the caudae retrorse
or incurved.
Guatemala, Honduras, Costa Rica, and Panama.
cocLÉ: north of El Valle de Antón, alt. about 1000 m., Allen 2004. CANAL ZONE:
Gatun, Hayes 138; forest, Rio Indio at Gatun, Maxon 4816.
(154)
49
19464
FLORA OF PANAMA (Orchidaceae)
)
> rr
re ЗИ
4228 Ww 24
мй >
Sage ORT;
%
Дае
y
SV
WE
mí
M
> 9x
44.
M
||
AY H
Ун
ЛЫ ;
ІШ 4
ШЕ
/)
M
М
Fig. 78. Spiranthes Prasophyllum
4. SPIRANTHES Моорзохи L. Wms. in Ann. Missouri Bot. Gard. 29:337. 1942
Rhizomes slender, rooting at
Stems slender,
Terrestrial palustrine herbs up to 5.5 dm. tall.
most of the nodes, with scarious sheaths arising from the nodes.
with well-developed leaves at the base which become bract-like above, glabrous
(155)
[Vor. 33
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN
below but becoming densely pubescent above. Leaves 3-10 cm. long, 1.3-2.8 cm.
broad, oblong-elliptic to oval, acute or obtuse, largest near the base of the stem
and reduced to amplexicaul bracts above. Inflorescence up to 10 cm. long, flowers
congested, becoming more open in fruit; bracts up to 4 cm. long and 1.6 cm.
broad, lanceolate, acuminate, pubescent dorsally. Flowers large, similar to those
of S. acaulis. Dorsal sepal 16.5-19 mm. long and 3.5-4 mm. broad, lanceolate,
acuminate, 3—5-nerved, densely pubescent dorsally. Lateral sepals long-decurrent
on the ovary, 35-40 mm. long from the apex to the base of the saccate mentum;
free part 16.5-19 mm. long and 4-4.5 mm. broad, lanceolate, acuminate, arcuate,
spreading, densely pubescent dorsally, 3—5-nerved. Petals 15-18 mm. long and
2-2.5 mm. broad, narrowly elliptic to narrowly oblanceolate, pubescent or ciliate
near the margins of the basal half. Lip 28-32 mm. long and 6-7 mm. broad
(apical lobe), linear-oblong and somewhat expanded and pandurate in the terminal
third, caudate, with two extremely pubescent converging callus-ridges on the
terminal third, the basal part densely pubescent and the remainder, except the
glabrous apex, less pubescent, glabrous dorsally except at the base; terminal lobe
oval or transversely rhombic; caudae about 5 mm. long, retrorse, fleshy but
flattened.
Panama.
то vicinity of El Boquete, alt. 1000-1500 m., b iier 2050; vicinity of
lions. alt. 1200-1500 m., Woodson 9 Schery 753; i mpy meadows, Finca Lérida
to Boquete, alt. about 1300-1700 m., Woodson, Allen bi Seibert 1148.
Very few species of Spiranthes have rhizomes.
5. SPIRANTHES ACAULIS (J.E.Sm.) Cogn. in Mart. Fl. Bras. 3*:221. 1895.
Neottia acaulis J. E. Smith, Exot. Bot. 2:91, /. 105. 1806.
Arethusa picta Anders. in Trans. Soc. Arts. 25. 1807.
Neottia жек Br. in Ait. Hort. Kew. ed. 2, 5:199. 1813; Sims in Bot. Mag. 37: #. 1562.
181
Ld picta Klotzsch in Allgem. Gartenz. 10:106. 1842; Schltr. in Beih. Bot.
© , Abt. 2: "r^ 1920.
Gyrostachys picta O. Ktze. Rev. Gen. 2:664. 1
Sarcoglottis hd. Schltr. in Fedde Rep. ш Nov. Beih. 17:13. 1922.
docent Powellii Schltr. loc. cit
Sarcoglottis Purpusiorum Schltr. in Fedde Rep. Sp. Nov. 21:333. 1925.
Large terrestrial herbs with basal leaves or with the leaves withered at flower-
ing time, up to 1 m. tall but more commonly 3-4 dm. tall. Leaves petiolate or
epetiolate; lamina 6-18 cm. long and 4-7 cm. broad when mature, elliptic to oval
to oblanceolate or obovate, acute or obtuse, membranaceous, mottled; petiole
short or none. Inflorescence up to 2 dm. long, few-many-flowered; bracts up to
4 cm. long, linear to linear-lanceolate, acuminate. Dorsal вера! 15-22 mm. long
and 3-4 mm. broad, ligulate, acute, pubescent dorsally. Lateral sepals 40-50 mm.
long and 4-7 mm. broad, free part 18-28 mm. long, lanceolate, acute, arcuate,
pubescent dorsally, basal part decurrent on the ovary and extending almost to
the base. Petals 14-24 mm. long and 2-4 mm. broad, linear-lanceolate or usually
(156)
19461 5
FLORA OF PANAMA (Orchidaceae) 51
linear-oblanceolate, acute or obtuse, slightly arcuate, coherent to the dorsal sepal.
Lip 30-40 mm. long and 7-11 mm. broad, essentially narrowly oblanceolate, ob-
tuse, the dilated apical portion constricted and with an oval or suborbicular
terminal lobe, disc pubescent below the constriction, bicaudate at the base, the
caudae fleshy, retrorse.
From Mexico through Central America and the West Indies, in South America
to northern Argentina.
ANAL ZONE: Culebra, Pittier 3427; near Panama City, San Juan, Manteca, Mata
қойады Juan Diaz, Ari as Hill, Е rijoles, Powell 147, 179, 388, 380, 300, 301, 302, 394,
БАЛ 30%, 309, 3435, 3521, 3525, 3531, 3532, 3534, 3536, 3541, 3551, 3567, 3568, 3569,
Known іп Panama only from the Canal Zone where it grows in fairly open
situations. A variable species.
6. SPIRANTHES sPECIOSA (J.F.Gmel.) A. Rich. іп La Sagra, Hist. Isla Cub.
ей. 2, 11:252. 1850.
Satyrium plantagineum L. SHE Veg. ed. 10, 1244. 1758-59, non Spiranthes plantaginea
Lindl., Spreng. vel Tor
Serapias speciosa J. F. rel. Syst. 59. 1791.
Neottia speciosa Jacq. Ic. Pl. Rar. 3:1. боо. 1793; Sims’ Bot. Mag. 33: 1. 1374. 1811;
DER 3, 4. 10
Stenorrhynchus speciosus L. C. Rich. in Мет. Mus. Par. 4:59. 1818.
Spiranthes colorata М. E. Br. in Gard. Chron. n. s. 19:210. 1883.
Erect terrestrial or epiphytic herbs up to 4 dm. tall, with a rosette of basal
leaves at flowering time. Leaves 4-18 cm. long and 1-8 cm. broad, еШіргіс to
ovate or obovate, acute or acuminate; petioles 2-15 cm. long. Stems with several
amplexicaul bracts. Inflorescence up to 10 cm. long, flowers congested; bracts
up to 5 cm. long, linear-lanceolate, acuminate, usually red. Dorsal вера! 12-14
mm. long and 3-4 mm. broad, lanceolate, acute or acuminate, glabrous. Lateral
sepals 12-15 mm. long and 3-4 mm. broad, lanceolate to narrowly lanceolate-
triangular, acute or acuminate, forming a short mentum at the base. Petals 12-14
mm. long and 3—3.5 mm. broad, lanceolate or elliptic-lanceolate, somewhat arcuate,
acute or acuminate, coherent to the dorsal sepal. Lip 13-15 mm. long and 4.5-5
mm. broad, terminal lobe lanceolate or oblong, acute, basal part oblong, pandurate,
the basal margins somewhat thickened, disc pubescent.
Mexico, Guatemala, Honduras, Costa Rica, Panama, the West Indies, and in
South America.
осі: vicinity of "New Switzerland," central valley of Río Chiriqui Viejo, alt.
1800-2000 m., Allen 1382; Salta Boquete, Boquete District, alt. 5500 feet, Terry 1267.
7. SPIRANTHES NAVARRENSIS (Ames) L. Wms. in Ann. Missouri Bot. Gard.
22:397, .1942.
Stenorrbyncbus navarrensis Ames, Sched. Orch. 9:15, f. 3. 1925.
Erect or ascending terrestrial herbs up to about 5 dm. tall. Leaves basal,
petiolate; lamina 7-16 cm. long and 3.5-8.5 cm. broad, narrowly to broadly
(157)
[Vor. 33
52 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ovate, acuminate, membranaceous, often oblique; petiole 9-20 cm. long, winged
and closed or sheathing at the base. Peduncle usually erect, with several mem-
branaceous, sheathing bracts. Inflorescence up to 10 cm. long, few-many-
flowered. Dorsal sepal 18-21 mm. long and 3.5-4.5 mm. broad, lanceolate, acute.
Lateral sepals 18-22 mm. long and 3-4 mm. broad, lanceolate, more or less ob-
lique. Petals 16-20 mm. long and about 2 mm. broad, linear-lanceolate, acute,
subsigmoid, connate to the dorsal sepal. Lip 16-18 mm. long and 4-7 mm. broad,
essentially lanceolate, constricted just above the base and again above the middle,
auriculate at the base, the auricles fleshy. Anther up to 11 mm. long.
Costa Rica and Panama.
CHIRIQUÍ: cloud forest, Cerro Horqueta, alt. about 2000 m., von Hagen 9 von Hagen
TJI.
Closely allied to S. speciosa.
8. SPIRANTHES ORCHIOIDES (Sw.) L. C. Rich. in La Sagra, Hist. Isla Cuba, ed. 2,
11:252. 1850.
Satyrium orchioides Sw. Nov. Gen. & Sp. Prodr. 118. 178
Neottia orchioides Sw. Fl. Ind. Occ. 3:1411. 1799; Sims in жүт” Mag. 25: f. 1036. 1807;
Edwards’ Bot. Reg. 9:7. 701. 1823.
Stenorrhynchus orchioides L. C. Rich. in Mém. Mus. Par, 4:59. 1818.
Spiranthes jaliscana S. Wats. in Proc. Am. Acad. 26:153. 1891.
Stenorrhynchus jaliscana Nash in Bull. Torr. Bot. Club 22:158. 1895.
Strict, leafless (at flowering time), terrestrial herbs up to 7 dm. tall but nor-
mally about 3 dm. tall. Leaves appearing after the flowering stage, 15-21 cm.
long and 2,5-4 cm. broad, elliptic, membranaceous. Stems with several amplexi-
caul bracts which occasionally become leaf-like below. Sepals connivent or spread-
ing at the tips, pubescent dorsally. Dorsal sepal 13-21 mm. long and 3-6 mm.
broad, lanceolate, acute. Lateral sepals 18-27 mm. long and 3-4.5 mm. broad,
lanceolate, acute, the bases decurrent on the ovary and at the base forming a pro-
truded, acute or obtuse mentum. Petals 12-15 mm. long and 2.2-4.5 mm. broad,
elliptic to elliptic-linear, acute or obtuse, coherent to the dorsal sepal. Lip 15-23
mm. long and 5.5-9 mm. broad, lanceolate, acute or acuminate, the margins of
the broad claw fleshy and pubescent.
Florida and Mexico through Central America to Panama, the West Indies, and
in South America to northern Argentina.
CANAL ZONE: EUN alt. 75 m., Allen 2453; Bella Vista, Ceiba Tierra, Monte Lirio,
near Gatun, La Сһоггега, Тесит Ts ia, Mt. McComber, Mata R Redonda, Powell 386,
387, 393, 396, 402, 404, 407, 408, 400, 410, 414, 3454, 3527, 3547, 3557. 3559.
The flowers of this species vary іп color from white to brilliant red.
кезі
52
Panama they are often greenish. А widespread and variable species.
9. SPIRANTHES FUNCKIANA Rich. & Gal. in Ann. Sci. Nat. ІП, 3:32. 1845.
Pelexia Pringlei Fernald in Proc. Am. Acad. 35:562. 1900.
Pelexia Funckiana Schltr. іп Fedde pud Sp. Nov. 15:197. 1918, as P. Funkiana; Ames
& Schweinf. Sched. Orch. 8:5.
(158)
1946]
FLORA OF PANAMA (Orchidaceae)
Fig. 79. Spiranthes orchioides
(159)
53
[Vor. 33
54 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Pelexia guatemalensis Schltr. loc. cit.
Pelexia congesta Ames & Schweinf. Sched. Orch. 10:5. 1930.
Slender terrestrial herbs up to 4.5 dm. tall. Leaves basal, petiolate; lamina
6-13 cm. long and 1.5—5 cm. broad, lanceolate to ovate, acute or acuminate,
membranaceous; petiole 3-10 cm. long. Inflorescence up to 13 cm. long, several-
many-flowered, lax or dense; bracts 1-3 cm. long, linear or linear-lanceolate,
acuminate, somewhat pubescent dorsally. Dorsal sepal 14-19 mm. long and 4-5
mm. broad, oblanceolate, obtuse or acute, pubescent dorsally. Lateral sepals 25—35
mm. long and 3-3.5 mm. broad, linear-oblanceolate, acute, pubescent dorsally,
oblique, free part strongly recurved, 14-18 mm. long, the basal part adnate to
and decurrent on the ovary, 12-16 mm. long, forming a free spur-like projection
at the base. Petals 14-19 mm. long and 2.5—3 mm. broad, linear-oblanceolate,
acute or obtuse, coherent to the dorsal sepal. Lip 18-27 mm. long and 4-6.5
mm. broad, linear, subpandurate, with two retrorse auricles at the base, the disc
puberulent toward the base.
Mexico, Guatemala, Honduras, Panama, and possibly South America.
cocLÉ: hills north of El Valle de Antón, vicinity of La Mesa, alt. about 1000 m., Allen
2315.
14. ERYTHRODES Blume
Екутнкоре5 Blume, Bijdr. Fl. Nederl. Ind. 410, #. 72. 1825.
Physurus L. C. Rich. іп Мет. Mus. Par. 4:55. 1818, nomen.
Terrestrial herbs with semi-basal or cauline leaves. Sepals free, nearly equal,
erect or spreading. Petals connate toward their apices and usually coherent to the
dorsal sepal and with it forming a galea. Lip simple or lobed, slightly adnate to
the column for a short way, extended into a simple or didymous spur at the base
which usually contains four or more mammillate calli or callus-like structures at
the base. Column short. Pollinia sectile or granular.
A genus found in the tropics and subtropics of both hemispheres. The species
are difficult to interpret.
A single species recognized in Panama.
1. Екүтнкореѕ Киллри Ames in Proc. Biol. Soc. Wash. 34:150. 1921.
Erect or ascending terrestrial herbs from a creeping rhizome, up to 1 m. or
more long. Stems leafy toward the base, slender. Leaves petiolate; lamina 6-13
cm. long and 2—4.5 cm. broad, lanceolate to ovate or suborbicular, acuminate or
acute, often oblique, membranaceous; petioles 2-5 cm. long, inflated and scarious
at the base, semiamplexicaul. Sepals connate nearly to their apices. Dorsal вера!
5-7 mm. long and 2-3 mm. broad, lanceolate to oblong-lanceolate, obtuse or
acute, pubescent dorsally. Lateral sepals 5.5—7 mm. long, 1.5-2.5 mm. broad,
narrowly elliptic to oblong-lanceolate, obtuse or acutish, pubescent dorsally. Petals
5-6.5 mm. long and 1.5-2.5 mm. broad, oblanceolate, obtuse. Lip saccate; lamina
(160)
19461
FLORA OF PANAMA (Orchidaceae) 55
5-6.5 mm. long and 2.5-3.5 mm. broad, the basal part oblong, canaliculate, fleshy,
the terminal lobe bi- or trilobulate, transverse, often ciliolate; sac 4-5 mm. long,
obtuse, straight or curved slightly.
CLE: vicinity of El Valle, alt. 1000 m., Allen 120I and 1702. cHrriQuf: Bajo
re бын alt. 6000 feet, Davidson 317; valley of the Río Caldera from El Boquete to
the Cordillera, alt. 1400-1600 m., Killip 3561.
More adequate material may show that the specimens from Coclé represent
a different species.
15. CORYMBORCHIS Thouars
CoryMBorCHIs Thouars, Orch. Iles Afr. №. 37, 38. 1822.
Corymbis Thouars, loc. cit., is Corymborchis by elision.
Terrestrial herbs with plicate or subplicate leaves and often with branched
stems. Sepals and petals connivent into a tube at the base, usually linear with
the petals dilated at the apex. Lip canaliculate, erect from the base of the column
and free from it, apex usually dilated and recurved. Column erect, slender, the
ze clavate, bilobed; anther erect, acuminate, E to the rostellum; pollinia
2, granular.
The genus is pantropic, with two species in the western hemisphere.
1. CORYMBORCHIS FLAVA (Sw.) О. Ktze. Rev. Gen. Pl. 2:658. 1891.
Serapias flava Sw. Nov. Gen. & Sp. Pl. Prodr. 119.
Corymbis flava Hemsl. in сө & Salv. Biol. Centr. Аға. Bot. 3:297. 1884.
ТАП, slender, terrestrial, leafy herbs up to 15 dm. tall. Stems up to about 6
mm. in diameter, covered with the leaf sheaths. Leaves when mature 25—45 cm.
long and 3.5-7 cm. broad, elliptic or elliptic-lanceolate, acuminate, plicate, sessile
or nearly so. Inflorescence a simple raceme or a panicle from the axils of the
upper leaves, up to 13 cm. long; bracts up to 10 mm. long, ovate, acute. Dorsal
вера! 13-15 mm. long and about 2.5 mm. broad, linear to linear-lanceolate, acute.
Lateral sepals 13-15 mm. long and about 2 mm. broad, linear or linear-lanceolate,
acute or arcuate. Petals 13-14 mm. long and about 4 mm. broad, oblanceolate,
acute. Lip 12-15 mm. long and 4-7 mm. broad, lanceolate to ovate-lanceolate,
acute, cucullate, with a callus ridge extending along the margins from the base
up to the middle or beyond.
British Honduras, Salvador, Costa Rica, Panama, the West Indies, also in South
America.
CHIRIQUÍ: llanos, on slope of Volcán de Chiriquí Viejo, alt. 1200 m., Allen 1000.
16. STELIS Swartz
STELIS Swartz іп Schrader's Jour. f. Bot. 2:239. 1799; in Svenska Vet.-Akad.
Handl. 21:248. 1800, nom. conserv.
(161)
[Vor. 33
56 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Humboltia Ruiz & Pavon, Fl. Peruv. et Chil. Prodr. 121, pl. 27. 1794, non Vahl.
Small to fairly large epiphytic herbs with caespitose or repent stems; secondary
stems terminated by a single, usually fleshy, often petiolate leaf. Sepals subequal or
the dorsal longest, more or less connate at the base, the laterals sometimes connate
to their tips (S. Allenii). Petals much smaller than the sepals, broad, often flabel-
late, anterior margin usually thickened. Lip sessile, simple or 3-lobed, usually fleshy
and about as long as the column. Column short, footless; stigmas confluent or
separated. Anther terminal, operculate, incumbent, biloculate; pollinia 2,
ceraceous.
A difficult genus to study because the petals and lip are usually very small and
often extremely fleshy.
a. ин, a Жар shorter than the dorsal sepal, connate to
middle
ddp ovate ae the ‘thick central callus concave in front 3. 8. DESPECTANS
bb. Lip not ovate, without a thickened ce Me 1 callus
c. Dorsal sepal more than 10 mm. "e посен 1. S. ALLENI
сс. Dorsal вера! less than 5 mm. long, 2. S. INAEQUALIS
aa. эж у sepals not Em shorter et ge dorsal sepal, connate
only at their bas
b. Lip up to more I twice as long as broad.
c" al sepal 3.5 mm. or more broad; leaves 3 cm. or more
oad 5. S. MONTANA
cc. = бергі less than 3 mm. broad; leaves 2 cm. or less broad.
d. Sepals 4 mm. or more long, теби пи -оуасе to oblong-lanceo-
late, inner құла glandular 7. $. котснп
dd. Sepals 3 mm. r less long, triangular to orbicular.
e. Lip 1-1.5 mm. long; sepals triangular.: 6. S. ATRORUBENS
0.5—0.6 mm. long; sepals ovate to orbicular 4. S. MICROCHILA
bb. Lip about as long as broad or broader than long
c. Lip uced into an apicule or acumen in Mis
ip produced into a broad acumen, acumen not erect.................- 8. S. HYMENANTHA
dd. Lip produced into an inconspicuous apicule igi 3) or
fleshy erect apicule.
e. Apicule fleshy, erect 9. S. CRESCENTIICOLA
ee. Apicule not fleshy, erect.
f. Sepals 2-4 mm. long, usually 3-nerved 11. S. ENDRESII*
ff. Sepals 4-9 mm. long, usually 5-nerved 10. S. LEUcoPocoN*
ront.
. Callus of н lip cruciform; bracts on rachis prominent, acute
or acuminat 12, S. PowrrLII
dd. Callus of чи Пр not crucifor
е. Callus of the li Рађен ан Пр with two minute auricles
at the base; plants mostly less than 5 cm. tall 3. 5.5токкп
ee. Callus of the lip те Бр without бды ас the base;
plants mostly more than 5 tall.
f. Inflorescence subequal to or n or but slightly longer
than the subtending leav 14. S. Махомп
апа 15. S. PANAMENSIS* *
ff. Inflorescence twice or more longer than the subtending
leaves.
2. Secondary к usually 3 cm. long or longer.
h. Sepals dens glandular-pubescent within 16. 8. VESTITA
hh. one not m glandular-pubescent within, rarely
pubescent 17. S. AEMULA
gg. Secondary = s 3 cm. or less long.
Һ. callus running at right angles to the trans-
verse 4: viii base of lip 18. S. COLLINA
(162)
1946)
FLORA OF PANAMA (Orchidaceae)
те“
EV
~ Q
а А 2 >>,
Ly 9а,
= >
--/4
|
|)
My
WE. V) M
D S
ЕМ “4
Fig. 80. Stelis Allenii
(163)
27
^ (Мог, 33 '
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN
hh. No callus-ridge running at right angles to the trans-
verse callus
i. Sepals 3. 54.5 mm. long 19. S. WILLIAMSI
ii. Sepals 1.5-2.5 mm. long.
j Bracts of the inflorescence мн. rne WR LE 17. S. AEMULA
jj. Bracts of the inflorescence well s ted. a 20. S. LONGIPETIOLATA
*'These two species appear somewhat different but il. are not dist
**$. panamensis would seem to key out here also, but the material available is so scanty that it
can not t be properly placed.
1. STELIS ALLENI Г. Wms. in Ann. Missouri Bot. Gard. 29:338. 1942.
Large caespitose epiphytic herbs up to 4 dm. tall. Secondary stems 7-15 cm.
long and 0.25-0.35 cm. in diameter, covered with two or three loose sheaths
which soon disentegrate, shorter than the leaves. Leaves 10-19 cm. long and
3.5—7 cm. broad, elliptic to elliptic-oval, acute or obtuse, coriaceous, attenuated
into a short petiole at the base. Inflorescence up to 30 cm. long, floriferous to
the base, one or more borne from the apex of the stems (if more than one then
presumably borne in different years); sheaths up to 2.5 cm. long, cucullate,
ample; bracts 2-18 mm. long, reduced upwards, ovate-lanceolate, acute or
acuminate, infundibuliform; flowers largest of the genus. Dorsal sepal 14-16
mm. long and 5-6 mm. broad, lanceolate, acute, 11-13-пегуед, cucullate. Lateral
sepals connate to their apices, together 10—12 mm. long and 8—10 mm. broad,
suborbicular-ovate, acute or obtuse, cucullate and gibbous at the base, many-
nerved. Petals about 1 mm. long and 1.5 mm. broad, broadly flabellate to trans-
versely oval, the apex much thickened. Lip 0.75-1 mm. long and 1—1.4 mm.
broad, about 0.75 mm. thick at the apex, flabellate, truncate, the transverse
callus at the apex of the lip, very like the petals but slightly smaller.
Endemic in Panama.
COCLÉ: hills north of El Valle de Antón, alt. 800 m., Allen 2952.
Stelis Allenii is perhaps the most distinctive species of the genus in Central
America and seems to be the largest-flowered. There are no near allies in Central
America but the species seems to belong to Lindley's section DrALISsA, a section
with but a few species in the Andes.
2. STELIS INAEQUALIS Ames, Sched. Orch. 4:12. 1923; in Bot. Mus. Leafl. Harv.
Univ. 3:155, £. 1955.
Small, caespitose, epiphytic herbs up to about 5 cm. tall. Secondary stems
about 3 mm. long, concealed by the sheathing bracts. Leaves 1.5-2.5 cm. long
and 0.4-0.5 cm. broad, oblanceolate, obtuse, emarginate, coriaceous. Inflorescence
up to 5 cm. long, exceeding the leaves; bracts about 1 mm. long, infundibuli-
form. Dorsal вера! 2-2.5 mm. long and 1-1.4 mm. broad, elliptic-oblong,
obtuse, puberulent ventrally. Lateral sepals 1—1.4 mm. long and together
1.6-2 mm. broad, connate to beyond the middle. Petals 0.5—0.75 mm. long
and 0.75-1 mm. broad, flabellate. Lip 0.6-0.75 mm. long and about 0.35—0.5
mm. broad, rather thin below the middle and much thickened above, strongly
concave in front of the obtuse apex.
(164)
19461
FLORA OF PANAMA (Orchidaceae) 59
Honduras and Panama.
PANAMA: foothills, upper reaches
of Chagres River near San Juan, alt.
near sea-level, Powell 271.
3. STELIS DESPECTANS Schltr. in
Fedde Rep. Sp. Nov. 8:453.
1910.
Stelis chiriquensis Schltr. in Beih. Bot.
Centralbl. 36, Abt. 2:386. 1918.
Stelis nutentiflora Schltr. loc. cit. 390.
Caespitose epiphytic herbs up
to about 25 cm. tall. Secondary
stems 2-12 cm. long, about as
long as or shorter than the leaves.
Leaves narrowed to a petiole at
the base; lamina 2-12 cm. long
and 2-7 mm. broad, linear-elliptic
or linear-oblanceolate; petiole up
to 4 cm. long, slender. Inflores-
cence about as long as or exceed-
ing the leaves in length. Sepals
connate at the base for a short
distance; dorsal sepal 2-3 mm.
| long and 0.75—1.5 mm. broad,
STELIS inaequalis Ames lanceolate-oblong to obovate-ob-
ong, acute or obtuse; lateral sepals
1.7-2.5 mm. long and 1.2-1.5
mm. broad, ovate, acute or obtuse. Petals 0.5—0.7 mm. long and 0.4—0.5 mm.
broad, suborbicular, apex truncate and thickened. Lip 0.6-0.75 mm. long and
0.3—0.6 mm. broad, rhombic to broadly ovate, thickened, excavate and concave in
front, the basal half much thickened and the callus submammillate.
Costa Rica and Panama.
Fig. 81
CHIRIQUÍ: between Alto de las Palmas and Cerro Horqueta, alt. 2100-2268 m.,
Pittier 3228.
The specimen cited is the largest known for the species and the maximum
measurements apply to it.
4. STELIS MICROCHILA Schltr. in Fedde Rep. Sp. Nov. 9:289. 1911.
Stelis barbata Rolfe in Kew Bull. 1913: 141. 1913.
Stelis costaricensis Schltr. in pos. Bot. Centralbl. 36, Abt. 2:388. 1918.
Stelis cinerea Schltr. loc. cit. 4
Stelis bryophila Schltr. in er p Sp. Nov. Beih. 19:16. 1923.
Small, caespitose, epiphytic herbs up to about 15 cm. tall. Secondary stems
(165)
[Vor 33
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN
up to about 2 cm. long, covered with one or more loose sheaths. Leaves 2-6 cm.
long and 0.2—0.8 cm. broad, narrowly to broadly oblanceolate, obtuse or acute,
attenuated into a narrow petiole at the base. Sepals connate for a short distance
at the base, puberulent within and sometimes dorsally; dorsal sepal 1.5-2 mm.
long and 1.3—1.5 mm. broad, ovate to suborbicular, obtuse or acutish; lateral
sepals 1.2-1.5 mm. long and 1.1-1.3 mm. broad, broadly ovate. Petals about 0.75
mm. long and 0.75-1.1 mm. broad, transversely oblong or subquadrate, the
truncate terminal margin thickened. Lip 0.5-0.6 mm. long and 0.25-0.4 mm.
broad; basal У; or 24 very fleshy, with two more or less longitudinal calluses ог
thickenings which have a distinct sinus; terminal lobe oblong or subquadrate,
obtuse, cochleate, relatively thin.
Guatemala, Costa Rica, and Panama.
COCLE: mountains beyond La Pintada, alt. 400-600 m., Hunter & Allen 590.
5. STELIS MONTANA L. Wms. in Ann. Missouri Bot.. Gard. 27:272. 1940; loc.
cit. 29:340, Ё. 32, figs. 0-12. 1942.
Caespitose epiphytic herbs up to about 53 cm. tall. Secondary stems 11-13
cm. long, covered with the sheathing cauline bracts, shorter than the leaves.
Leaves 17-20 cm. long and 3-3.5 cm. broad, oblanceolate, obtuse, coriaceous,
gradually narrowed into a short petiole. Inflorescence racemose, as long as or
longer than the subtending leaves. Sepals connate for a short distance at their
bases. Dorsal вера! about 6 mm. long and 3.5 mm. broad, ovate to lanceolate-
ovate, acute, apiculate. Lateral sepals about 5 mm. long and 4.5 mm. broad,
broadly ovate, somewhat oblique, acute, apiculate. Petals about 1.5 mm. long
and 1.5 broad, nearly orbicular, the apex thickened. Lip about 2-2.25 mm. long
and 0.8—1 mm. broad, oblong to oblong-obovate, obscurely 3-lobed, the basal half
filled with a large callus, the terminal part thin, minutely puberulent dorsally.
Endemic to Panama.
CHIRIQUÍ: trail from Cerro Punta to headwaters of Río Caldera, alt. 2250-2500 m.,
6 `
Allen 1463.
6. STELIS ATRORUBENS L. Wms. in Ann. Missouri Bot. Gard. 29:239. 1942.
Small, caespitose, epiphytic herbs up to about 18 cm. tall. Secondary stems
1.5-3.5 cm. long, slender, covered with bracts. Leaves 6-9 cm. long, petiolate,
much longer than the secondary stems; lamina about 4-6.5 cm. long and 0.5-1
cm. broad, elliptic to elliptic-oblanceolate, obtuse, fleshy; petiole 2-3 cm. long.
Inflorescence up to 15 cm. long, upper half more or less densely flowered; bracts
about 2 mm. long, infundibuliform, acute, scarious. Sepals connate at the base,
rotate, 2-2.5 mm. long and 1.5-2 mm. broad, triangular, acute, 3-nerved. Petals
about 0.75 mm. long and 0.75 mm. broad, suborbicular, 1-nerved, the terminal
part thickened, fleshy. Lip 1-1.5 mm. long and 0.5-0.6 mm. broad, obscurely 3-
lobed, oblong or oblong-oval, basal part of the lip oblong, somewhat concave,
fleshy, terminating into two small, suberect lateral lobes, terminal lobe of the lip
about 0.5 mm. long, suborbicular, strongly concave, fleshy.
(166)
1946)
FLORA OF PANAMA (Orchidaceae) 61
Fig. 82. Stelis atrorubens
Endemic to Panama.
COCLE: vicinity of El Valle, alt. 600-1000 m., Allen 1234; hills north of El Valle de
Anton, trail to Las Minas, Allen 2876.
Allied to the preceding and the following species.
7. STELIS SkurcHm Ames in Bot. Mus. Leafl. Harv. Univ. 6:17, 2. 1938.
Caespitose epiphytic herbs up to 3 dm. tall. Secondary stems 6-10 cm. long,
slender, covered with scarious sheaths. Leaves petiolate; lamina 5-12 cm. long
and 0.8—2 cm. broad, elliptic, obtuse or acutish; petioles 2—4 cm. long, sulcate.
Inflorescence up to 22 cm. long, exceeding the subtending leaf; bracts about 2
mm. long, infundibuliform. Sepals connate at the base, papilliferous within;
dorsal sepal about 6 mm. long and 2 mm. broad, lanceolate-ovate, apiculate; lateral
sepals about 4.5 mm. long and 2 mm. broad, ovate-lanceolate, slightly oblique.
Petals about 1 mm. long and about 1.3 mm. broad, flabellate, callus-thickened
toward the truncate apex. Lip about 1 mm. long and 0.5 mm. broad, linguiform,
with an erect but obscure lobule on each side near the middle, callus-thickened
below, terminal part slightly concave above and slightly narrower than the basal
portion.
Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, Boquete District, alt. 6000 feet, Davidson 270.
(167)
ANNALS ОҒ THE MISSOURI BOTANICAL GARDEN
Fig. 83. Stelis Skutchii
(168)
[Vor. 33
19461
FLORA OF PANAMA (Orchidaceae) 63
8. SrELIS HYMENANTHA Schltr. in Fedde Rep. Sp. Nov. 10. 291. 1912.
Stelis cuspidilabia Schltr. in Fedde Rep. Sp. Nov. Beih. 19:17. 1923.
Stelis seleniglossa Schltr. loc. cit. 97
Small, caespitose, epiphytic herbs up to about 20 cm. long. Secondary stems
1-5 cm. long, shorter than the leaves. Leaves 4-10 cm. long and 0.2-1 cm. broad,
linear, ligulate to oblanceolate, obtuse or acute, coriaceous. Inflorescence up to
13 cm. long, longer than or subequal to the subtending leaves. Sepals 1-1.4 mm.
long and 0.75-1.2 mm. broad, broadly oval to suborbicular, obtuse or acute, free
almost to their bases. Petals about 0.5 mm. long and 0.25-0.3 mm. broad, oblong,
truncate, apex thickened. Lip 0.5-0.7 mm. long and about 0.4 mm. broad, sub-
quadrate to suborbicular-ovate, the apex produced into a broad, acute or acuminate
apicule about 14 the length of the lip, disc provided with a bipartite callus on
the basal half.
Mexico, Guatemala, Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, alt. 6000 feet, Terry 1286.
m
STELIS CRESCENTIICOLA Schltr. іп Fedde Rep. Sp. Nov. 16:442. 1920; Ames,
Sched. Orch. 5:11. 1923; Ames in Bot. Mus. Leafl. Harv. Univ. 3:175, t.
1933.
Stelis flexuosa Lindl., sensu Kränzl., in Engl. Bot. Jahrb. 26:450. 1899.
Stelis Isthmii Schltr. in Fedde 21 Sp. Nov. Beih. 17:16. 1922.
Stelis praemorsa Schltr. loc. cit. 17.
Small, caespitose, epiphytic herbs up to 20 cm. tall. Secondary stems 1-2 cm.
long, slender, much shorter than the leaves. Leaves 4-9.5 cm. long and 0.5-1 cm.
broad, oblanceolate, acute or obtuse, attenuated into a petiole at the base, coria-
ceous. Inflorescence up to about 18 cm. long, much exceeding the leaves, raceme
densely flowered; bracts 1-2 mm. long, infundibuliform. Sepals 1-1.5 mm. long
and 1-1.5 mm. broad, suborbicular to subrhombic-ovate, acute or obtuse, papilli-
ferous within. Petals about 0.5 mm. long and about 0.75 mm. broad, flabellate,
not thickened at the apex. Lip about 0.5 mm. long and 0.5 mm. broad and nearly
as thick, rhombic-ovate, with a transverse callus through the middle and the short
apex incurved.
Costa Rica and Panama.
CANAL ZONE (?): in tidal belt, Pittier 6505. PANAMA: Rio Chagres, Lehmann 4540;
hills east of [Panama] City and San Juan, all near sea-level, Powell 232, 234, 284, 285,
3312, 3313, 3314, 3522.
The smallest-flowered of the Panamanian species.
10. STELIS LEUCOPOGON Reichb. f. Beitr. Orch. Centr.-Am. 95, 7. 9, figs. 1, 1-4.
° 1866.
Stelis cascajalensis Ames, Sched. ОЧ 4 11. 1923.
Stelis eximia Ames, loc. cit. 6:54
(169)
64
ANNALS OF
S TELIS
THE MISSOURI BOTANICAL GARDEN
Ж
4 XS UNS oA t.
Fig. 84
(170)
[Vor. 33
19461
FLORA OF PANAMA (Orchidaceae) 65
Caespitose epiphytic herbs up to about 40 cm. tall. Secondary stems 2-15
cm. long, either longer or shorter than the leaves. Leaves 6-15 cm. long and
1-4.5 cm. broad, ligulate, oblanceolate to narrowly oval, acute or obtuse, coria-
ceous, attenuated into a petiole at the base. Inflorescence up to 30 cm. long, one
or more from the apex of each secondary stem, subequal to usually much exceeding
the leaves in length. Sepals 4-9 mm. long and 4-8 mm. broad, broadly ovate-
triangular to suborbicular, acute or obtuse, puberulent or glabrous within, usually
5-nerved. Petals 0.75-1.3 mm. long and 1.75-2.2 mm. broad, flabellate, truncate
apex very fleshy. Lip 0.5-1 mm. long and 0.7-1.2 mm. broad, transversely oblong
to suborbicular-quadrate, apiculate or triapiculate at the apex, disc very fleshy,
the callus more or less transverse.
Costa Rica and Panama; possibly also in South America.
CHIRIQUÍ: vicinity of Bajo Chorro, alt. 1900 m., Woodson t$ Schery 624; “СагатШа,”
alt. 4000-6000 feet, Powell 312, 334. сосі.Е: hills north of El Valle de Antón, alt. 1000
m., Allen 2173; vicinity of La Mesa, north of El Valle de Antón, alt. about 1000 m.,
Allen 2301; north rim, El Valle de Antón, Alston & Allen 1840.
A variable species which may include Stelis Endresii Reichb. f. Тһе specimens
from Coclé differ somewhat from the others in having a triapiculate apex on the
lip.
11. STELLIS ENpresu Reichb. f. іп Gard. Chron. 1373. 1870; Ames іп Bot. Mus.
Leafl. Harv. Univ. 1?: fig. p. 9. 1933.
Stelis parvibracteata Ames, Orch. 7:131. 1922.
Small, caespitose, epiphytic herbs up to 22 cm. таП. Secondary stems 1.5—5
cm. long, slender, shorter than the leaves.
Leaves 4—13 cm. long and 0.5-2 cm.
broad, elliptic to oblanceolate, obtuse or
acute, attenuated into a petiole at the base.
Inflorescence up to 18 cm. long, floriferous
to the middle or below, much exceeding
the leaves, 1 or more from a stem; bracts
1-2 mm. long, infundibuliform. Sepals 2—4
mm. long and 2-5 mm. broad, suborbicular
to subtriangular, acute or obtuse, glabrous
to puberulent within, 2-(or rarely 5-)
nerved. Petals 0.75—1 mm. long and 1-1.5
mm. broad, flabellate, truncate, apical part
thickened and fleshy. Lip about 0.75 long
and 1 mm. broad, transversely oval, with a
small, erect apicule at the apex.
Mexico, Guatemala, Costa Rica and
(171)
[Vor. 33
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN
em. possibly also in South America.
Cana and vicinity, alt. 6000 feet, Williams 072. сосіЖ: hills north of El
Valle өү рее alt. 800-1000 m.. Allen 2273.
Critical work on the genus may prove this species to be the same as S. leuco-
pogon Reichb. f
12. STELIS Рохкілл Schltr. in Fedde Rep. Sp. Nov. Већ. 17:16. 1922; Ames in
Bot. Mus. Leafl. Harv. Univ. 1°: fig. p. 0. 1933.
Small, caespitose, epiphytic herbs up to
STELIS Powellii Schltr. 15 cm. tall. Secondary stems 1-3 cm. long,
shorter than the leaves. Leaves attenuated
into a petiole at the base; lamina 2-7 cm.
long and 0.5—1.5 cm. broad, oblanceolate to
ligulate-oblanceolate, obtuse; petiole up to
3 cm. long. Inflorescence longer than the
leaves; bracts 2-4 mm. long, ovate, acute,
large and conspicuous for the genus, infundi-
buliform. Sepals 2.5-3 mm. long and 3-3.5
mm. broad, connate for a short distance at
the base, suborbicular to ovate-suborbicular
or ovate-triangular, obtuse, 5-nerved, puber-
ulent or papillose within. Petals 1-1.5 mm.
long and 1.5-2 mm. broad, flabellate, sub-
truncate and thickened terminally. Lip
ыы 1.2-1,5 mm. long and 0.75-1.2 mm. broad,
Fig. 86 ovate, concave, relatively thin with a cruci-
form callus in the middle.
Costa Rica and Panama.
CHIRIQUÍ: alt. 4000 feet, Powell 247.
Conspicuous among Panamanian species by the relatively conspicuous bracts.
13. STELIS Ѕтоккп Ames in Bot. Mus. Leafl. Harv. Univ. 3:54, fig. 1935; І.
Wms. in Ann. Missouri Bot. Gard. 27:273. 1940.
Small, caespitose, epiphytic herbs up to 6 cm. tall. Secondary stems 1-2 cm.
long, covered with sheathing bracts, shorter than the leaves. Leaves 1—4 cm.
long and 0.25—0.5 cm. broad, oblanceolate, acute or obtuse, inconspicuously
marginate, longer than the secondary stems, equal to or usually shorter than the
inflorescence. Inflorescence up to about 4 cm. long; bracts 1-1.5 mm. long,
infundibuliform, apiculate. Sepals connate for a short distance at their bases,
1-1.3 mm. long and 1—1.3 mm. broad, nearly orbicular, obtuse, 3-nerved. Petals
0.5—0.75 mm. long and about 0.6 mm. broad, broadly obovate or cuneate.
(172)
19461
FLORA OF PANAMA (Orchidaceae) 67
flabellate, slightly cochleate, the apex thick-
ened, Lip about 0.5-0.75 mm. long and
as broad, nearly orbicular, fleshy, with a
large central callus extending through the
middle of the lamina and prominently
raised at the base.
Costa Rica and Panama.
OCLÉ: vicinity of El Valle, alt. 800-1000
m., Allen 1828.
14. STELIS Махомп Schltr. in Већ. Bot.
Centralbl. 36, Abt. 2:389. 1918.
Caespitose epiphytic herbs up to 40 cm.
tall. Secondary stems up to 23 cm. long,
slender. Leaves 9-15 cm. long and 1-2.7
cm. broad, elliptic to lanceolate, obtuse,
coriaceous, attenuated into a short petiole at
the base. Inflorescence up to about 15 cm.
long, subequal to the leaves; bracts about 3
mm. long, infundibuliform, acute or shortly
acuminate. Sepals 2-3 mm. long and 2.2-
3.5 mm. broad, suborbicular. Petals about
1 mm. long and 1 mm. broad, suborbicular,
flabellate, the terminal part thickened and
fleshy. Lip about 1.2 mm. long and 1 mm.
broad, broadly oval in outline, with a trans-
verse callus across the middle of the lip, the
lip fleshy.
Costa Rica and Panama.
IRIQUÍ: Volcán de Chiriquí, alt. 10,000 STELIS «оғ Chnes
T "Devin 088; above El Boquete, alt. /
1450-1650 m., Maxon p southern slope of Fig. 87
La Horqueta, alt. 1700-2100 m., Pittier 3242
Closely allied to S. thecoglossa Reichb. f. with which it has been placed.
15. STELIS PANAMENSIS Schltr. in Већ. Bot. Centralbl. 37, Abt. 2:391. 1918.
Erect epiphytic herbs up to 18 cm. tall Secondary stems 2-3 cm. long, much
shorter than the leaves. Leaves 8—10 cm. long and 0.8—1 cm. broad, oblanceolate-
ligulate, obtuse, narrowed to a petiole at the base. Inflorescence equal to the
leaves or shorter; bracts cucullate. Sepals broadly ovate, obtuse; dorsal sepal
about 2.5 mm. long; the lateral sepals distinctly smaller, oblique. Petals broadly
rhombic-reniform, oblique, obtuse, apex thickened. Lip subequal to the petals,
reniform, obtuse, excavated at the base and the middle with a transverse callus.
(173)
[Vor. 33
68 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
Panama.
PANAMA: forests of the upper Mamoni River, alt. 150-400 m., РИНег 4490.
This species is known to us only by the description and an analysis of the type.
STELIS vestita Ames 16. STELIS VESTITA Ames, Sched. Orch. 6:56.
B 1923; in Bot. Mus. Leafl. Harv. Univ.
б; 1°: fig. р. 9. 1933.
Caespitose epiphytic herbs up to 45 cm.
tall. Secondary stems 4-10 cm. long, sub-
equal or shorter than the leaves. Leaves
5-16 cm. long and 1-2 cm. broad, elliptic-
oblong to oblanceolate, attenuated into a
petiole at the base, coriaceous. Inflorescence
up to about 35 cm. long, much longer than
the subtending leaf; bracts 2-3 mm. long,
infundibuliform. Sepals 2-2.5 mm. long
and 2-2.3 mm. broad, ovate to suborbicular,
obtuse, densely glandular-pubescent within.
Petals 0.8-1 mm. long and 1.5-1.75 mm.
broad, cuneate-flabellate, fleshy, truncate.
Lip 0.8-1 mm. long and 1-1.2 mm. broad,
suborbicular or transversely subrhombic,
callus-thickened in the middle and the basal part with a smaller spongy» callus,
the margins of the lip thin.
Costa Rica and Panama.
снікюоі: Boquete, alt. 4000 feet, Davidson боб.
Allied to S. Endresii Reichb. f. and to S. leucopogon Reichb. f.
17. STELIS AEMULA Schltr. in Већ. Bot. Centralbl. 36, Abt. 2:385. 1918.
Stelis sarcodantba Schltr. loc. cit. 392.
Small, caespitose, epiphytic herbs up to about 2.5 cm. tall. Secondary stems
1.5—7.5 cm. long, slender, shorter than the leaves. Leaves 4-11.5 cm. long and
0.5—1.3 cm. broad, ligulate to oblanceolate, obtuse or acute, coriaceous, attenuate
at the base into a petiole. Inflorescence up to 20 cm. long (in Panama mostly 12
cm. or less), much exceeding the leaves; bracts 1-2 mm. long, infundibuliform.
Sepals 1.5—2.2 mm. long and 1.75-2.5 mm. broad, broadly ovate-orbicular or
suborbicular, obtuse or acutish, marginate or the margins obscurely involute,
puberulent or glabrous within. Petals 0.5-0.6 mm. long and 0.6-1 mm. broad,
oblong to flabellate, the apex thickened. Lip 0.5—0.75 mm. long and 0.6-1 mm.
broad, suborbicular, obtuse, with a transverse callus in the middle, margins not
thickened.
(174)
19461 |
FLORA OF PANAMA (Orchidaceae) 69
(Guatemala, Honduras — $. perplexa Ames) ?, Costa Rica, and Panama.
CHIRIQUÍ: Palo Alto, alt. 4500-5000 feet, Powell 200, 201, 203, 204. сосіЁ: hills
north of E] Valle de Antón, alt. 1000 m., Allen 2154, 2157.
18. STELIs COLLINA Schltr. іп Fedde Rep. Sp. Nov. Beih. 17:15. 1922.
Small, caespitose, epiphytic herbs up to 28 cm. long. Secondary stems up to
3 cm. long, slender. Leaves 3-10 cm. long and 0.7-1.2 cm. broad, oblanceolate-
ligulate, obtuse, attenuated into a petiole at the base. Inflorescence elongated,
much exceeding the leaves; bracts about 2 mm. long, infundibuliform. Sepals
3-3.5 mm. long and 3—3.5 mm. broad, triangular-ovate, obtuse, 3-nerved. Petals
about 1 mm. long and 1.5 mm. broad, flabellate, truncate apex thickened. Lip
about 0.8 mm. long and 1 mm. broad, suborbicular, obtuse, with a transverse
raised callus in the middle, the basal half with two lateral cavities and a small
longitudinal ridge between them.
Panama.
PANAMA: foot-hills near [Panamá] City, near sea-level, Powell 180.
Formery referred to S. Williamsii Ames, but probably distinct, though closely
allied.
19. STELIS WirLiIAMsm Ames, Orch. 7:133. 1922.
Small, caespitose, epiphytic herbs up to 25 cm. tall. Secondary stems about
1.5 cm. long. Leaves 4-10 cm. long and 0.6-1.1 cm. broad, attenuated into a
short petiole at the base, oblanceolate-ligulate. Inflorescence up to 23 cm. long,
much exceeding the leaves; bracts 2-5 mm. long, infundibuliform, free part tri-
angular-lanceolate or broadly ovate, acute or acuminate. Sepals about 3.5—4.5
mm. long and as broad, suborbicular, obtuse. Petals about 1.5 mm. long and 1.7
mm. broad, cuneate-flabellate. Lip about 1 mm. long and 1.2 mm. broad, trulli-
form, obtuse, with a raised transverse callus.
Panama.
DARIÉN: Cana and vicinity, alt. 6000 feet, Williams 970.
Has been confused with S. collina Schltr., a closely allied sea-level species.
20. STELIS LONGIPETIOLATA Ames, Sched. Orch. 1:6. 1922.
Small, caespitose, epiphytic herbs up to 10 cm. tall. Leaves 4.5-8 cm. long and
0.7-1.1 cm. broad, elliptic, petiolate, the lamina up to 5.5 cm. long. Secondary
stems 1.5—2.5 cm. long, slender, much shorter than the leaves. Inflorescence
about equalling the leaves in length; bracts about 1.5 mm. long, infundibuliform.
Sepals 2-2.5 mm. long, rotate, deltoid-ovate. Petals about 1 mm. long, ovate to
suborbicular, thickened nearly to the base, abruptly excavated below the thicken-
ing, apex obscurely and minutely papillose. Lip about 0.75 mm. long and 1.5
mm. broad, reniform, very fleshy, somewhat concave above with a prominent and
obscurely bilobed callus in the middle, the sides of which are decurrent laterally
(175)
[Vor. 33
70 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and form a transverse ridge.
Panama.
cHIRIQUI: humid forest between Alto de las Palmas and Cerro de la Horqueta, alt.
2100-2200 m., Maxon 5460.
17. PHYSOSIPHON Lindl.
Рнүѕоѕрном Lindl. in Bot. Кер. 21: sub 7. 1797. 1836.
Small, caespitose, epiphytic herbs. Secondary stems slender, terminated by a
single leaf. Sepals connate at the base into an (usually) inflated tube, free above,
erect or spreading, not caudate. Petals very small in comparison to the sepals.
Lip small, articulated to the column-foot, entire or three-lobed. Anther terminal,
incumbent, operculate; pollinia 2, ceraceous.
A small genus of rather diverse species, of which the Panama one is not typical
of the genus.
1. Рнузоярном MINUTIFLORUS Ames & Schweinf. Sched. Orch. 8:11, 7. 22, figs.
1-6 and habit. 1925.
Small caespitose epiphytic herbs up to 30 cm. tall. Secondary stems 2-10 cm.
long, invested by 2-3 loose sheaths. Leaves 3.5-12 cm. long and 0.5-1.5 cm.
broad, oblong-ligulate, coriaceous, attenuated at the base into a petiole. Inflores-
cence up to 25 cm. long, equalling or exceeding the leaves, densely flowered almost
to the base; bracts 2-3 mm. long, infundibuliform. Sepals connate into a short
tube at the base, 2-3 mm. long and about 0.6-0.8 mm. broad, free portion ovate-
lanceolate, acute or obtuse. Petals about 0.75 mm. long and about 0.2 mm. broad,
lanceolate, acute or obtuse. Lip about 0.75 mm. long and 0.75 mm. broad,
ovate-quadrate to suborbicular, apiculate.
Costa Rica and Panama; possibly Honduras.
CHIRIQUÍ: Cerro Vaca, alt. 900-1136 m., Pittier 5340; "Caramillo," alt. 5000 feet,
Powell 345.
18. CRYPTOPHORANTHUS Barb. Rodr.
Cr¥PTOPHORANTHUS Barb. Rodr. Gen. & Sp. Orch. Nov. 2:79. 1882; Kranzl. in
Fedde Rep. Sp. Nov. Beih. 34:220-232. 1925.
Small epiphytic herbs, the secondary stem terminated by a single leaf. Іп-
florescence short, 1-flowered or with a fascicle of flowers. Sepals connate at the
base and at the apices, leaving an opening ("window") on either side between the
dorsal and lateral sepals, the lateral sepals forming a mentum at the base. Petals
small in comparison to the sepals. Lip free, simple, obscurely lobed or hastate,
with two parallel lamellae, subequal to the petals in length. Column small, terete,
erect or arcuate, produced into a short foot at the base. Anther terminal, opercu-
late, incumbent; pollinia 2, ceraceous.
(176)
19461
FLORA OF PANAMA (Orchidaceae)
Fig. 89. Physosiphon minutiflorus
A small genus closely allied to Masdevallia and Pleurothallis.
or four species in Costa Rica and one in Panama.
(177)
71
There are three
[Vor. 33
ya ANNALS OF THE MISSOURI BOTANICAL GARDEN
1. CRYPTOPHORANTHUS LEPIDOTUS L. Wms. in Ann. Missouri Bot. Gard. 29:340,
і. 30, figs. 4-5. 1942.
Caespitose epiphytic herbs up to about 17 cm. tall. Secondary stems 2-7 cm.
long and 1-2 mm. in diameter, covered with 4-5 pergameneous, infundibuliform
sheaths which soon disentegrate. Leaves oblanceolate to narrowly obovate, obtuse
or acutish, coriaceous, contracted into a distinct petiole; lamina 3.5-10 cm. long;
petiole 1-2 cm. long, conduplicate. Inflorescence consisting of 1-6 long-peduncu-
late flowers at the apex of the secondary stem; the peduncle with 1—3 short
infundibuliform sheaths. Sepals joined together at the base and at the tip, leaving
Fig. 90. Cryptopborantbus lepidotus
a small opening between the dorsal and lateral ones; dorsal вера! about 15-20 mm.
long and 6—7 mm. broad, oblong-lanceolate, strongly cucullate, fleshy, 7-nerved,
ridged dorsally and the ridges verrucose; lateral sepals connate to their apices,
about 12-18 mm. long and together 6-8 mm. broad, fleshy, each about 7-nerved,
with verrucose ridges dorsally. Petals 5-6 mm. long and 2.5-3.5 mm. broad,
broadly ovate-lanceolate, acute or acuminate, 3-(5-)nerved. Lip 5-6 mm. long,
hastate, unguiculate; lamina about 4 mm. long, 1.5-2 mm. broad, 3-nerved,
verrucose or lepidote, with two longitudinal, lamellate calluses extending from
the auricles to about the middle; auricles about 1 mm. long, retrorse, subulate;
claw 1.5-2 mm. long and 1—1.5 mm. broad, verrucose-scurfy or lepidote toward
(178)
19461
FLORA OF PANAMA (Orchidaceae) 73
its apex. Column of the genus.
COCLÉ: trail to Las Minas, hills north of El Valle de Antón, alt. about 1000 m., Allen
2718.
19. MASDEVALLIA Ruiz & Pavon
MaspDEVALLIA Ruiz & Pavon, Fl. Peruv. & Chil. Prodr. ed. 1, 122. 1794; ed. 2,
110, Ё. 27. 1797; Woolward, Genus Masdevallia. 1896; Kranzl. in Fedde
Rep. Sp. Nov. Beih. 34:1-202. 1925.
Caespitose or repent, epiphytic or terrestrial herbs. Secondary stems usually
very short, bearing a single leaf and a peduncle or the leaves and peduncles borne
on separate stems. Leaves most often broadest toward the apex and attenuated
into a petiole at the base, with an abscission at the end of the secondary stem and
another slightly above. Peduncles usually 1- to 2-flowered. Sepals connate into
a tube at the base or to the middle or beyond, the free portions often spreading,
the tips usually developed into slender tails, produced into a mentum at the base.
Petals very much smaller than the sepals, usually narrow. Lip small, articulated
to the column-foot, about as long as the column. Column erect, produced into
a column-foot at the base; margins of the clinandrium usually membranaceous
and denticulate. Anther terminal, operculate, incumbent; pollinia 2, ceraceous.
A large and complex genus of some 250 species. The species are most ab andant
in coo! mountain regions of the tropics of the New World. The genus was once
very popular with orchid fanciers, some of the rarer or more beautiful species
commanding fabulous prices.
Masdevallia connects to the genus Pleurothallis at several points, notably
through such species as М. Allenii L. Wms., which differs but slightly from the
species of Schlechter's Barbosella (previously reduced to Pleurothallis). Scapho-
sepalum has several species which are "connecting links" between Мазде а а and
Pleurothallis. Lindley’s genus Restrepia, which I have reduced to Pleurothallis,
contains some intermediate species between Pleurothallis and Masdevallia, and if
Restrepia were to be maintained as a genus most of the species which were re-
ferred to Barbosella and many of the species of Masdevallia § Triaristella (M.
triaristella Reichb. f., M. Allenii L. Wms., etc.) should be placed there. There is
little or no technical reason why Pleurothallis (and Stelis too) should not be re-
duced to Masdevallia, but it would serve no practical purpose to do so. Pleuro-
thallis and Masdevallia are both large and technical and most of the species can
be placed in the proper genus readily.
a Tails of the lateral sepals preme: below the apex 8. M. ALLENII
a. Tails of the lateral als, if present, terminal.
b. Lateral — with transverse pr des at the base 7. M. SIMULA
bb. Lateral sepals wi transverse calluses at the bas
с: кер = the base of the petal free and appearing to be a салда... 1. M. COLLINA
сс. Callus lacking at 52. base of the petal, or if present not free
d. Lip less than 2.5 . long 6. M. TENUISSIMA
dd. Lip more than El mm. long.
(179)
[Vor. 33
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN
е. Du sepal prolonged into a subfiliform tail.
Peduncle қ. ri - uam Law: as long as the leaves or
ыдыс» er; lip up t 3. M. CHONTALENSIS
ff. iva lowered em shorter than the leaves; lip
than 4 2. M. ATTENUATA
ee. Doral sepal either ore i tails, or with tails not filif
f. am t of the dorsal sepal lanceolate or edit
. M. ECAUDATA
ff. Free part of the dorsal sepal ligulate or lanceolate-ligulate,
recurved 5. M. LivINGSTONEANA
1. MASDEVALLIA COLLINA L. Wms. in Am. Orchid Soc. Bull. 11:93, 7. 3. 1942.
Small caespitose epiphytic herbs up to 15 cm. tall. Secondary stems very
short, bearing a leaf and ап inflorescence. Leaves 6-9 cm. long and 1.1-1.5 cm
broad, oblanceolate, obtuse, attenuated into a petiole at the base, coriaceous, 3-
nerved. Inflorescence slender, (1?—)2-flowered, peduncle much exceeding the
leaves. Sepals about 25 mm. long, connate into a tube for about У; their length;
tube 11-14 mm. long, free part of the sepals 9-14 mm. long, consisting of sub-
filiform caudae from a very short free lamina. Petals about 4 mm. long and 1.5
mm. broad, linear-oblong, truncate and tridentate at the apex, 1-nerved, with
an elongated, sublamellate callus which extends from near the base to near the
apex along the anterior margin; another callus begins near the base and extends
to and off at the base forming a free, curved, cauda-like process, the free part
about 1 mm. long. Lip about 6 mm. long and 2 mm. broad, linear-oblong, trun-
cate and retuse at the base, the apex rounded, somewhat dilated and ciliate, fleshy,
somewhat canaliculate, keeled on the mid-nerve below.
Panama.
COCLÉ: hills north of El Valle de Antón, alt. about 1000 m., Allen 2158.
2. MASDEVALLIA ATTENUATA Reichb. f. іл Gard. Chron. 834. 1871; Woolw.
Genus Masdevallia 7. [35]. 1896; Hook. f. in Bot. Mag. 33: £. 6273. 1877.
Masdevallia Laucheana vier Pda ex Woolw. loc. cit. 7. [19]; Kránzl. іп Fedde Rep.
Sp. Nov. Beih. 34:166.
Small, caespitose, epiphytic herbs up to about 15 cm. tall. Secondary stems
very short. Leaves 5-13 cm. long and 0.5—1.8 cm. broad, linear-oblanceolate to
oblanceolate, acute or obtuse, attenuated into a petiole at the base. Inflorescence
1-flowered; peduncles slender, shorter than the leaves. Sepals 18-22 mm. long,
connate into a tube for about № their length, the caudate apices about as long as
the broad part of the sepals, puberulent within, the broad, free part of the sepals
oblong-ovate to triangular-ovate. Petals 3.5—5 mm. long and 1.2 mm. broad,
subrhombic to oblong-lanceolate, acute, somewhat oblique, with a longitudinal
callus along one margin. Lip 4.2-5 mm. long and 1.5-2 mm. broad, oblong-
lanceolate to oblong-pandurate, obtuse or acute, apical portion denticulate, with
two longitudinal keels which converge toward the apex.
Costa Rica and Panama.
CHIRIQUÍ: Boquete, alt. about 1360 m., Davidson 1271; “Chiriqui,” Pfau; "Cerro
Horconcito," alt. about 1360 m., Powell 268.
(180)
19461
| FLORA OF PANAMA (Orchidaceae) 75
SS
са У US
И
Ж fy A м NN 4
Fig. 91. Masdevallia collina
(181)
[Vor. 33
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Possibly the original collection of this species was from Chiriqui (Pfau),
Powell’s collection differs slightly from the others. The species is rare and more
specimens are desired for study.
3. MASDEVALLIA CHONTALENSIS Reichb. f. іп Otia Bot. Hamb. 1:274. 1878; L.
Wms. in Ann. Missouri Bot. Gard. 27:274. 1940.
Masdevallia diantha Schltr. in Вей. Bot. Centralbl. 36, Abt. 2:384. 1918.
Small, caespitose, epiphytic herbs up to about 11 cm. tall. Secondary stems
up to about 2 mm. long. Leaves 2-9 cm. long and 0.2-0.8 cm. broad, oblanceo-
late to linear-oblanceolate, acute or obtuse, attenuated into a petiole at the base.
Inflorescence 1- or usually 2-flowered; peduncles from almost as long as the leaves
to twice as long. Sepals 9-18 mm. long, connate into a tube for about № or 35
their length, the free part ovate to ovate-lanceolate and caudate. Petals 2.5—3.5
mm. long and 0.7-1.2 mm. broad, linear-oblong to oblong-lanceolate, apex tri-
dentate, truncate or obtuse, with a small lateral callus extending from about the
middle toward the apex. Lip 2.8-4 mm. long and 0.9-1.7 mm. broad, oblong-
oblanceolate to oblong-obovate, biauriculate at the base, with two longitudinal
callus-ridges extending from the base nearly to the apex where they converge,
dilated apical part denticulate.
Nicaragua, Costa Rica and Panama.
ocLÉ: vicinity of El Valle, alt. 600-1000 m., Allen 1231; vicinity of El IE north
rim, alt. 800-1000 m., Allen 1826; trail to Las и, El Valle de Antón, alt.
Allen 2876.
4. MASDEVALLIA ECAUDATA Schltr. in Beih. Bot. Centralbl. 36, Abt. 2:384. 1918;
L. Wms. in Ann. Missouri Bot. Gard. 27:274. 1940
Small, caespitose, epiphytic herbs up to about 12 cm. tall. Secondary stem
short, 2-10 mm. long. Leaves 3-9 cm. long and 0.6—1.1 cm. broad, oblanceolate,
obtuse, coriaceous, attenuated into a petiole at the base. Inflorescence about as
long as or exceeding the leaves, 1-flowered, the peduncle slender. Sepals puberulent
within, about 15-22 mm. long, connate into а tube about 24 of their length, the
free parts lanceolate to ovate-lanceolate, acute or acuminate, usually ecaudate but
sometimes caudate, the apices thickened. Petals 5-8 mm. long and 1.5—3.5 mm.
broad, oblong-ovate, short-unguiculate, usually angled on one side. Lip 6.5-9
mm. long and 2-3.5 mm. broad, oblong-lanceolate, auriculate at the base, disc
with a pair of longitudinal fleshy calli.
Costa Rica and Panama.
cHiRIQUÍ: valley of the upper Río Chiriqui Viejo, White 9 White 81.
Very closely allied to M. Livingstoneana, from which it differs mainly in having
slightly different sepals and petals and a larger lip.
5. MaASDEVALLIA LIvINGSTONEANA Reichb. f. іп Gard. Chron. II, 2:322. 1874;
Ames, Sched. Orch. 5:9, fig. 2. 1923.
Scaphosepalum panamense Schltr. in Fedde Rep. Sp. е 12:205. 1913.
Masdevallia panamensis Ames, Sched. Orch. 4:9.
(182)
19461
FLORA OF PANAMA (Orchidaceae)
М КІЛТ
/ dj T V |
UM Ју
/, Й
2 | : у
PAIN
Fig. 92. Masdevallia Livingstoneana
Small, caespitose, epiphytic herbs up to 13 cm. tall. Secondary stems short.
Inflorescence 1-flowered, peduncle shorter than the leaves. Sepals 15-20 mm.
long, connate into a tube for about У; their length; free portion of dorsal sepal
up to 12 mm. long and 3 mm. broad at the base, ligulate or lanceolate-ligulate,
(183)
77
[ VoL. 33
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN .
somewhat thickened and fleshy toward the apex; free part of the lateral sepals up
to 12 mm. long and 5 mm. broad, lanceolate-triangular, arcuate, acute or obtuse.
Petals 4.5-5 mm. long and 1.5-2 mm. broad, oblong, obtuse, with a single longi-
tudinal sublamellate callus extending from below the middle to the apex. Lip
4-6 mm. long and 1-1.2 mm. broad, oblong to oblong-lanceolate, acute or obtuse,
sometimes more or less pandurate, the disc thickened and with two raised, longi-
tudinal calli.
Costa Rica (?) and Panama.
CANAL ZONE: forest along Río Indio de Gatun, alt. near sea-level, Pittier 2803; west
of canal near village of Arrajan, alt. near sea-level, Powe 3. PANAMA: foothills east
of Panama City and San Juan, alt. near sea-level, Powell 273, 411, 3282, 3287.
6. MASDEVALLIA TENUISSIMA С. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 4:111.
1997.
Small, caespitose, epiphytic herbs up to about 3 cm. tall. Secondary stems
very short, up to about 1 mm. long. Leaves 1-2.2 cm. long and 1-2.5 mm.
broad, linear-oblanceolate, obtuse or acute, attenuated into a narrow petiole at the
base. Inflorescence up to 3 cm. long, exceeding the leaves; scape filiform. Sepals
connate into a tube at the base; dorsal sepal about 6 mm. long and 2.5 mm. broad,
caudate, the lamina about 2.5 mm. long and as broad, suborbicular. Lateral sepals
about 7.5 mm. long and each about 1 mm. broad, caudate, free part of lamina
lanceolate. Petals about 2 mm. long and about 0.75 mm. broad, oblong-lanceolate,
subtruncate, triapiculate. Lip about 2 mm. long and 0.7 mm. broad, oblong-
lanceolate, obtuse, with 2 inconspicuous, submarginal, lamellate calli extending
from the base to the middle, shortly bicaudate at the base, the caudae about 0.2
mm. long. Ovary inconspicuously muricate on the angles.
Panama.
СОСІ.Е: mountains beyond La Pintada, alt. 400-600 m., Hunter & Allen 587.
Masdevallia pygmaea Kránzl. will possibly include this species when the type
can be studied
Fig. 93. Masdevallia simula
(184)
19461
FLORA OF PANAMA (Orchidaceae) 79
MASDEVALLIA SIMULA Reichb. f. in Gard. Chron. n. s. 3:8. 1875; Woolw.
Genus Masdevallia, #. [71]. 1890; L. Wms.’ in Ann. Missouri Bot. Gard.
27:274, pl. 31, figs. 1-8. 1940.
Small caespitose or repent epiphytic herbs up to about 9 cm. tall. Secondary
stems up to about 1 mm. long. Leaves 1-10 cm. long and 0.2—0.5 cm. broad,
linear to oblanceolate, attenuated into a petiole at the base. Inflorescences 1—3-
24
flowered, much shorter than the leaves. Sepals connate for only a short distance
at the base; dorsal вера! 3.5-9 mm. long and 2-4 mm. broad, lanceolate to ovate-
lanceolate or triangular-lanceolate, acute, acuminate or caudate; lateral sepals
similar to the dorsal except usually somewhat falcate and each provided with a
transverse, semi-lunate callus at the base. Petals 1.5-3 mm. long and 1.5—3 mm.
broad, subquadrate or subrhombic, angulate. Lip 2-3.5 mm. long and 1.2-3
mm. broad, unguiculate; lamina subquadrate to subquadrate-obovate, biauriculate
at the base, with two lamellate calli near the junction with the claw.
Guatemala, Honduras, Costa Rica, Panama, Colombia, and Ecuador.
CANAL ZONE: in tops of high trees, Quebrada Lopez, alt. about 30 m., Allen 2115.
Masdevallia simula is quite a diverse species
in Central America and may prove to be either
an aggregate or a polymorphic. It is not a
typical Masdevallia and might almost as well
be placed іп Pleurothallis. The forms М.
guatemalensis Schltr. and M. linearifolia Ames
are included in the description.
8. MASDEVALLIA ALLENI L. Wms. in Ann.
Missouri Bot. Gard. 27:273, pl. 31, figs. 12-
16. 1940.
Small caespitose epiphytic herbs up to about
5 cm. tall. Secondary stems about 3 mm. long,
covered with white chartaceous sheaths. Leaves
10-20 mm. long and up to 2 mm. broad,
linear, acute, coriaceous, subterete and canal-
iculate. Inflorescence 1-flowered, longer than
the leaves; peduncle about 3—4 cm. long. Sepals
forming a short tube at the base; lamina of
the dorsal sepal 2.5—3 mm. long and 3-3.5 mm.
broad, free portion of the lamina triangular,
long-caudate, the cauda filiform and about 10
mm. long; lateral sepals 12-15 mm. long, con-
nate to their apices or nearly so, the lamina
lanceolate, with a lateral, filiform appendage
about 5-6 mm. long near the apex on either
Fig. 94. Masdevallia Allenii
(185)
[Vor. 33
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN
side. Petals 1.5-2 mm. long and about 0.75 mm. broad, oblong, obtuse. Lip
about 3 mm. long and 1 mm. broad, lanceolate or lanceolate-oblong, obtuse, bi-
caudate at the base; the lamina with two longitudinal calli.
Panama.
cocLÉ: vicinity of El Valle, alt. 600-1000 m., Allen 1230.
20. SCAPHOSEPALUM Pfizer
Scar UM Pfitz. in Engler & Prantl, Nat. Pflanzenfam. 29:139. 1888;
Kränzl. in Fedde Rep. Sp. Nov. Већ. 34:204-219. 1925
Caespitose or repent epiphytic or terrestrial herbs. Secondary stems very short,
unifoliate. Leaves mostly broadest near the apex and attenuated into a petiole at
the base. Inflorescence а distichous raceme. Sepals connate at the bases; the
dorsal sepal free or nearly so, sometimes caudate; lateral sepals more or less united
usually forming a cymbiform synsepal, often callus-thickened near the apices, the
apices often attenuated into tails. Petals much shorter than the sepals. Lip
small, articulated to the column-foot. Column erect, produced into a foot at
the base; margins of the clinandrium membranaceous and denticulate. Anther
terminal, operculate, incumbent; pollinia 2, ceraceous.
Scaphosepalum is hardly distinct frora Masdevallia. The one Panamanian
species might well be placed in Pleurothallis.
1. SCAPHOSEPALUM ELASMOTOPUS Schltr. in Fedde Rep. Sp. Nov. 12:204. 1913.
Scaphosepalum longirepens Ames in Proc. Biol. Soc. Wash. 34:155. 1921.
Small, repent, epiphytic herbs. Rhizome elongated. Secondary stems short,
up to about 5 mm. long, bearing a leaf or a leaf and a peduncle, with an addi-
tional abscission layer in the petiole above the apex of the secondary stem. Leaves
7.5-15 cm. long and 0.7-1.4 cm. broad, oblanceolate, acute, coriaceous, attenuated
into a slender petiole at the base. Inflorescence about as long as or shorter than
the leaves; raceme distichous, several-flowered, up to about 6 cm. long. Dorsal
sepal free nearly to the base, about 5-8 mm. long, obtuse, oblong-ligulate, the
apex thickened. Lateral sepals joined nearly to their apices, 6-10 mm. long, the
lamina oblong, short-caudate at the apex. Petals 3-3.5 mm. long, elliptic-oblong
to elliptic-ovate, oblique, acute. Lip about 3 mm. long, short-unguiculate; lamina
oblong-pandurate or ovate-oblong and constricted toward the middle, the terminal
lobe rounded, ciliate.
Panama.
CHIRIQUÍ: east of Rio Caldera, alt. 2. га., Killip 3567; between Alto de las Palmas
and top of Cerro de la Horqueta, alt. 2100-2268 m., Pittier 3220, 5730. CANAL ZONE:
Quebrada Lopez, alt. 30 m., Allen 2 et
This species has been reduced to Scaphosepalum macrodactylon Rolfe by
Kranzlin. Kranzlin probably did not see material from Panama.
(186)
1946)
FLORA OF PANAMA (Orchidaceae) 81
21. LEPANTHES Sw.
LEPANTHES Sw. in Nova Acta Soc. Sci. Upsal. 6:85. 1799; in Svensk. Vet-Akad.
Handl. 21:249. 1800.
Small repent or caespitose epiphytic herbs. Stems (secondary stems) generally
unifoliate, with several loose, often ciliate and pubescent, sheaths below. Leaves
coriaceous. Inflorescence terminal, 1 or few from the base of the leaves, the
flowers small, often distichous. Sepals subequal, often more or less connate at
the bases or the laterals connate nearly to their apices. Petals shorter than the
sepals, bipartite or transversely bilobed, usually short-unguiculate and adnate to
the column, except L. eximia. Lip bilobed or sometimes trilobed, with the mid-
lobe small and inconspicuous, adnate to the column, the lobes usually appearing
as wings. Column short, footless, wingless, dilated at the apex; anther terminal,
operculate, incumbent; pollinia 2, waxy.
An easily distinguished genus with many closely allied and poorte defined
species.
a. Petals simple, not bilobed and transverse 8. L. EXIMIA
aa. Petals not simple, bilobed and transverse.
b. Sepals linear-lanceolate, at least 5 times longer than broad................ 7. L. INSECTIFLORA
bb. Sepals broader than linear-lanceolate, less than 5 times longer than
broad.
c. Anterior lobe > the petal as long ав the dorsal 5 5. L. ROTUNDIFOLIA
сс. а obe the абл Beet re ve the Mind. x
d. sal sepal ү than m. lon
e. езд 3-nerved 6. L. TURIALVAE
ee. Petals 1-nerved 1. L. CHIRIQUENSIS
dd. Dorsal sepal 4 mm. long or longer.
f. Petal 1-nerved; dorsal sepal 14—17 mm. long 4. L. MaxoNm
ff. Petals 3-(5-)nerved; dorsal вера! 4—8 mm. long.
в. Dorsal вера! about 4-6 mm. long, ovate 3. L. ЕСПЛАТА
gg. Dorsal siam about 8 mm. long, Іапсеојасе........................ 2. L. ELATA
1. LEPANTHES CHIRIQUENSIS Schltr. іп Fedde Rep. Sp. Nov. Већ. 17:17. 1922.
Lepantbes micrantba Ames, Sched. Orch. 4:31. 1923.
Small, caespitose, epiphytic herbs up to 10 cm. tall. Secondary stems slender,
up to about 8 mm. long. Leaves 1—4 cm. long and 0.3—1.1 cm. broad, variable,
linear-lanceolate to ovate. Inflorescence few-flowered, shorter than the leaves.
Dorsal sepal 2-2.5 mm. long and 1.2-1.5 mm. broad, oblong-ovate to lanceolate-
ovate, acute. Lateral sepals 1.5-2 mm. long and 0.75—1.2 mm. broad, ovate-
lanceolate, acute, arcuate, sometimes denticulate toward the apex. Petals 0.5-0.7
mm. long and 2—2.5 mm. broad, transverse, ciliate, the lobes lanceolate to ovate-
lanceolate. Lip 0.75-1.2 mm. long and 0.75-1 mm. broad, 3-lobed; the lateral
lobes malleoliform, about 0.75—1.2 mm. long; mid-lobe small, apiculiform, about
0.2 mm. long.
Costa Rica and Panama.
RIQUÍ: vicinity of Monte Lirio, valley of upper Río Chiriqui 524 alt. 1300-1900
m., Seibert 174, 104, 198; "Province of Chiriqui,” alt. 4500 ft., Powell 2
(187)
[Vor. 33
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Lepanthes chiriquensis has been referred to L. Lindleyana Oerst. & Reichb. f.
as a synonym, which it may be, but the original description and figures of L.
Lindleyana are either very inaccurate or the species is quite different.
2. LEPANTHES ELATA Reichb. f. Beitr. Orch. Centr.-Am. 90. 1866.
Rather large, caespitose, epiphytic herbs up to 6 dm. tall. Secondary stems
up to 5 dm. long, covered with scarious sheaths which are glabrous or at most
ciliate. Leaves 3.5-12 cm. long and 1.8-7 cm. broad, lanceolate-ovate to oval,
acuminate. Inflorescences 1 or more from the axils of each leaf, densely flowered;
bracts very crowded, ovate-acuminate, infundibuliform. Dorsal sepal about 8 mm.
long and 3 mm. broad, lanceolate, acuminate. Lateral sepals about 8 mm. long
and 3 mm. broad, lanceolate, acuminate, somewhat oblique, connate nearly to the
middle. Petals transverse, about 5 mm. broad; the posterior lobe about 2.5 mm.
long, oblong-obovate; the anterior lobe about 2.5 mm. long and about 2 mm.
broad, the lateral lobes malleoliform, the mid-lobe small, apiculiform.
Costa Rica and Panama.
cHiRIQUÍ: Bajo Chorro, alt. 6000 ft., Davidson 237.
3. LEPANTHES ECILIATA Schltr. in Fedde Rep. Sp. Nov. 12:203. 1913.
Erect, caespitose, epiphytic herbs up to about 18 cm. tall. Secondary stems
slender, covered with several sheaths, the mouths of which are eciliate but puberu-
lous. Leaves 3.5—7.5 cm. long and 0.8—1.8 cm. broad, elliptic or lanceolate, erect,
attenuated to the base. Inflorescences several, about half as long as the leaves.
Sepals ovate, ciliate; dorsal sepals about 4—6 mm. long and about 3.5 mm. broad,
short-acuminate; lateral sepals about 4—5 mm. long and together about 3.5 mm.
broad, acuminate, oblique, connate to about the middle. Petals about 4 mm.
broad, transverse, glabrous; anterior lobe oblong, oblique, obtuse; posterior lobe
narrowly oblong, obtuse, oblique. Lip about 1-1.5 mm. long, bipartite and with
an apicule in the sinus, lobes semirhombic, obtuse, glabrous.
Panama.
cHIRIQUÍ: Bajo Chorro, alt. about 1800 m., Davidson 128 in part; Cerro de la Hor-
queta, alt. 1700 m., Pittier 3173.
4. LEPANTHES Махомп Schltr. in Fedde Rep. Sp. Nov. 12:204. 1913.
Small, caespitose, epiphytic herbs up to about 15 cm. tall Secondary stems
up to about 9 cm. long, covered with several pubescent or scabrous infundibuli-
form bracts. Leaves 3.5—5.5 cm. long and 1.5—2.5 cm. broad, ovate, acuminate,
apex tridenticulate. Inflorescence distichous, fractiflex, exceeding the leaves;
flowers large for the genus. Sepals connate at the base; dorsal sepal 14—17 mm.
long and about 4—5 mm. broad, triangular-lanceolate or lanceolate, long-acuminate;
lateral sepals connate to the middle or beyond, 12-17 mm. long and together 5-6
mm. broad, broadly lanceolate, acute or acuminate. Petals transverse, about 3—4
mm. broad; anterior lobe narrowly oblong to lanceolate, arcuate, acute or obtuse,
about equal to the posterior lobe in length; posterior lobe narrowly lanceolate,
(188)
19461
FLORA OF PANAMA (Orchidaceae) 83
arcuate, acute. Lip about 1 mm. long and 1 mm. broad, bipartite, the lobes sub-
rhombic.
Panama.
CHIRIQUÍ: between Alto de las Palmas and Cerro do la Horqueta, alt. 2100-2268 m.,
Maxon 3256; same locality, Pittier 5404
5. LEPANTHES ROTUNDIFOLIA L.W ms.
in Ann. Missouri Bot. Gard. 27:
275, pl. 31, figs. 0-11. 1940.
Small, caespitose, epiphytic herbs
up to about 8 cm. tall. Secondary
stems slender, covered with sheaths
orbicular-ovate, or even broader than
long, coriaceous. Inflorescence dis-
tichous, shorter than the leaves. Dorsal
sepal about 3 mm. long and 2 mm.
broad, suborbicular, abruptly acumi-
nate. Lateral sepals about 2.5 mm.
long and 2 mm. broad, suborbicular,
obtuse or acutish, connate ас their
bases. Petals bipartite; lobes near the
dorsal sepal longest, about 3.5 mm.
long and 1 mm. broad, lanceolate,
acute, oblique; lobes near the lateral
sepals about 2 mm. long and 0.8 mm.
broad, lanceolate, acute, oblique. Lip
bipartite, about 1.5 mm. long and 2
mm. broad, the lobes malleoliform.
Panama.
rth rim, vicinity of El Fig. 95. Lepanthes rotundifolia
Valle, ale, '800— 1000 m., Allen 1835.
6. LEPANTHES TURIALVAE Reichb. f. in Bonplandia 3:225. 1855; Xenia Orch.
1:151, £. 50, figs. V, 15-16. 1855.
Small epiphytic herbs up to 15 cm. tall. Secondary stems slender, covered
with sheaths which are usually dark-colored. Leaves 1.5—4.5 cm. long and 0.9—2
cm. broad, elliptic-oblong to suborbicular, attenuated into a short petiole at the
base. Sepals lanceolate to ovate-lanceolate, acute; dorsal вера! about 2.5—3.5 mm.
long and 1.5-2.2 mm. broad; lateral sepals about 2.273 mm. long and 1-1.5 mm.
broad, connate to the middle or beyond. Petals about 0.5-1.5 mm. long and 3-4
mm. broad, transverse, posterior lobe the larger. Lip about 1-1.5 mm. long and
1.5-2 mm. broad, trilobate; the lateral lobes large and. peltate or flattened; mid-
lobe apiculiform.
(189)
[Vor. 33
84 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico to Panama.
CHIRIQUÍ: vicinity of Casita Alta, alt. 1500-2000 m., Woodson, Allen & Seibert 054.
COCLÉ: vicinity of El Valle, alt. 600-1000 m., Allen 1241; north of El Valle de Antón,
alt. 800—1000 m., Allen 2265.
The description is based on Panamanian material only. Lepanthes turialvae, as
now delimited, is possibly an aggregate of several species.
7. LEPANTHES INSECTIFLORA C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 7:152.
1939.
Small, caespitose, epiphytic herbs up to about 9 cm. tall. Secondary stems
slender, 2.5-7.5 cm. long. Leaves up to 2.5 cm. long and 0.9 mm. broad, elliptic-
oval to subrhombic, apex tridenticulate. Inflorescence several-flowered, shorter
than the leaves. Sepals about 5 mm. long and 0.5-1 mm. broad, linear-lanceolate,
acuminate. Petals transversely bipartite; posterior lobe about 3 mm. long, sub-
filiform; anterior lobe about 0.7 mm. long, subovate. Lip about 1 mm. long and
as broad, suborbicular, deeply emarginate and bilobed, with a minute apicula in
the sinus.
Panama.
CHIRIQUÍ: Bajo Chorro, alt. about 1800 m., Davidson 185.
Differs from L. tipulifera Reichb. f. only in having the anterior lobe of the
petal shorter. Material seen inadequate.
8. LEPANTHES ЕХІМІА Ames, Sched. Orch. 5:21. 1923.
Lepanthes abnormis Schltr. in Fedde Rep. Sp. Nov. Beih. 19:21. 1923.
Small, caespitose, epiphytic herbs up to 6 cm. tall. Secondary stems very
slender, up to 4 cm. long, covered with infundibuliform sheaths. Leaves 0.6-2
cm. long, 0.4-0.7 cm. broad, elliptic to oval, acute or obtuse, attenuated to the
base. Sepals 2-3 mm. long and 1-1.5 mm. broad, ovate-lanceolate, acute or
acuminate, denticulate. Petals 1-1.5 mm. long and 0.2-0.4 mm. broad, linear,
auriculate on the anterior margin at the base, not transverse. Lip 1-1.4 mm.
long and 0.6-0.8 mm. broad, oblong to subquadrate, short-unguiculate, retuse
or emarginate, ecallose.
Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, alt. about 1800 m., Davidson 186.
Unusual in having petals which are not transverse.
22. ACOSTAEA Schltr.
AcOSTAEA Schltr. in Fedde Rep. Sp. Nov. Већ. 19:283. 1923.
Small, caespitose, epiphytic herbs. Secondary stems very short, unifoliate,
covered with a loose sheath. Leaves subcoriaceous, attenuated into a petiole at
the base and with an abscission layer above the one at the apex of the secondary
stem. Inflorescence terminal, usually one from each stem, few-flowered; flowers
small. Dorsal вера! broad, cucullate, enclosing the column. Lateral sepals con-
nate into an oblong, bidentate lamina, longer than the dorsal sepal. Petals small,
(190)
1946]
FLORA OF PANAMA (Orchidaceae) 85
linear, inserted on the apex of the column-foot, retuse at the apex and with a
large, auriculate, hood-shaped callus or ligule toward the base. Column arcuate,
with a broad, membranaceous wing on each side; column-foot elongated, subequal
to the column; pollinia 2, ceraceous. -
A small genus of only two species, one Panamanian. Perhaps the most dis-
tinctive of the Pleurothalloid genera in Central America.
1. ACOSTAEA CosTARICENSIs Schltr. in Fedde Rep. Sp. Nov. Beih. 19:284. 1923.
Very small, caespitose, epiphytic herbs up to about 3 cm. tall. Secondary
stems up to about 1 mm. long. Leaves 9-15 mm. long and 2.5-6 mm. broad,
obovate-spatulate, obtuse, attenuated into a petiole at the base. Inflorescence up
to about 3 cm. long, few-flowered; peduncle slender, exceeding the leaves; flowers
small. Dorsal sepal 2.5-3 mm. long and 3-4 mm. broad, suborbicular, transverse,
strongly cucullate, obtuse. Lateral sepals 4-5.5 mm. long and together 2-2.5 mm.
broad, elliptic to elliptic-oblong, connate nearly to their apices and with the free
parts acute. Petals 1.5-2 mm. long and 0.3—0.5 mm. broad, ligulate, obtuse or
acute, falcate. Lip 2-2.5 mm. long and 0.5-1 mm. broad, with а large, thin,
hood-shaped callus near the base, apex emarginate and the lobules divergent.
Column arcuate, broadly winged, the wings membranaceous and subquadrate.
Costa Rica and Panama.
CLÉ: hills north of El Valle de Antón, alt. about 1000 m., Allen 2352; trail to Las
Minas, El Valle de Antón, alt. 1000 m., Allen 2892; mountains beyond La Pintada, alt.
400—600 m., Hunter & Allen 593.
23. PLEUROTHALLIS R. Br.
PLEUROTHALLIs К. Br. in Aiton, Hort. Kew. ed. 2, 5:211. 1813; Lindl. Folia
Orch. Pleuroth. 1859.
Restrepia HBK. ке» E & Sp. Pl. 1:366, #. 94. 1816.
Kraenzlinella O. Ktz ees & Kuntze, Gen. Phan. 310. 1904.
Platystele Schltr. in Fedde R p. Sp. Nov. 8:565. 1910.
Small to medium-sized pue herbs with caespitose or repent primary stems.
Inflorescence terminal or rarely pseudo-radical. Sepals subequal, erect or spread-
ing; dorsal sepal free or connate with the lateral sepals for a short distance, apex
sometimes clavellate; lateral sepals from nearly free to connate to their tips, often
gibbous at the base. Petals usually shorter and narrower than the sepals, apices
plain or clavellate. Lip simple or three-lobed, shorter or about as long as, or some-
times longer than, the petals, often unguiculate. Column about as long or shorter
than the lip, winged or wingless, produced into a foot, or sometimes footless at the
base; column-foot, when present, from short to as long as the column. Anther
terminal, operculate; pollinia 2 or 4, ceraceous.
Pleurothallis is one of the largest of the New World genera of orchids with
many diverse forms. The division of the genus most often used is that which
was proposed by Pfitzer (in Engl. & Prantl, Nat. Pflanzenfam., Nachtrage zu Teil
II-IV:105. 1897), but that is wholly artificial and quite impossible to use. The
(191)
[Vor. 33
86 ANNALS OF THE MISSOURI BOTANICAL GARDEN
separations used here are quite as artificial as those of Pfitzer and have nothing to
recommend them except, as far as the Panamanian species are concerned, they are
convenient even though they are not absolute. Further, it may cause allied species
to be widely separated.
a. Apices of the dorsal sepals or of the vent not clavellat
b. Sheaths not appressed to the stems, ciliat their mou i: „Series I. LEPANTHIFORMES (p. 86)
bb. gon usually appressed to the stems, EL mouths always glabrou
Inflorescence exceeded by the муены чи leaf, 1-flowered, кешік
or racemose Series П. BRACHYSTACHEAE (p. 86)
cc. Inflorescence subeq ual to or hing - cea 4 leaf,
flowered, a fascicle on a long ncle, mose......Series III. је (р. 87)
аз. Арісев 4 the dorsal se and id vm eer dd ENS Series ГУ. КЕЗТВЕРТАЕ (p. 88)
Series І. по EPANTHIFORMES.—A small, rather natural group of species
which are easily m d we loose Vra sheaths; inflorescence
= айе ог ап «һе ending
a. Petals not is y nor ха its петље de apex; stems with 1-3
leaves 1. P. BRoapwayi
aa. Petals ciliate or ciliate-lacerate toward the apex.
b. Sepals glandular-pubescent dorsally
bb. Sepals glabrous and smooth dorsally.
c. Leaves narrowly elliptic to oval, margins not recurved
2. Р, GNOMONIFERA
. P. BLAISDELL
. Lea
cc. Leaves broadly "e to Беке rar qnibos sphacelate margins
кин recur 4. Р. ROTUNDATA
Series П. BRACHYSTACHEAE.—An artificial group which probably has no
phylogenetic mra but is of value in separating and keying species of this
ifficult genus. Plants in which the inflorescence is exceeded by the
subtending leaf. Inflorescence a single flower, a fascicle of flowers, a
raceme or fascicle of racemes.
a. oe а fascicle ү” опе to many flowers ас the apex of the
stem.
ondar
ы” pec now cordate at the "T mostly bro:
c. Lip about 6 mm. long; petals 2- to 3- dae ‘flowin relatively
lar Р. CARDIOCHILA
сс. Lip 4 m. or less long; eng 1-nerved; flowers relatively small.
d. josh. — or denticu
e. Base of the lamina of che lip with a transverse E-shaped
callus Р. RHODOGLOSSA
ee. Base of a lamina of the lip without an E-shaped callus.
‚ Lip st po or iir ый ai gm А n smooth........ . P. TRIANGULABIA
Lip LL oblong-c Low 8. P. ANTONENSIS
ff.
dd. we -— ci iliate nor den id
e. Bas the lip with i ы ы arms; petals semi-terete
above 9. Р. ARIETINA
ee. Base of the lip without lateral arms; petals not semi-terete.
f рое of lamina of the lip with a transverse E-shaped
6. Р. RHODOGLOSSA
ff. Base J the lamina of the lip without a transverse E-shaped
callus 10. P. HOMOLANTHA
bb. Leaves not Же 26 at the base, usually attenuated.
c. Secondary stems 25. P. ALPINA
nge
ec. Muda stems not ged.
d. Lateral in "те ас a mel above the middle, usually nearly to
20. P. TRIBULOIDES
at the base 15. P. TRACHYCHLAMYS
te.
f. Petals abruptly dilated
ff. he als not к dilated at the base.
E 18. P. OCTOMERIAE
a small falcate lateral lobes y ^A Val Базе.......... 19. P. UNCINATA
dd. Pol mith connate to their apices or nearly
(192)
19461
FLORA OF PANAMA (Orchidaceae)
e. Lip deeply saccate-
ее. Lip not saccate-concave.
f. Petals serrulate (or ciliate).
2 Dorsal sepal bois lip not ^a с к 5.5.) 13;
. Dorsal sepal мин ж, lip EE elis- 11.
if Petals entire (sometimes ciliolat
в. = forming a ee: һы at the base, hispidulous
N
—
55. == not forming a distinct tube at the base, the
do a. 5. from the laterals, glabrous. |
echinate 20.
hh. басе пог егі т
i Lip about as ped. s long, € me basal lobes...... 14.
i. Lip longer dam чи; simple t least not with
е basal lobes
. Pe ii els of the flowers less than / as long as the
gn 17,
jj- Pedicels of the flowers at least half as long as the
eaves 16.
aa. Inflorescence а few- to several-flowered raceme.
. Secondary stems less than half as long as the leaves they bear.
C. Sep als pube escent within 46.
се Sepals MES. ithin.
. Inflorescence a "ad of short racemes, much shorter than
the subtending 36
4а. net cal : eds raceme nearly as long as the subtending
lea
bb. eo stems at least half as long as җе leaves that they bear.
c. Pet "r s linear or linear- lanceolate, enti
d. Lip lepidote-verrucose a 28.
ip not сени slate 42.
а. T:
cc. Petals broader than linear or linear-lanceolate (or if not then
serrulate), entire or serrulate
d.
Petals very vds ad recurved- -arcuate; denticul 33.
dd. Mir not stro ved-arcuate; dencieulare or entire.
recurve
Lip broader d Жн base ellip 34.
ee. к А ос broader th nor transversely Ten
. Lip with small acral grew r the middle
g- pope stems with кайс: т wings.
(Cf. also P. vELATICAULIS
ch вера! 9-11 mm. long, elliptic-obovate................ 29.
1
. Do
hh. Dorsal вера! 6-6.5 mm. long, elliptic to oblong-
Е u ond . 30.
hh. Petals not denticulate, oblong to adi oblanceolate.... 31.
iddle
ff. Lip without lateral он near the mi
g. Petals not serru lat
h. Lip with erect ruses near the middle; petals oblong
to oblong- ames. <
t
oblon cbe to su ubr ho
gg. Petals а at least
h. Lateral sepals connate to » i apices; petals elliptic to
elli
ptic-lance
hh. а макту: ree for at least № their length; petals
broader than elliptic or spines зак
^ Petals oblong-oblanceolat 27.
шегі subrhombic-obova "№.
бегіев "d ELON eries of species not = eei е” allied
but ing the 2.2 босса іп common. Plant apris the
Sci exceeds the subtending leaves in length. Inflo orescence a sin-
le-fl d to a many-flowered raceme or a fascicle of flowers on an
1
gle-flowere
elongated peduncle.
a. Inflorescence a 1- to several-flowered fascicle on a long peduncle.
(193)
87
. P. CONCAVIFLORA
P. COBRAEFORMIS
P. sIMULANS
P. HISPIDA
P. TRIBULOIDES
P. ALLENII
P. RUsCIFOLIA
P. EUMECOCAULON
P. sEGOVIENSIS
. P. STENOSTACHYA
7. P. OVATILABIA
P. LEPIDOTA
P. PRUINOSA
P. ROWLEE!I
P. PRAEGRANDIS
P. COGNIAUXIANA
25. P. ALPINA
P. УІТТАТА
P. VELATICAULIS
31. P. vELATICAULIS
hh. Lip without erect pens near the middle; petals
n ond 32. P. GELIDA
35. P. ELLIPSOPHYLLA
P. VERECUNDA
P. PTEROCAULIS
[Vor. 33
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN
b. Lateral lobes of the ҰЗ erect; lamina with a pair of parallel lamellate
calluses near the mi 23
bb. Lateral ad of the гк if present, not erect; lamina without par-
allel lamellate calluses.
с. Pedunc e puberulent 24.
с. Peduncle след 5%
aa. Inflorescence a
Ь. Peduncle бга and win
nged.
с. ap of the lip p or retuse 50.
pex of the lip 49.
bb. [S eh not ме nor winged.
c. Lateral sepals free nearly to Si bases, at least not connate to
nea middle.
r the
d. rrii ary muricate 51.
dd. Ovary not muricate.
e. Disc of the lip with 3 prominent longitudinal callus
ee. Disc of the lip without prominent longitudinal «кек
$
>
oo
f. Sepals 10 mm. or more long 52.
5
ff. hag 19 mm. or les
s long.
tals fimbriate or мел са 41.
s above.
. Lip narrowly oblong; pem Mee acuminate 39.
ip ovate; Dod sepals acu obtuse 37
cc. Lateral sepals connate to about the middle « or usually to near their
d. Secondaty stems less than half as long as the leaves they bear.
e. Ovary 40
мә CV v
P. FULGENS
P. GLANDULOSA
P. BRIGHAMII
P. sPECTABILIS
РЕАУП
. MURICATA
Р
. P. DOLICHOPUS
CRENATA
ARISTATA
CALYPTROSTELE
OVATILABIA
echinate . P. РОЕСИ var
ee. Ovary not echinate. MEN
f. Petals fimbriate or д рен -fimbriate 41. P. ARISTATA
ff. Petals not fimbriate nor r lacerate-fimbriate,
Ж T4 3- lobed; sepals pubescent with 46. P. SEGOVIENSIS
imple; se glabrous eb, 38. P. Сковут
dd. р stems at. is half ei long as the leaves they bear.
e. Petals linear or anceolat ‚ 42. P. PRUINOSA
ee. Petals not linear nor linear- теке T either lanceolate or
oblong or broader.
f. Petals truncate 4
>
als pubescent within 46
gg. Sepals glabrous —
Lateral sepals 5-9 mm. long.
1. m als alliptic- бм to ovate-lanceolate, 6-8.5
ong 47
г жені oblong-obovate to subrhombic, 2.5-3.5 mm.
long 3
hh. Lateral sepals 12-25 mm. lon
erect basal sien or lobes; petals ver-
rucose at the apex
к ғғ. basal auricles or lobes; petals not ver-
ucose
Series ТУ. ResrREPIAE.—AÀA 42. cnin group of species mainly distin-
guished by th poete vn ке of the хун sepal and the petals.
Sheaths of yw sec usually maculat
a. Disc of the te an par rad ew lip verrucose; VM NN more than
half as long as the lea
b. iei wich a short y PRO appendage on the anterior side just
b
5
1.
53
bb. Pedicel idunt pend
aa. Disc of the янги с hes of the lip not verrucose; inflorescence less
than half as long as the lea
. P. MACRANTHA
. P. SEGOVIENSIS
. P. GHIESBREGHTIANA
P. cELIDA
. P. TuERCKHEIMII
. Р. PowEerrn
P. FILAMENTOSA
54. P. SUBSERRATA
55. P. XANTHOPHTHALMA
1. PLEUROTHALLIs BRoADwAYI Ames, Orch. 2:267. 1908; С. Schweinf. in Bot.
Mus. Leaf. Harv. Univ. 8:41. 1940.
Pleurothallis Williamsii Ames, Orch. 7:120. 1922; Sched. mpr 7:31,
Pleurothallis Broadwayi var. tricarinata C. Schweinf. loc. cit
(194)
t. 15. 1924.
19461
FLORA OF PANAMA (Orchidaceae) 89
Small, caespitose, epiphytic herbs
up to about 15 cm. tall. Secondary
stems up to about 6 cm. long, slen-
der, bearing 1-3 leaves. Inflorescence
exceeding the leaves; peduncle sub-
filiform, rachis somewhat fractiflex.
Leaves 0.6—1.6 cm. long and 0.3—0.8
cm. broad, elliptic-oval to obovate,
obtuse, short-petiolate. Dorsal sepal
3.5-6 mm. long and 1.2-2 mm.
broad, lanceolate to ovate-lanceolate,
acute or acuminate, somewhat cucul-
late. Lateral sepals 3—5.5 mm. long
and 0.6—1.2 mm. broad, free almost
to the base, lanceolate, acute or
acuminate, oblique. Petals 1.1—1.5
mm. long and 0.6-0.8 mm. broad,
oval to obovate, obtuse, slightly ob-
lique, short-clawed. Lip 1-2 mm.
Fig. 96. Pleurothallis Broadwayi
long and 0.7-1.1 mm. broad, ellip-
tic-ovate, obtuse, sometimes obscurely trilobate, the lamina bi-tricarinate, papilli-
ferous, the apex thickened.
Mexico, Honduras, Costa Rica, Panama, and the West Indies.
DARIEN: Cana and vicinity, alt. about 600—1950 m., Williams 976.
2. PLEUROTHALLIS GNOMONIFERA Ames, Sched. Orch. 6:61. 1923.
Pleurothallis brevis Schltr. in Fedde Rep. Sp. Nov. Beih. 19:183. 1923.
Small, caespitose, epiphytic herbs up to 12 cm. tall. Secondary stems up to
about 8 cm. long, covered with infundibuliform, hispidulous sheaths. Leaves
1.4-4.5 cm. long and 0.6-2 cm. broad, elliptic to ovate, acute or obtuse, coriaceous,
attenuated into a short petiole at the base. Inflorescence one to several racemes
from the axil of the leaf, usually less than half as long as the leaves, few-flowered.
Sepals glandular-pubescent dorsally; dorsal вера! about 3-4 mm. long and 1 mm.
broad, oblong-lanceolate, acute or obtuse; lateral sepals 3-4 mm. long and together
2.5-3 mm. broad, connate at the base or as far as the middle, free parts oblong,
obtuse. Petals about 1.5 mm. long and 0.5 mm. broad, oblong, obtuse, ciliate or
lacerate at the apex. Lip 1.6-2 mm. long and 0.5-0.6 mm. broad, narrowly ob-
long, obtuse, subpandurate, biauriculate at the base, with a small submammillate
callus near the base of the disc.
Costa Rica and Panama.
cHIRIQUÍ: Cerro Punta to headwaters of the Rio Caldera, alt. 2250-2500 m., Allen
1466; Cerro Horqueta, alt. 1500-1800 m., Powell 311.
(195)
[Vor. -33
90 ANNALS OF THE MISSOURI BOTANICAL GARDEN
3. PLEUROTHALLIS BLAISDELLI S. Wats. in Proc.
Am. Acad. 23:284. 1888.
Pleurothallis peraltensis Ames, Sched. Orch. 6:65.
1923; loc. cit. 7: f. II, figs. 1-5. 1924.
Small, caespitose, epiphytic herbs up to about
15 cm. tall. Secondary stems up to 10 cm. long,
covered with several more or less pubescent,
loose sheaths which disintegrate with age. Leaves
2—6.5 cm. long and 0.4—2 cm. broad, narrowly
elliptic to oval, acute or obtuse, coriaceous, at-
tenuated into a short petiole at the base. In-
florescence a raceme or fascicle of racemes, short,
usually less than half as long as the leaves, few-
flowered. Dorsal sepal 4-6 mm. long and 1.5-2
mm. broad, lanceolate to oblong-lanceolate,
acute, usually eciliate. Lateral sepals 4-6 mm.
long and together 2.5-3 mm. broad, connate at
the bases or nearly to the middle, free portions
oblong-lanceolate, acute or obtuse, usually ciliate
toward the base on the outer side. Petals 2-2.5
mm. long and 1-1.5 mm. broad, oblong, obtuse,
apical part ciliate or fimbriate. Lip 2.5-3 mm.
long and 0.75-1 mm. broad, oblong-lanceolate
or oblong-pandurate, obtuse, ciliate, auriculate at
the base, short-unguiculate.
Fig. 97. Pleurothallis Blaisdellii Mexico, British Honduras, Honduras, Costa
Rica, and Panama.
CHIRIQUÍ: Bajo Chorro, alt. about 1800 m., Davidson 160; “Chiriquí Province,” alt.
about 1200 m., Powell 424.
4. PLEUROTHALLIs ROTUNDATA С. Schweinf. in Bot. Mus. Leafl. Нагу. Univ.
4.113. 1937.
Small, caespitose, epiphytic herbs up to about 13 cm. tall. Secondary stems
up to about 9 cm. long, slender, covered with several hispid, infundibuliform
sheaths. Leaves 2.5-4.5 cm. long and 2-3.7 cm. broad, broadly oval to sub-
orbicular-ovate, obtuse, with a very short petiole, the margins sphacelate and
rolled toward the dorsal surface. Inflorescence a fascicle of 1(—few?) -flowered
racemes, shorter than the leaves. Dorsal вера! 4.5-5.5 mm. long and 1.5-2.1 mm.
broad, lanceolate, acute or acuminate, ciliate or eciliate. Lateral sepals about 5
mm. long and together 2.5—3 mm. broad, ovate-lanceolate, connate to about the
middle, free apices fleshy-thickened, ciliate laterally toward the base and near the
sinus within. Petals about 1.5 mm. long and 0.75 mm. broad, oblong or oblong-
obovate, obtuse, the apex ciliate-lacerate. Lip 2.5-3 mm. long and 0.8-1 mm.
broad, lanceolate to ligulate-lanceolate, acute or acuminate, ciliolate, short-
unguiculate.
(196)
91
19461
FLORA OF PANAMA (Orchidaceae)
Endemic to Panama.
COCLE: vicinity of El Valle, alt. 600-1000 m., Allen 1242; north rim of El Valle de
Anton, alt. 600-1000 m., Allen 1676; hills north of El Valle de Anton, alt. about 1000
m., Allen 2206, 2259; mountains beyond La Pintada, alt. 400-600 m., Hunter 9 Allen
Fig. 98. Pleurothallis cardiochila
5. PLEUROTHALLIS CARDIOCHILA L. Wms. in Ann. Missouri Bot. Gard. 29:343.
1942.
Small, caespitose, epiphytic herbs up to about 21 cm. tall. Secondary stems up
to about 11 cm. long, slender, with 1-2 loose chartaceous sheaths toward the base,
naked above. Leaves 9-10 cm. long and 3.6-3.9 cm. broad, lanceolate-cordate,
acuminate, coriaceous. Inflorescence а 1(—few?)-flowered fascicle subtended by
а chartaceous sheath about 1-1.5 cm. long, much shorter than the leaves. Flowers
very large for the group, yellowish with the dorsal sepal tinged with dark red, lip
(197)
[Vor. 33
92 ANNALS OF THE MISSOURI BOTANICAL GARDEN
deep orange. Dorsal sepal about 20 mm. long and 12.5 mm. broad, oblong-oval,
obtuse, with 7—9 principal nerves, strongly cucullate. Lateral sepals connate to
their apices, about 18 mm. long and 10 mm. broad, ovate, acute, with 7-9 prin-
cipal nerves. Petals about 13 mm. long and 2.5 mm. broad, linear-oblong, acute,
arcuate, subauriculate on the posterior margin at the base, entire, 3-nerved at the
base, the posterior nerve short, the anterior nerve branched near the base, hence
the apical part of the petal’ 3-nerved. Lip unguiculate; lamina about 6 mm. long
and 5.5 mm. broad, suborbicular-cordate, obtuse, callus thickened along the basal
sinus and below the subapiculate apex, 3-nerved; claw about 2 mm. long, oblong.
Endemic in Panama.
CHIRIQUÍ: Bajo Chorro, alt. about 1800 m., Davidson 119.
Pleurothallis cardiochila is most closely allied to P. palliolata Ames, from which
it is distinguished by the suborbicular-cordate lip and the 3-nerved, entire petals.
6. PLEUROTHALLIS RHODOGLOSSA Schltr. in Fedde Rep. Sp. Nov. Beih. 17:22.
1922.
Small, caespitose, epiphytic herbs up to about 2 dm. tall. Secondary stems up
to about 13 cm. long, slender, with 1-2 sheaths at the base, otherwise naked.
Leaves 4-7 cm. long and 2.2-4 cm. broad, broadly ovate-cordate, acute or acumi-
nate, coriaceous. Flowers rather small, greenish with a purple lip. Dorsal sepal
about 3.5—4 mm. long and 1.8-2.2 mm. broad, oblong-obovate, obtuse, 3-nerved.
Lateral sepals about 3 mm. long and 3 mm. broad, orbicular, connate to the apices.
Petals about 2-2.5 mm. long and 0.5-0.7 mm. broad, linear-ligulate, acute, slight-
ly arcuate, obscurely denticulate (ciliolate, fide Schltr.). Lip about 1.4-1.7 mm.
long and 1.1-1.3 mm. broad, cordate or narrowly cordate, obtuse; lamina fleshy
and with a transverse E-shaped callus toward the base (open part of the "E"
toward the base).
Panama.
HIRIQUÍ: "Province of Chiriqui," alt. m., Powell 182. (Schlechter i in his de-
Sube cites Powell 182 but gives the alien as “Auf Hiigeln bei Panama-city”).
Two specimens of this have been seen. The one in the Ames Herbarium gives
the data cited above. The specimen in the Herbarium of the Missouri Botanical
Garden bears the data given by Schlechter. The Chiriqui locality is doubtless
correct.
7. PLEUROTHALLIS TRIANGULABIA C. Schweinf. in Ann. Missouri Bot. Gard,
24:183. 1937.
" Caespitose epiphytic herbs up to about 4 dm. tall. Secondary stems up to
about 3 dm. long, slender, with loose sheaths which soon disintegrate. Leaves
3.5-12 cm. long and 1.3-6 cm. broad, oblong-cordate to ovate-cordate, acuminate,
coriaceous. Inflorescence a 1- to few-flowered fascicle, the flowers rather large.
Dorsal sepal 9-12 mm. long and 6-9 mm. broad, oblong-ovate to oval, acute or
obtuse, larger than the combined laterals. Lateral sepals 9-12.5 mm. long and
(198)
19461
FLORA OF PANAMA (Orchidaceae) 93
together 5-8 mm. broad, ovate-lanceolate to broadly ovate, acute or obtuse, con-
nate to their tips. Petals 6-8 mm. long and 1.5-2.1 mm. broad, elliptic-linear,
acute, ciliate to lacerate-dentate, arcuate. Lip 3.2-4 mm. long and 3-4 mm.
broad, triangular to triangular-subquadrate, surface verrucose, base truncate with
a submammillate callus in the middle with cavities on either side.
Costa Rica and Panama.
IRIQUÍ: Cerro Punta to headwaters of the Rio Caldera, alt. 2250-2500 m., Allen
ту m us of the upper d Chiriquí Viejo, alt. 1300—1900 m., Seibert 269; Bajo
Chorro, alt. 1800 m., Davidson 223.
8. PLEUROTHALLIS ANTONENSIS L. Wms. in Ann. Missouri Bot. Gard. 29:341.
Caespitose epiphytic herbs up to about 3 dm. tall. Secondary stems slender,
with one or two scarious sheaths at the base, naked above. Leaves 4—9 cm. long
and 1.5—4.7 cm. broad, lanceolate-cordate to broadly cordate (juvenile leaves
elliptic), acute or acuminate, coriaceous. Inflorescence a 1- to several-flowered
fascicle from the apex of the secondary stems, much shorter than the subtending
leaves. Dorsal берг! 6-7.5 mm. long and 3-4 mm. broad, elliptic-obovate, obtuse
or acute, 3-nerved. Lateral sepals connate to their apices, 5.5—7 mm. long and
4-5 mm. broad, ovate to broadly ovate, obtuse or acute, 6-nerved. Petals 3.8—4.5
mm. long and 0.6-1 mm. broad, linear-oblong, acute, serrulate, strongly arcuate,
1-пегуед. Lip unguiculate; lamina 2.5—3 mm. long and 2-2.5 mm. broad, cordate
to oblong-cordate, denticulate, fleshy, the surface obscurely verrucose or smooth,
with a small central cavity near the base; claw short.
Endemic іп Panama.
LÉ: hills north of El Valle de Antón, alt. about 1000 m., Allen 2156; same rons
Pring pe^ 2267, 2701; mountains beyond La Pintada, alt. about 400—600 m., Hunte
& Allen 504.
Specimens of Pleurothallis antonensis have been referred to Р. phyllocardia
Reichb. f., a rather obscure and poorly described species.
9. PLEUROTHALLIS ARIETINA Ames, Sched. Orch. 4:16. 1923; loc. cit. 7: £. 16.
1924.
Caespitose epiphytic herbs up to 3.5 dm. tall. Secondary stems up to 2.5 dm.
long, slender, ancipitous or winged, at least when dry, with one or more basal
sheaths, naked above. Leaves 6—13 cm. long and 1.5—3.5 cm. broad, lanceolate
to ovate-lanceolate, acute or acuminate, cordate at the base when mature, coria-
ceous. Inflorescence a fascicle of 1—several, rather long-pedicellate flowers, shorter
than the leaves. Dorsal вера! 9-10 mm. long and 2.5—4 mm. broad, oblong to
oblong-lanceolate, obtuse, cucullate, fleshy. Lateral sepals 9—10 mm. long and
together 2-4 mm. broad, oblong or oblong-lanceolate, obtuse, cucullate, fleshy,
connate to their apices. Petals 7-10 mm. long and 1.4—1.8 mm. broad, linear-
oblong or linear-lanceolate, obtuse, attenuated into a thin claw at the base, the
apical half semi-terete. Lip 1.5—2.5 mm. long and 2-3 mm. broad when expanded,
triangular-crescentiform and with two linear-lanceolate arms at the base, the
(199)
[Vor. 33
94
ANNALS OF THE MISSOURI BOTANICAL GARDEN
\
М
|
N \ ы
N N
Pleurothallis arietina
Fig. 99.
(200)
19461 i
FLORA OF PANAMA (Orchidaceae) 95
whole when spread out suggesting a ram’s head.
Costa Rica and Panama.
IRIQUÍ: Cerro Horqueta, alt. 1500—1660 m., Powell 288. состЕ: mountains be-
кеі ‘La Pintada, alt. 400-600 m., Hunter 9 Allen 505.
10. PLEUROTHALLIS HOMOLANTHA Schltr. in Fedde Rep. Sp. Nov. Beih. 19:106.
1923.
Pleurothallis nemorum Schltr. loc. cit. 191.
Small, caespitose, epiphytic herbs up to about 3.5 dm. tall. Secondary stems
up to about 23 cm. long, slender, with one or more sheaths toward the base, naked
above. Leaves 3.5-10 cm. long and 0.9-3.5 mm. broad, elliptic-lanceolate to
ovate, cordate at the base, acute or acuminate. Inflorescence a fascicle of 1—few
flowers, much shorter than the leaves. Dorsal вера! 6-10 mm. long and 3-4.5
mm. broad, elliptic-oblong to elliptic-obovate, acute. Lateral sepals connate to
their apices, 6-10 mm. long and 3,5-6 mm. broad, lanceolate-ovate to ovate,
acute, somewhat cucullate. Petals 3.5-5 mm. long and 0.5-1 mm. broad, elliptic-
linear, acute, arcuate, glabrous. Lip unguiculate; lamina 2-3.5 mm. long and
1.5-3 mm. broad, oblong-cordate, obtuse (rarely acutish), fleshy.
Costa Rica and Panama.
HIRIQUÍ: valley of the upper Río Chiriquí Viejo, White t$ White 80. сосіЁ: El
Valle, alt. 600-1000 m., Allen 1236.
11. PLEUROTHALLIS SIMULANS L. Wms. in Ann. Missouri Bot. Gard. 27:281,
pl. 33, figs. 8-10. 1940.
Small, caespitose, epiphytic herbs up to about 10 cm. tall. Secondary stems
2.5—6 cm. long, slender, usually a little longer than the leaves, with one or two
sheathing bracts at the base. Leaves 3-5 cm. long and 0.5—0.8 cm. broad, ligulate
to elliptic-oblanceolate, obtuse, coriaceous, submarginate. Inflorescence 1-flowered
or а few-flowered fascicle, shorter than the subtending leaf. Dorsal вера! about
10 mm. long and 3.5—4 mm. broad, lanceolate, acuminate. Lateral sepals connate
to their apices, about 10 mm. long and 4.5—5 mm. broad, broadly lanceolate,
acuminate. Petals 5-6 mm. long and 2-2.5 mm. broad, broadly elliptic to sub-
rhombic, acute or acuminate, lacerate-ciliate nearly to the base. Lip 3—3.5 mm.
long and 2-2.5 mm. broad, simple, subcordate in outline, acuminate, the anterior
margin somewhat serrulate, the disc covered with a thick callus simulating that
found in species of Stelis; the callus with an anterior rim, an inconspicuous de-
pression toward its apex and sharply declined to the attachment of the lip.
Panama.
coc north rim of El Valle de Antón, Allen 1912; trail to Las Minas, hills north
of El Valle de Antón, Allen 2
An interesting and conspicuous species of Pleurothallis because of the unusual
structure of the lip.
(201)
[Vor. 33
96
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 100. Pleurothallis simulans
12. PLEUROTHALLIS CONCAVIFLORA C. Schweinf. in Bot. Mus. Leafl. Harv. Univ
4:114. 1937; L. Wms. in Ann. Missouri Bot. Gard. 28:416. 1941.
Small, caespitose, epiphytic herbs up to about 11 cm. tall. Secondary stems
up to about 7 cm. long, slender. Leaves 2.5-4 cm. long and 0.7-1.5 cm. broad,
elliptic-oblong, obtuse or acute, coriaceous. Inflorescence a fascicle of 1- to
several-flowered peduncles. Sepals obscurely puberulent or glabrous within. Dorsal
sepal 4-5 mm. long and 2.5-3.3 mm. broad, oblong-ovate to oblong-obovate,
acute or obtuse, cucullate, fleshy. Lateral sepals connate to their apices, about
4.5 mm. long and 4-5 mm. broad, ovate-suborbicular to suborbicular, cucullate,
obtuse or acute. Petals 1.7-2.5 mm. long and 0.6-0.9 mm. broad, linear-oblong
со elliptic-oblong, acute or obtuse. Lip 2-2.5 mm. long and 1-1.5 mm. deep,
3-lobed or at least the base produced into prominent retrorse auricles, saccate-
(202)
19461
FLORA OF PANAMA (Orchidaceae) 97
concave, apiculate at the apex; lateral lobes or auricles retrorse or erect, triangular-
lanceolate; the disc with a callus connecting the bases of the lobes or auricles.
Costa Rica and Panama.
COCLÉ: vicinity of El Valle de Antón, alt. about 600 m., Allen 2058; vicinity of La
Mesa, alt. about 1000 m., Allen 2317.
This species seems to differ from P. excavata Schltr. only in having slightly
smaller flowers. Мо material of P. excavata is available for study. The types of
the two units are from the same locality in Costa Rica.
13. PLEUROTHALLIS COBRAEFORMIS L. Wms. in Ann. Missouri Bot. Gard. 27:277,
pl. 34, figs. 7-14. 1940. |
Small, caespitose, epiphytic herbs up to about 13 cm. tall. Secondary stems
mostly up to 4-6 cm. long, slender, subequal to the leaves or a little longer.
Fig. 101. Pleurothallis cobraeformis
(203)
[Vor. 33
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Leaves 4-6 cm. long and 0.8-1.1 cm. broad when mature, obtuse or acute, fleshy.
Inflorescence a 1- to several-flowered fascicle (or possibly an abbreviated raceme?),
much shorter than the leaves. Dorsal вера! 8-10 mm. long and 7-9 mm. broad,
suborbicular, subacute, strongly cucullate. Lateral sepals connate to their apices,
about 6-7 mm. long and 4.5-5 mm. broad, broadly oval. Petals 4-6 mm. long
and 1.2-2.5 mm. broad, elliptic, acute, arcuate, dentate-ciliate. Lip about 3.5
mm. long and as broad, short-clawed; lamina apiculate, broadly cordate, fleshy,
Fig. 102. Pleurothallis Allenii
(204)
1946]
FLORA OF PANAMA (Orchidaceae) 99
sharply deflexed, the basal auricles rounded.
Panama.
COCLÉ: vicinity of El Valle de Antón, alt. about 600 m., Allen 2057.
14. PLEUROTHALLIs ALLENI Г. Wms. in Ann. Missouri Bot. Gard. 27:275, pl.
33, figs. 5-7. 1940
Small, caespitose, epiphytic herbs up to about 15 cm. tall. Secondary stems
3-6 cm. long, slender, shorter than the leaves, with one or two sheaths at the
base. Leaves 5-9 cm. long and 0.3—0.6 cm. broad, ligulate to linear-ligulate,
obtuse or tridentate, coriaceous. Inflorescence a 1- to several-flowered fascicle,
shorter than the leaves, the flowers large for the size of the plant. Dorsal sepal
about 15 mm. long and 4 mm. broad, lanceolate, long attenutate-acuminate.
Lateral sepals connate nearly to their apices, about 15 mm. long and together
about 5 mm. broad, lanceolate, acute. Petals about 12 mm. long and 3 mm.
broad, lanceolate, acute or acuminate, slightly oblique. Lip about 4 mm. long
and as broad, 3-lobed; the lateral lobes erect, about 1.25 mm. long and 1 mm.
broad, broadly oblong, the apices rounded; the mid-lobe about 2 mm. long and
1.5 mm. broad at the base, narrowly triangular, acute; disc very fleshy, with a
tripartite raised callus near the middle, and the mid-lobe covered with echinulate
protuberances.
Panama.
OCLÉ: vicinity of El Valle de Antón, alt. 600-1000 m., Allen 1240; vicinity of
La n alt. about 1000 m., Allen 2338.
15. PLEUROTHALLIS TRACHYCHLAMYS Schltr. in Fedde Rep. Sp. Nov. Већ. 17:23.
1922; Ames, Sched. Orch. 7:31, #. 10. 1924.
Small, repent, epiphytic herbs. Secondary stems up to about 6 cm. long,
covered with scarious sheaths which are scurfy or at least verrucose. Leaves 4.5—7
cm. long and 0.4-1 cm. broad, elliptic to lanceolate, acute. Inflorescence a 1- to
few-flowered fascicle. Dorsal sepal 4-5 mm. long and 1.5—1.8 mm. broad, ovate
or oblong-lanceolate, acute. Lateral sepals 4-5 mm. long and about 1.5 mm.
broad, free to the bases, oblong-lanceolate, acute. Petals about 4 mm. long and
1.5 mm. broad, lanceolate and abruptly expanded near the bases. Lip about 2
mm. long and 1 mm. broad, oblong-oval, obtuse, with 2 small, erect, lateral lobes
near the middle, with a small callus at the base.
Costa Rica (?), Panama, and dubiously in Peru.
PANAMÁ: foothills east of Panama City, Powell 215.
16. PLEUROTHALLIS EUMECOCAULON Schltr. in Fedde Rep. Sp. Nov. Beih.
19:187. 1923; L. Wms. in Ann. Missouri Bot. Gard. 27:277. 1940.
Small, caespitose, epiphytic herbs up to about 2 dm. tall Secondary stems
slender, strict, with one or two sheaths. Leaves 2.5-7.5 cm. long and 0.4-1.3
cm. broad, linear, elliptic or usually lagceolate, acute, coriaceous. Inflorescence
a fascicle of long (2-3 cm.) -pedicellate flowers; one or more short, scarious bracts
(205)
(Моб 33
100 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
at the base. Dorsal sepal 5-7 mm. long and 2.5—3.5 mm. broad, lanceolate to
ovate-lanceolate, acute or acuminate, cucullate. Lateral sepals connate to their
apices, 4.5-5.5 mm. long and 2.5-3.5 mm. broad, broadly lanceolate to ovate-
lanceolate, obtuse or acute, fleshy. Petals 4.5-5 mm. long and about 1-1.5 mm.
broad, linear-lanceolate to lanceolate, acuminate, papilliferous, the acumination
subterete. Lip 3-3.5 mm. long and 1.5-2 mm. broad, papilliferous to subciliate,
lanceolate to oblong-ovate, acute, truncate at the base, slightly constricted above
the middle, fleshy, with a small V-shaped excavation above the middle and with
3 raised lines on the dorsal surface.
COCLE: vicinity of El Valle, alt. 600-1000 m., Allen 1237; hills north of El Valle de
Antón in the vicinity of La Mesa, alt. 1000 m., Allen 2 2330.
17. PLEUROTHALLIs RUSCIFOLIA (Jacq.) К. Br. in Aiton, Hort. Kew. ed. 2, 5:211.
1813; Ames in Bot. Mus. Leafl. Harv. Univ. 192. р. 5. 1933.
Epidendrum ruscifolium Jacq. Enum. Pl. Carib. 29. 176
Dendrobium ruscifolium Sw. in Nova Acta Soc. Sci. Upsala 6:84. 1799.
Pleurothallis glomerata Ames, Sched. Orch. 4:21. 1923; loc. cit. 7: f. 10. 1924.
Caespitose, epiphytic herbs up to about 5 dm. tall but usually much smaller.
Secondary stems up to about 4 dm. long, slender. Leaves 4-20 cm. long and 1-5
cm. broad, linear-lanceolate to elliptic to lanceolate-ovate, acute or acuminate,
contracted into a short cucullate petiole at the base. Inflorescence a 1- to many-
flowered fascicle. Dorsal sepal 6-10 mm. long and 1.3-3 mm. broad, narrowly
to broadly lanceolate, cucullate. Lateral sepals 6-10 mm. long and together
2.5-3 mm. broad, lanceolate to ovate-lanceolate, cucullate, acuminate, connate to
their apices. Petals 4-8 mm. long and 0.31 mm. broad, subfiliform to narrowly
lanceolate, acute. Lip 1.5-2 mm. long and about 1.2-1.4 mm. broad, ovate to
oblong-cordate or ovate-cordate, acute or acuminate.
Costa Rica and Panama, the West Indies, and South America. Reported from
Guatemala and Salvador.
CHIRIQUÍ: Bajo coe alt. 1800 m., Davidson 262. cocré: hills north of El Valle
de Antón, alt. 800-1000 m., Allen 2260; trail to Las Minas, hills north of El Valle de
Anton, alt. 1000 m., proe
This variable and widespread species is the generic type of Pleurothallis.
18. PLEUROTHALLIS OCTOMERIAE Schltr. in Fedde Rep. Sp. Nov. Beih. 17:21.
1922.
Pleurothallis cerea Ames, Sched. Orch. 4:19. 1923; loc. cit. 7:26, fig. 4 and 1. 8. 1924.
Erect, caespitose, epiphytic herbs up to about 5 dm. tall. Secondary stems up
to about 3 dm. long and 0.4 cm. in diameter, covered with closely appressed
sheaths which may be scurfy, the sheaths disintegrating with age. Leaves 4—20
cm. long and 0.8-6 cm. broad, narrowly elliptic to broadly lanceolate, acute or
obtuse, coriaceous. Inflorescence a few- to many-flowered fascicle. Dorsal sepal
5.5-8 mm. long and 2-3 mm. broad, lanceolate to oblong-lanceolate, obtuse.
Lateral sepals 4.5-7 mm. long and 2-2.8 mm. broad, broadly lanceolate, obtuse
(206)
19461
FLORA OF PANAMA (Orchidaceae)
PLEUROTHALLIS
ruscttolia
Fig. 103. Pleurothallis ruscifolia
(207)
101 `
| [Vor. 33
102 ANNALS OF THE MISSOURI BOTANICAL GARDEN
or acute, usually arcuate, connate at the base and forming a short mentum. Petals
4-8 mm. long and 1.5-2.2 mm. broad, lanceolate, obtuse or acute, somewhat
arcuate. Lip 2-3 mm. long and 0.8—1.3 mm. broad, oblong to oblong-ovate,
obtuse, short-clawed, with 2 submarginal, elongated, sublamellate calluses which
are about as long as the lip.
Costa Rica and Panama; possibly also Honduras.
IRIQUÍ: e of Chiriqui,” Powell 252, 3432. COCLÉ: hills north of El Valle
de рее alt. 1000 m., Allen 2710. РАМАМА: hills above Campana, alt. 600-800 m.,
Allen 1881.
Allen 2719 consists of a fine series of plants which show that P. cerea is only
a robust phase of P. octomeriae.
19. PLEUROTHALLIS UNCINATA Fawc. in Jour. Bot. 33:12. 1895; Fawc. &
Rendle, Fl. Jam. 1:63, Ё. 10. 1910; C. Schweinf. in Bot. Mus. Leafl. Harv.
Univ. 7:151. 1939.
Pleurotballis Alexandrae Schltr. in Fedde Rep. Sp. Nov. Beih. 19:103. 1923.
Tall, rather coarse, subcaespitose, epiphytic herbs up to 5.5 dm. tall. Secondary
stems up to about 3.5 dm. long, covered with several loose, ancipitous, chartaceous
sheaths which soon disintegrate. Leaves 8-22 cm. long and 1-5 cm. broad, linear-
elliptic to oblong-lanceolate, acute, coriaceous. Inflorescence a 1- to few-flowered
fascicle with the flowers on long pedicels. Dorsal sepal 10-20 mm. long and 4-6
mm. broad, linear-oblanceolate to oblong, acute or obtuse, puberulent dorsally.
Lateral sepals 12-20 mm. long and 3-4 mm. broad, linear-lanceolate to oblong-
lanceolate, acute, puberulent on both sides. Petals 5-9 mm. long and 1.3-2.5
mm. broad, oblanceolate, acuminate, denticulate above, 3-nerved. Lip 6-8 mm.
long and 2.5-3 mm. broad, oblong-ovate, usually subpandurate, obtuse, with two
narrow, falcate lateral lobes above the base, typically with thick calluses near the
base of the lateral lobes and without a retrorse callus at the base. (Cf. note below.)
Costa Rica, Panama, and the West Indies.
осі: Boquete, alt. about 1800 m., Davidson 220 (sterile specimen). СОСІ.Е:
Valle ys pé RA about 800 m., Cope s.
Schweinfurth, loc. cit., has reduced P. Alexandrae Schltr. to P. uncinata with-
out comment. Adequate material may prove that it is distinct and that the Cope
specimen should be referred there. Тһе lip on the Cope specimen is relatively
small, has a prominent retrorse callus at the base (which typical P. uncinata lacks),
and is membranaceous at the base of the side lobes (while in P. uncinata a large
callus is situated at the base of each of the side lobes).
20. PLEUROTHALLIS TRIBULOIDES (Sw.) Lindl. Gen. & Sp. Orch. РІ. 6. 1930.
Epidendrum tribuloides Sw. Nov. Сеп. & Sp. РІ. Do "dd 1788.
Pleurotballis fallax Reichb. f. in B Е: 3:224.
Cryptophoranthus acaulis Kranzl. in Fedde Rep. Sp. Nov. Beih. 34:232. 1925.
Small, caespitose, epiphytic herbs up to 8 cm. tall. Secondary stems about 1
(208)
19461
FLORA ОЕ PANAMA (Orchidaceae) 103
cm. long, covered with loose, chartaceous bracts, slender. Leaves 2-7 cm. long
and 0.4-1.4 cm. broad, narrowly to broadly oblanceolate, obtuse, attenuated into
a petiole ас the base, coriaceous. Inflorescence a fascicle of 1—few brick-red flow-
ers. Dorsal вера! 5-7 mm. long and 2-2.2 mm. broad, oblong-lanceolate or oblong-
oblanceolate, acute, papilliferous, constricted just above the base. Lateral sepals
often connate to their apices, 4.5-7.5 mm. long and about 3 mm. broad, oblong or
oblong-lanceolate, acute, cucullate, papilliferous. Petals 2.5-3 mm. long and
1.2-1.4 mm. broad, oblong-oblanceolate, oblique, lobulate, acute, fleshy especially
toward the apex. Lip 2.5-3 mm. long and 0.7-1.5 mm. broad, oblong or oblong-
lanceolate, lobulate below the middle, obscurely ciliate toward the apex. Capsule
echinate. |
Mexico, British Honduras, Guatemala, Honduras, Costa Rica, Panama, and
the West Indies.
CHIRIQUÍ: banks of Caldera River, alt. 1360 m., Powell 237, 3376.
21. PLEUROTHALLIs HISPIDA L. Wms. in Ann. Missouri Bot. Gard. 27:277, pl. 33,
figs. I-4.
Ни Powellii Ames, Sched. Orch. 4:8. 1925, non Pleurothallis Powellii
chltr.
Small, caespitose, epiphytic herbs up
to about 5 cm. tall. Secondary stems
0.5-2 mm. long, shorter than the leaves,
with 2 or 3 cauline bracts, the upper
infundibuliform, maculate with maroon
dots and hispid at least along the angles.
Leaves 1—3.5 cm. long and 0.3—1.3 mm.
broad, narrowly elliptic to elliptic-oval,
obtuse, coriaceous, margins hispid or his-
pidulous. Inflorescence a one- to few-
flowered fascicle. Dorsal sepal connate
with the laterals for half its length, 5—6
mm. long, lanceolate, acute, hispidulous
dorsally. Lateral sepals about 6 mm.
long and together about 6 mm. broad,
connate to their tips, strongly cucullate,
in natural position calceolate, carinate,
hispidulous dorsally and papillose with-
in. Petals about 4 mm. long and 1 mm.
broad, linear-oblong, the apex obliquely
truncate and acute. Lip about 3 mm.
long and 1.5 mm. broad, unguiculate;
Fig. 104. Pleurothallis hispida lamina oblong-lanceolate, with a lamel-
late callus extending from each lateral
5
(209)
[Vor. 33
104 ANNALS OF THE MISSOURI] BOTANICAL GARDEN
angle toward the apex and with a mammillate callus at the junction of the lamina
and the claw, apical margins entire or denticulate; claw about 0.5 mm. long,
minutely biauriculate at the base.
Endemic in Panama.
cocLÉ: vicinity of El Valle, alt. 600-1000 m., Allen 782, 1243. PANAMA: San
Juan, Powell 278.
22. PLEUROTHALLIs BricHaMu S. Wats. in Proc. Am. Acad. 23:285. 1888;
Ames, Sched. Orch. 7:19, £. у. 1924.
Pleurothallis barboselloides Schltr. іп Fedde Rep. Sp. Nov. Већ. 17:18. 1922; loc. cit.
59: 1. 20, fig. ІІ
Pleurothallis periodica Ames, loc. cit. 7:21, t. 4.
Қ... acrisepala Ames & Schweinf. Sched. Orch.
923,
Small, caespitose, epiphytic herbs up to 12 cm.
tall. Secondary stems up to about 6 mm. long,
very short and inconspicuous, enclosed by sheaths.
Leaves 1.8-9 cm. long and 0.3-1.2 cm. broad,
oblanceolate to linear-oblanceolate, obtuse or acute,
attenuated to the base, coriaceous. Inflorescence
a 1- to several-flowered fascicle borne on a long
slender peduncle, up to 12 cm. long, subequal to
or exceeding the leaves. Flowers small, opening
one at a time. Sepals connate for a short distance
at the base; dorsal вера! 6-10 mm. long and 1.2-3
mm. broad, elliptic to oblong-lanceolate, obtuse
to acuminate. Lateral sepals 6-9 mm. long and
together 3-4.5 mm. broad, oblong to oblong-
ovate, connate to the middle or beyond, acute.
Petals 2-3.5 mm. long and 0.75-1.7 mm. broad,
obliquely spatulate to obliquely oval, the anterior
margin produced, acute or acuminate. Lip 2.1-4
mm. long and 0,8-2 mm. broad, oblong-ligulate
to oblong-lanceolate, obtuse, the apex ciliate or
denticulate, with an auriculate lateral lobe on
Fig. 105. Pleurothallis Brighamit either side near the base of the lamina, claw with
a prominent callus.
A not uncommon plant at low elevations from Guatemala to Panama.
AS DEL TORO: without locality, Wedel 185; Isla Colón, Wedel 486. CANAL ZONE:
dacs Cond, Island, Kenoyer 256, Sbattuck 205, 567; drowned forest between Tumba
Vieja and Salamanca, Steyermark & Allen 16750; Salamanca Hydrographic Station, Wood-
son, Allen & Seibert 1585; hills east of or near Panama City, alt. near sea- ris Powell
02, 219; Gatün Lake, Powell 275. DARIÉN: vicinity of El Real, alt. about 1 , Allen
043. PANAMÁ: Río La Maestra, alt. 0—25 m., Allen 54; Río Tecámen, ides of "Chepo
Road, alt. about 30 m., Hunter 9 Allen 225; 1 near Tapía River, Juan Díaz region, Maxon
& Harvey 6699; Casa Ladre, San Juan, hills east of Panama City, Powell 274, 275, 420,
3508.
(210)
19461
FLORA ОЕ PANAMA (Orchidaceae) 105
Dorsal se pal
Fig. 106. Pleurothallis alpina
Pleurotballis Brighamii is a variable species which may eventually prove to be
the same as P. pyrsodes Reichb. f.
23. PLEUROTHALLIS FULGENS Reichb. f. in Gard. Chron. n. s. 4:516. 1875.
Small, caespitose, epiphytic herbs up to about 10 cm. tall. Secondary stems.
up to about 1 mm. long. Leaves 4—8.5 cm. long and 1—1.8 cm. broad, elliptic to
broadly oblanceolate, attenuated into a petiole at the base. Inflorescence a single
(211)
қ [Vor. 33
106 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
flower or a few-flowered fascicle borne on a long slender peduncle, about as long
as or exceeding the leaves. Dorsal sepal 8-9 mm. long and about 3.5 mm. broad,
ovate-lanceolate, acuminate, somewhat cucullate. Lateral sepals 10-12 mm. long,
connate nearly to the middle, the bases gibbous; free part of the sepals about 6-7
mm. long and 3 mm. broad, lanceolate, acute or acuminate. Petals about 4 mm.
long and 1.5 mm. broad, sublanceolate but oblique, produced anteriorally into an
auricle at the middle, acute. Lip about 5 mm. long and 2.5 mm. broad, elliptic,
obtuse, with a pair of small, erect, lateral lobes below the middle and a pair of
parallel, submarginal, lamellate calluses near the middle.
Costa Rica and Panama.
cHIRIQUÍ: Cerro Horqueta, alt. 1200-1500 m., Powell 335, 340.
The description is based on a record of the type and the specimen cited. Speci-
mens from Costa Rica which have been referred here are very much larger, have
relatively short inflorescences, and show slight floral differences.
24. PLEUROTHALLIS GLANDULOSA Ames, Sched. Orch. 6:60. 1923.
Pleurothallis vittariifolia Schltr. in Fedde Rep. Sp. Nov. Beih. 19:26. 1923.
Small, caespitose, epiphytic herbs up to about 5 cm. tall. Secondary stems 1
mm. long or less. Leaves 1.5-2.5 cm. long and 0.1-0.2 cm. broad, obtuse, at-
tenuated to the base. Inflorescence a single flower or a few-flowered fascicle;
peduncle slender, puberulent, up to about 4 cm. long, exceeding the leaves. Pedicel
and ovary up to about 1 cm. long, puberulent. Dorsal вера! 6-9 mm. long and
about 1.5 mm. broad, linear-lanceolate, acute or acuminate, puberulent and lightly
keeled dorsally. Lateral sepals 6-8 mm. long and together about 2.5 mm. broad,
connate to about the middle, gibbous at the base, puberulent and lightly keeled
dorsally. Petals about 3 mm. long and 1.3 mm. broad, subdolabriform, cuneate
at the base, oblique, apiculate. Lip about 3.5 mm. long and 1 mm. broad, unguic-
ulate; lamina subpandurate and auriculate at the base, glandular-puberulent on
the margin and upper surface.
Costa Rica and Panama.
CANAL ZONE: near Mes Ж San Juan, Dodge, Steyermark & Allen 16578. PROVINCE
(PA ANAL ZONE m Тапа Grande Range, alt. near sea-level,
Powell pi Ns perdis di, Purdom
25. PrEUROTHALLIS ALPINA Ames, Sched. Orch. 5:14, fig. 3. 1923.
Caespitose epiphytic herbs up to about 4 dm. tall. Secondary stems slender,
up to about 3 dm. long, prominently 2- or 3-alate, expanded at the apex and the
wings passing into the leaves. Inflorescence a raceme or fascicle of racemes, few-
flowered, short, not half as long as the leaf. Leaves 5-12 cm. long and 0.6-2.5
cm. broad, ligulate to elliptic-lanceolate or elliptic-oblanceolate, acute or obtuse,
coriaceous. Sepals acute, fleshy, obscurely puberulent dorsally; dorsal sepal 6—6.5
mm. long and 1.5—2.5 mm. broad, elliptic to oblong-oblanceolate, cucullate;
lateral sepals 5-6 mm. long and 1.5-2 mm. broad, lanceolate, arcuate. Petals
about 2.5 mm. long and 1-1.2 mm. broad, elliptic-oblong, acute, serrulate, arcuate.
Пір 2.5-4.5 mm. long and 1.5-2 mm. broad, elliptic-oblong, obtuse, with two
(212)
19461
FLORA OF PANAMA (Orchidaceae) 107
Gordon а ЖТ.
Fig. 107. Pleurothallis pterocaulis
small lanceolate-arcuate lateral lobes about 0.5 mm. long near the middle, apex
fimbriate, with two carinate, longitudinal calluses near the middle of the disc.
(213)
[Vor. 33
108 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
Costa Rica and Panama.
CHIRIQUÍ: Caramillia, alt. 1360 m., Powell 282. сосіЁ: vicinity of El Valle, alt.
600-1000 m., Allen 1238; hills north of El Valle de Antón, alt. about 800-1000 m.,
Allen 2280.
A not uncommon species which is allied to Р. sicaria Lindl.
26. PLEUROTHALLIS PTEROCAULIS L. Wms. in Ann. Missouri Bot. Gard. 27:280,
t. 32, figs. 1-7. 1940
Repent epiphytic herbs up to about 16.5 cm. tall. Secondary stems 2.5-7 cm.
long, winged or strongly angled (at least when dry), shorter than the leaves.
Leaves 4-11 cm. long and 1-1.5 cm. broad, elliptic to elliptic-lanceolate, acute,
coriaceous. Inflorescence a fascicle of one- to few-flowered racemes, shorter than
the leaves. Dorsal вера! about 6 mm. long and 2-2.5 mm. broad, oblong-lanceolate,
strongly cucullate, carinate with the apex thickest. Lateral sepals connate for
about half or more their length, 5-6 mm. long and together about 5 mm. broad,
each sepal oblong-lanceolate, somewhat oblique, acute, carinate and with a thicker
carinate cushion near the outer margins. Petals about 2.5 mm. long and 1.7 mm.
broad, subrhombic-obovate, the apical margin serrulate. Lip about 4 mm. long
and 1.5 mm. broad, unguiculate, oblong-lanceolate, obtuse, carinate, with two in-
conspicuous calluses at the junction of the lamina and the claw, the claw incon-
spicuously biauriculate at the base.
Endemic in Panama.
COCLE: vicinity of El Valle de Antón, alt. 600-1000 m., Allen 1230.
Originally said to be closely allied to P. hondurensis Ames, it is probably more
closely allied to P. verecunda Schltr.
27. PLEUROTHALLIS VERECUNDA Schltr. in Fedde Rep. Sp. Nov. Beih. 17:24.
22
Erect, repent, epiphytic herbs up to about 3.5 dm. tall. Secondary stems up
to about 19 cm. long, terete or angled (at least when dry). Leaves 8.5-16 cm.
long and 1-2.2 cm. broad, elliptic or lanceolate-ligulate, acute, coriaceous. In-
florescence a few-flowered raceme about half as long as the leaves. Dorsal sepal
5.5-7 mm. long and 1.5-2 mm. broad, lanceolate to oblong-lanceolate, acute.
Lateral sepals 5.5-7 mm. long and each 1.5-2.5 mm. broad, lanceolate, acute,
apex thickened, free nearly to the base. Petals 2.2-3.2 mm. long and 0.75-1 mm.
broad, oblong-oblanceolate, oblique, apical half denticulate. Lip 3-3.5 mm. long
and 1.5-2 mm. broad, oblong to oblong-ovate, obtuse, truncate at the shortly
unguiculate base; lamina with two short auricles at the base, fleshy, with a sub-
mammillate callus at the apex of claw and two lateral callus thickenings.
Costa Rica and Panama.
CANAL ZONE: Rio Indio, near mouth of Chagres River, alt. about sea-level, Powell 01;
Barro Colorado Island, Shattuck 722.
(214)
19461
FLORA OF PANAMA (Orchidaceae) 109
Fig. 108. Pleurothallis verecunda
(215)
[Vor. 33
110 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 109. Pleurothallis lepidota
(216)
19461
FLORA ОЕ PANAMA (Orchidaceae) 111
28. PLEUROTHALLIS LEPIDOTA L. Wms. in Ann. Missouri Bot. Gard. 27:279, pl.
32, figs. 8-12. 1940.
Small, caespitose, epiphytic herbs up to about 2.2 dm. tall. Secondary stems
about 9-12 cm. long, partially invested with 2-3 scarious sheaths. Leaves 7-10
cm. long and 3-3.5 cm. broad, elliptic to elliptic-lanceolate, short-acuminate.
Inflorescence one or more few-flowered racemes, shorter than the leaves. Dorsal
sepal 13-14 mm. long and about 2 mm. broad, linear-lanceolate, acute. Lateral
sepals about 14 mm. long and together 12 mm. broad at their bases, subtriangular,
connate nearly to their apices, acuminate or acute, carinate-winged along the mid-
nerve dorsally, outer surface papilliferous. Petals about 5 mm. long and 0.25.
mm. broad, linear, acute. Lip about 8 mm. long and 5 mm. broad, unguiculate,
obscurely 3-lobed, upper surface lepidote-verrucose, with a mammillate callus at
the junction of the lamina and the claw, margins serrulate to serrate; lateral lobes
2 mm. long and as broad, rounded; mid-lobe oblong-lanceolate, acute, about 4—5
mm. long and 2 mm. broad.
Panama.
CHIRIQUÍ: Llanos del Volcán, alt. about 1300 m., Allen 1552.
29. PLEUROTHALLIS COGNIAUXIANA Schltr. in Fedde Rep. Sp. Nov. 3:246. 1907.
Repent or caespitose epiphytic herbs up to about 3.2 dm. tall. Secondary
stems up to about 19 cm. long, slender, naked or with 1—2 sheaths at the base,
with 2-3 narrow but prominent wings. Leaves 8-15 cm. long and 3-9 cm.
broad, elliptic-lanceolate to ovate-lanceolate. Inflorescence 1 or more short
racemes up to half as long as the leaves. Dorsal sepal 9-11 mm. long and 3-4.5
mm. broad, elliptic-obovate, obtuse or acute, slightly verrucose within. Lateral
sepals 5-7.5 mm. long and together 5-7 mm. broad, suborbicular, connate to their
apices or nearly so, slightly verrucose within. Petals 3.5-4.5 mm. long and
1.5-2.4 mm. broad, lanceolate-subrhombic, acute, terminal half denticulate. Lip
4-5 mm. long and 2-3 mm. broad, subrhombic, unguiculate, with two inflexed
auricles or lateral lobes below the middle and two auricles at the base of the claw,
surface of the apical half verrucose and the margin denticulate, with 2 parallel
callus thickenings near the mi
Costa Rica and Panama.
Qui: vicinity of “New Switzerland,” alt. 1800-2000 m., Allen 1383; Bajo
к: at 1800 m., Dies 100, 362
30. PLEUROTHALLIs VITTATA Lindl. in Bot. Reg. 24: Misc. р. 73. 1838; L. Wms.
in Ann. Missouri Bot. Gard. 26:280. 1939.
Pleurothallis polystachya Rich. & Gal. in Ann. Sci. Nat. ІП, 3:16. 1845.
Pleurotballis mandibularis Kránzl. in Vid. Medd. ab. Foren 71:169. 1920.
Pleurothallis В Kranzl. in Ark. f. Bot. 168:15.
Repent or caespitose epiphytic herbs up to about 2.5 dm. tall. Secondary
stems up to 15 cm. long, prominently angled, at least when dry. Leaves 5-12
cm. long and 1-2.3 cm. broad, lanceolate to elliptic-ligulate, acute or obtuse,
(217)
[VoL. 33
112 ANNALS OF THE MISSOURI BOTANICAL GARDEN
coriaceous. Inflorescence 1-several short racemes, % as long as the leaves or less.
Sepals densely puberulent dorsally; dorsal sepal 5-8 mm. long and 1.5-2 mm.
broad, oblanceolate, acute; lateral sepals 5-8 mm. long and together 3-5 mm.
broad, ovate or oblong-ovate, connate to their apices, slightly gibbous at the base.
Petals 2.2-2.7 mm. long and 1.2-1.5 mm. broad, obovate, acute or obtuse, serru-
late above. Lip 2-3 mm. long and 1.2-1.5 mm. broad, oval, obtuse, denticulate
above, unguiculate, with 2 small inflexed, crescent-shaped lateral lobes near the
middle of the lip and 2 auricles at the base of the claw; lamina with 2 parallel
callus-like thickenings near the middle.
Mexico, Guatemala, Honduras, Costa Rica, Panama, and possibly Venezuela.
LÉ: hills south of El Valle de Antón, alt. 700 m., Allen 2513; between Las Маг-
ws and El Valle, Woodson, Allen 9 Seibert 1282.
31. PLEUROTHALLIS VELATICAULIs Reichb. f. in Linnaea 22:824. 1849.
Pleurothallis Pittieri Schltr. in Fedde Rep. Sp. Nov. 3:247. 1907.
Small to large, caespitose, epiphytic herbs up to about 6 dm. tall. Secondary
stems up to 4 dm. long, slender to stout, partly to entirely covered with loose
sheaths. Leaves extremely variable, 8-22 cm. long and 1.2-9 cm. broad, ob-
lanceolate-ligulate to oval, attenuated into a petiole at the base. Inflorescence a
raceme ог fascicle of racemes, shorter than the leaf. Dorsal вера! 3-5.5 mm. long
and 1.2-1.8 mm. broad, lanceolate or lanceolate-oblong, acute, somewhat cucullate.
Lateral sepals 3.5-5 mm. long and 1-1.8 mm. broad, lanceolate, acute, lightly
connate nearly to the apices but becoming free. Petals 1.5-3 mm. long and
0.5-0.75 mm. broad, oblong to oblong-oblanceolate, obtuse. Lip 1-2 mm. long
and 1-1.5 mm. broad, oblong-oval to subrhombic, obtuse, with fleshy, erect,
auriculate lateral lobes near the middle.
Costa Rica, Panama, the West Indies, and from Venezuela to Peru.
CHIRIQUÍ: forests between Las Palmas and top of Cerro de la Horqueta, alt. 2100-
2268 m., Pittier 3227. cocrí: hills north of El Valle de Antón, alt. 1000 m., Allen 2245.
One of the most variable of the species of Pleurothallis in Panama.
32. PLEUROTHALLIs GELIDA Lindl. in Bot. Reg. 27: Misc. р. 91. 1841.
Pleurotballis chiriquensis Schltr. in Fedde Rep. Sp. Nov. Beih. 17:19. 1922.
Caespitose epiphytic herbs up to about 5.5 dm. tall. Secondary stems up to
about 3.5 dm. long and about 0.4 cm. in diameter toward the base, partially
covered with chartaceous sheaths. Leaves 8—23 cm. long and 1.5—7 cm. broad,
elliptic-oblong to oblong-ovate, obtuse or acute, coriaceous, attenuated into a short
petiole at the base. Inflorescence 1-several racemes, either shorter, subequal to, or
longer than the subtending leaf, few- to many-flowered. Dorsal sepal 5-7.5 mm.
long and 2-2.5 mm. broad, oblong-lanceolate, acute, cucullate, puberulent within.
Lateral sepals 5-6 mm. long and each 2-2.5 mm. broad, acute, connate to the
middle or nearly so, puberulent within. Petals 2.5-3.5 mm. long and 1.5-2.5 mm.
broad, oblong-obovate to subrhombic, obtuse or truncate and the apex often
dentate or lobulate. Lip 2-2.5 mm. long and 0.9-1.4 mm. broad, dilated from a
(218)
1946)
FLORA OF PANAMA (Orchidaceae) 113
ZA
752 >
повишите"
|
|
H
veal
he
om
SNO
~ са
=o
Fig. 110. Pleurothallis gelida
(219)
114
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
narrow base into an oblong-cuneate lamina, truncate, with 2 parallel, lamellate
calluses near the middle.
Florida, Mexico to Panama, the West Indies, South America.
BOCAS DEL TORO: "Chiriqui," alt. 1200—1500 m., Powell 227.
A variable species and one of the most widespread of the genus.
Fig. 111.
Pleurothallis Rowleei
(220)
19461
FLORA OF PANAMA (Orchidaceae) 115
33. PLEUROTHALLIs КОУТЕЕГ Ames, Sched. Orch. 1:11. 1922; loc. cit. 7: Ё. 18.
1924. |
Pleurothallis cucullata Ames, loc. cit. 3:5. 1923; loc. cit. 7: £. 17. 1924.
Caespitose or repent epiphytic herbs up to 3.5 dm. tall. Secondary stems up
to 25 cm. long, slender, partially covered with one or two sheaths. Leaves 3-13
Fig. 112. Pleurothallis praegrandis
(221)
[Vor. 33
116 ANNALS OF THE MISSOUR! BOTANICAL GARDEN
cm. long and 1—5.4 cm. broad, elliptic-lanceolate to ovate-lanceolate, acuminate,
subcordate at the base. Inflorescence 1 to few slender racemes, shorter or rarely
longer than the subtending leaf. Dorsal sepal 6-11 mm. long and 2.5—5.5 mm.
broad, lanceolate to ovate-lanceolate, acute or acuminate, cucullate. Lateral sepals
7-10 mm. long and together 2.2-7 mm. broad, elliptic-oval to ovate, acute or
obtuse, connate to the apices. Petals 6-8.5 mm. long and 1.2-2.5 mm. broad,
subsigmoid-elliptic, denticulate, acute. Lip 2.5-4 mm. long and 1.5-2.5 mm.
broad, subsagittate; the basal portion broad and fleshy; the apical portion nar-
rowly triangular.
Costa Rica and Panama.
снікюоі: Bajo Chorro, alt. 1810 m., Davidson. DARIEN: Cana-Cuasi trail, alt. 1650
m., Terry & Terry 156.
34. PLEUROTHALLIS PRAEGRANDIS Ames, Sched. Orch. 5:17. 1923; loc. cit.
7:5. 17. 1924.
Large, caespitose, epiphytic herbs up to about 12 dm. tall. Secondary stems
up to about 9 dm. long. Leaf 31.5 cm. long and 20.5 cm. broad, cordate, acumi-
nate. Inflorescence consisting of several racemes which are shorter than the leaf.
Dorsal вера! 2,5-3 mm. long and about 1-1.5 mm. broad, elliptic-oblong, acute.
Lateral sepals about 3.5 mm. long, connate to their apices, suborbicular, cochleate.
Petals about 2 mm. long and about 1 mm. broad, oblong-spatulate, obtuse, the
apex fleshy-thickened. Lip about 1 mm. long and 2.5-3 mm. broad, transversely
elliptic; disc with a transversely rhombic thickening in the middle.
Panama.
DARIEN: Cana and vicinity, alt. about 1800 m., Williams 973.
Description based on a photograph and a flower of the type, and the original
description.
35. PLEUROTHALLIS ELLIPSOPHYLLA L. Wms. in Ann. Missouri Bot. Gard. 29:344.
Small, repent or caespitose, epiphytic herbs up to about 2 dm. tall. Secondary
stems up to 9.5 cm. long, prominently angled when dry, with one or more loose
sheaths covering the basal part. Leaves 4—9.5 cm. long and 0.7-2.7 cm. broad,
elliptic to elliptic-oblong, acute or obtuse, coriaceous. Inflorescence of 1 to several
short, few-flowered racemes, about half as long as the leaves. Dorsal sepal 9-14
mm. long and 1.5-2.5 mm. broad, narrowly lanceolate to elliptic-oblanceolate,
acute or acuminate, cucullate, puberulent dorsally. Lateral sepals 10-13
mm. long and together 3—4 mm. broad, connate except at the very tip, the
lamina lanceolate, acute or acuminate, pubescent dorsally, gibbous and with a
mentum at the base. Petals 5.5—6.5 mm. long and 1.4-2 mm. broad, elliptic or
elliptic-lanceolate, acute or acuminate, arcuate, denticulate or denticulate-lacerate
toward the middle but the base and apex usually entire. Lip 4—5 mm. long and
1—1.3 mm. broad, lanceolate, acute or subaristate, biauriculate at the base, claw
(222)
19461
FLORA OF PANAMA (Orchidaceae) 117
Бір. 113. MEN ellipsopbylla
short; lamina with an inconspicuous V-shaped callus on the basal 14.
Panama.
AS DEL TORO: Maccaw Hill, Isla Colón, alt. 0—125 m., von Wedel 560; Río Crica-
mola, between St. Louis and Konkintóe, alt. about 10-15 m., Woodson, Allen & Seibert
1684.
36. PLEUROTHALLIS STENOSTACHYA Reichb. f. in Linnaea 18:399. 1844.
Pleurot ballis myriantba Lehm. & Kranzl. in Engl. Bot. m 26:445. 1899.
Pleurotballis Lankesteri Rolfe in Kew. Bull. 1914: 210.
Pleurotballis dubia Rich. & Gal. var. myriantba Schltr. in Реда Кер. 5р. Моу. Већ.
220221922;
Pleurothallis stenostachya Reichb. f. хат. Lankesteri Ames, Sched. Orch. 7:31, Ё. 15 pars.
1924.
Small, caespitose, epiphytic herbs up to about 1 dm. tall. Secondary stems up
to about 2 cm. long, very slender, covered with loose sheaths. Leaves long-
petiolate; lamina 0.5-5 cm. long and 0.3—0.9 cm. broad, linear-elliptic to oblong-
elliptic or usually oblanceolate, attenuated into the petiole; petiole very slender,
0.3—3 cm. long. Inflorescence a fascicle of short, few-flowered racemes; the
flowers from greenish to brick-red and very small. Dorsal вера! 1.2-1.7 mm.
(223)
(Vor. 33
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN
long, lanceolate-ovate to ovate, strongly cucullate, acute. Lateral sepals 1.2-1.5
mm. long and 0.5-0.75 mm. broad, segments lanceolate-ovate, acuminate, connate
below the middle. Petals 1-1.5 mm. long and 0.2-0.6 mm. broad, linear-lanceolate,
acute, arcuate. Lip 1.1—1.3 mm. long and 0.5—0.7 mm. broad, elliptic-oval, obtuse,
glandular-puberulent above, short-unguiculate.
Mexico, British Honduras, Guatemala, Honduras, Costa Rica, and Panama.
у
Quí: Boquete, alt. 1500 m., Davidson 1268; El Boquete, alt. 1000-1300 m.,
Pittier үреді “Chiriqui,” alt. 1060 m., Powell 224; vicinity of Boquete, alt. 1200-1500
, Woodson & Schery 800
А small-flowered and distinctive species.
37. PLEUROTHALLIS OVATILABIA Ames & Schweinf. Sched. Orch. 10:33. 1930.
Small, caespitose, epiphytic herbs up to about 8 cm. tall. Secondary stems up
to about 6 mm. long, covered with scarious sheaths. Leaves 2.5-6 cm. long and
0.25-0.6 cm. broad, narrowly oblanceolate,’ obtuse, attenuated into a long petiole
at the base. Inflorescence surpassing the leaves or rarely subequal; raceme slender,
unilateral, up to about 3 cm. long. Sepals 1.5-1.8 mm. long and 0.9-1 mm.
broad, subovate or oblong-ovate, acute or obtuse, cucullate. Petals 1.4-1.5 mm.
long and 0.3-0.4 mm. broad, linear-lanceolate, acute, falcate. Lip 1.5-2.5 mm.
long and about 1 mm. broad, ovate, acute.
Honduras, Costa Rica, and Panama.
CHIRIQUÍ: El Boquete, alt. 1000 m., Pittier
20.
Possibly only a variety of P. lancilabris
(Reichb. f.) Schltr
38. PLEUROTHALLIs Сковуг Batem. ех
Lindl. in Bot. Reg. 21: /. 1707. 1835;
Hook. in Bot. Mag. 65: /. 3682. 1838;
L. Wms. ex A. C. Sm. in Lloydia 2:172.
1737.
Pleurothallis picta Lindl. in Bot. Reg. 21: sub
1835; Lindl. loc. cit. 21: f. 1825.
меги marginata Lindl. loc. cit. 24:Misc.
1838.
Pett Hm - іп Fedde Кер.
жалы бан en in Fedde R
Sp. Nov. Beih. 19:186. 1923, non д
КА
—— integrilabia Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ.
3: 39.
Small, densely саеѕрісоѕе, epiphytic
herbs up to about 15 cm. tall. Leaves 1.5—7
Fig. 114. Pleurothallis Grobyi cm. long and 0.15-0.8 cm. broad, ob-
(224)
19461
FLORA OF PANAMA (Orchidaceae) 119
lanceolate to ovate-lanceolate to suborbicular, obtuse or acute, often marginate,
attenuated into a petiole at the base. Inflorescence a few- to several-flowered
raceme, usually much exceeding the leaf. Dorsal sepal 3.2-9 mm. long and 1.2-3
mm. broad, lanceolate to elliptic-ovate, acute or acuminate, cucullate. Lateral
sepals 3-11 mm. long and together 1.2-3 mm. broad, lanceolate to oblong-
lanceolate to ovate-lanceolate, acute or acuminate, gibbous at the base, connate to
the apex or nearly so. Petals 1.5-2.5 mm. long and 0.4-0.8 mm. broad, ovate-
oblanceolate or nearly so, acute or obtuse, more or less oblique. Lip 2-3 mm. long
and 0.6-1.3 mm. broad, oblong, obtuse, short-unguiculate.
Mexico to Panama, the West Indies and in South America.
ZONE: Quebrada Lopez, Allen 2137; Colorado poa Aviles 55; Sbattuck
226, с Standley 31400, 41176; near Vigia and San Juan, alt. 66 m., Dodge, Steyermark
& Allen 16526; around Gamboa, alt. 2-100 m., tms 4787; hills north of Frijoles,
Standley 27665; Salamanca Hydrographic іс Station, alt. about 80 m., Woodson, Allen 8
Seibert 1560, 1580. CHIRIQUÍ and VERA auf Pampasbáumen, Wa agner 24. COLON:
Rio Indio, alt. near sea-level, Pittier pu PANAMÁ: Río La Maestra, alt. 0—25 m.,
Allen 53; hill east of Panama City, Powell 106.
Pleurothallis Grobyi is quite a variable and widespread species. It is found
mostly near sea-level, rarely as high as 900 meters.
39. PLEUROTHALLIS CALYPTROSTELE Schltr. in Fedde Rep. Sp. Nov. Beih. 19:23.
1923; І. Wms. in Ann. Missouri Bot. Gard. 27:276, £. 33, figs. 11-15. 1940.
Pleurothallis biflora Schltr. loc. cit. 181, oc
а geminiflora Ames, Hub. & Schweinf. i in Bot. Mus. Leafl. Harv. Univ. 3:39.
Fig. 115. Pleurothallis calyptrostele
Small, repent or caespitose, epiphytic herbs up to about 3 cm. tall. Secondary
stems about 2-3 mm. long, covered with scarious sheaths. Leaves 5-15 mm. long
(225)
[Vor. 33
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN
and 1-4 mm. broad, oblanceolate, obtuse, attenuated into a petiole at the base.
Inflorescence a few-flowered raceme, longer than the leaves. Dorsal sepal 4-5.5
mm. long and 1.2-1.5 mm. broad, oblong-lanceolate, acuminate. Lateral sepals
3.5-6 mm. long and 1-1.6 mm. broad, lanceolate, acuminate, arcuate, free nearly
to the bases. Petals 1.8-2 mm. long and 0.5-0.6 mm. broad, oblanceolate, acute,
slightly oblique. Lip 2-2.5 mm. long and 0.5-1 mm. broad, narrowly oblong,
obtuse.
Costa Rica and Panama.
COCLE: vicinity of El Valle, alt. 600-1000 m., Allen 1233.
40. PLEUROTHALLIs ЕОЕСИ var. echinata L. Wms., var. nov.!
Small, caespitose, epiphytic herbs up to about 4 cm. tall. Secondary stems very
short, up to about 3 mm. long, covered with loose sheaths. Leaves 10-17 mm.
long and 3-6 mm. broad, oval to oblong-obovate, obtuse, attenuated into а
slender petiole at the base. Inflorescence exceeding the leaves, a simple few-
flowered raceme. Dorsal sepal 8-9 mm. long and 1.5-2 mm. broad, lanceolate,
long-acuminate. Lateral sepals connate to about the middle, about 7-9 mm. long
and together about 3 mm. broad, the segments lanceolate, long-acuminate. Petals
2.5-3 mm. long and 0.6-1 mm. broad, oblong-lanceolate, obtuse. Lip about 3.5
mm. long and about 1.5 mm. broad, short-unguiculate, oblong-ovate, obtuse,
constricted toward the middle. Ovary echinate.
Endemic in Panama.
CHIRIQUÍ: Volcán de Chiriquí, alt. about 2720 m., Davidson 081 (ТУРЕ in Herb.
mes).
The variety echinata differs from the species in having an echinate ovary,
narrower petals, larger flowers, and larger leaves. The material is not as good as
might be desired, and more adequate material may show it to be a distinct species.
41. PLEUROTHALLIS ARISTATA Hook. in Ann. & Mag. Nat. Hist. 2:329, 7. 15.
1839.
Pleurothallis dicbotoma Ames, Sched. "una 6: PA pe non Schltr.
Pleurotballis divexa Ames, loc. cit. 7:20,
Small, caespitose, epiphytic herbs up to about 7 cm. tall. Secondary stems
very short, 1-3 mm. long. Leaves 8-24 mm. long and 1.5—4.5 mm. broad, oblong
to oblanceolate, obtuse, attenuated into a petiole at the base. Inflorescence much
exceeding the leaves, a fractiflex raceme with relatively few flowers. Dorsal sepal
4.5—9 mm. long and 1-1.8 mm. broad, lanceolate to ovate-lanceolate, aristate-
acuminate, cucullate. Lateral sepals 4.5-9 mm. long and 1-1.6 mm. broad,
lanceolate, aristate-acuminate, connate about 1 mm. at the base. Petals about
2—3 mm. long and 1—1.5 mm. broad, lanceolate-ovate to ovate, acute or acuminate,
fimbriate or lacerate-fimbriate. Lip 2-3 mm. long and 0.6-1.5 mm. broad, ob-
lHerbae parvae Vena epiphyticae usque = 4 i altae. Folia ovalia vel oblongo-ovata,
obtusa. Sepalum sale lanceolatum, acuminatu Sepala lateralia usque ad medium MEN
lanceolata, acuminata. Petala oblongo- dm obtusa. Labellum oblongo-ovatum, obtusu
Ovarium echin
(226)
19461
FLORA OF PANAMA (Orchidaceae) 121
long, obtuse, subauriculate at the base, with 2 erect side lobes, keeled on the
mid-nerve below, apical half of lip more or less fleshy and terete, lepidote-verrucose.
Costa Rica, Panama, and the West Indies.
CHIRIQUÍ: Bajo Chorro, alt. 1800 m., Davidson 187.
The specimen upon which this record is based is unsatisfactory but perhaps it
belongs to this species.
42, PLEUROTHALIS PRUINOSA Lindl. in Bot. Reg. 28: Misc. p. 75. 1842.
Pleurothallis pauciflora Schltr. in Fedde Rep. Sp. Nov. Beih. 19:192. 1923.
Small, caespitose or repent, epiphytic herbs up to about 14 cm. tall. Secondary
stems slender, usually with but one sheath, longer than the leaves. Leaves 1-5
cm. long and 0.4—0.7 cm. broad, linear-elliptic to linear-lanceolate (rarely
broader), acute or obtuse, coriaceous. Inflorescence a slender, few-flowered raceme,
subequal to or exceeding the leaves (rarely shorter). Dorsal sepal 2.5-4 mm.
long and 1,5-1.8 mm. broad, lanceolate to ovate-lanceolate, acute or acuminate,
cucullate. Lateral sepals 2.5-4 mm. long and together 1.5—2.5 mm. broad, ovate
or ovate-lanceolate, cochleate, connate to their apices. Petals 2-3 mm. long and
0.3—0.5 mm. broad, linear or linear-lanceolate, acute. Lip 1.5-2 mm. long and
0.8—1 mm. broad, ovate-lanceolate to triangular-lanceolate, acute or acuminate,
fleshy.
Honduras, Costa Rica, Panama, the West Indies, Surinam, French Guiana, and
possibly Peru.
CHIRIQUÍ: Boquete, alt. 1150 m., Davidson 708; Chiriquí, alt. 1200 m., Powell 434.
A variable species. The Central American material is usually smaller than that
from the West Indies.
43. PLEUROTHALLIS MACRANTHA L. Wms. in Ann. Missouri Bot. Gard. 28:417,
pl. 21, figs. 1-6. 1941.
Caespitose, epiphytic or terrestrial herbs up to 23 cm. tall. Secondary stems
6-8 cm. long, subequal to the leaves in length on mature plants, almost com-
pletely invested with one or two loose, scarious sheaths, unifoliate. Leaves 4-7
cm. long and 1,5-2.8 cm. broad, elliptic, obtuse, coriaceous. Inflorescence a
several-flowered raceme, about twice exceeding the subtending leaves; sheath at
the base of the peduncle short, about 9-11 mm. long; bracts 6-7 mm. long, ovate-
lanceolate, scarious, semiamplexicaul. Dorsal вера! 16-20 mm. long and 6-8 mm.
broad, elliptic to oblanceolate-elliptic, acute, 3-nerved, keeled dorsally along the
mid-nerve. Lateral sepals 16-18 mm. long and together 6-8 mm. broad, connate
almost to their apices, oblong-oblanceolate, acute, together 4-пегуед, somewhat
gibbous at the base. Petals about 9-11 mm. long and 5-6 mm. broad near the
apex, oblong-obovate, slightly oblique, truncate or slightly retuse, 3-nerved, with
an inconspicuous cushion-like callus near the base. Lip 10-12 mm. long and 3-4
mm. broad toward the apex when expanded, about 2 mm. broad near the middle
and 3 mm. broad across the auriculate base, subterete, and sulcate above, somewhat
(227)
[Vor. 33
122 ANNALS OF THE MISSOURI BOTANICAL GARDEN
laterally compressed, expanding toward the apex and into two auricles at the base,
with three small calluses between the basal auricles.
Endemic in Panama.
CHIRIQUÍ: Bajo Chorro, alt. 1900 m., Woodson 9 Schery 666.
A very large-flowered species of Pleurotballis.
44. PrEUROTHALLIs TuERCKHEIMIXI Schltr. in Fedde Rep. Sp. Nov. 10:292. 1912.
Pleurothallis megachlamys Schltr. in Fedde Rep. Sp. Nov. Већ. 19:108. 1923.
Caespitose epiphytic herbs up to about 7.5 dm. tall, but usually about 3 dm.
tall. Secondary stems 5.5—42 cm. long, usually provided with 1-2 loose, ancipi-
tous, brown, chartaceous sheaths. Leaves %-27 cm. long and 2.5-7.5 cm. broad,
elliptic to oval, obtuse or rarely acute, attenuated into a short petiole at the base,
either longer or shorter than the secondary stem, usually shorter than the in-
florescence or occasionally subequal. Inflorescence an erect raceme which usually
exceeds the leaves, the flowers among the largest of the genus; sheath up to about
9 cm. long, conspicuous. Погва! sepal 11-25 mm. long and 4-6 mm. broad,
lanceolate, acute or acuminate, cucullate, puberulent dorsally. Lateral sepals con-
nate nearly to their apices, 14-25 mm. long and 6-7 mm. broad, elliptic to oblong-
lanceolate, acute, puberulent dorsally. Petals 4.5-8 mm. long and 2.5-4 mm
broad, oblong-oval to oval, arcuate, verrucose and subcochleate toward the apex.
Lip 5-9 mm. long and about 3—3.5 mm. broad, auriculate at the base, the auricles
erect; lamina ligulate, rostrate, fleshy, smooth or verrucose.
Mexico, Guatemala, Costa Rica, and Panama.
RIQUÍ: upper Río Chiriqui Viejo, alt. 1300-1900 m., White & White 18; vicinity
of Mon Lirio, alt. 1300-1900 m., Seibert 142.
45. PLEUROTHALLIS PowEL.u Schltr. іп Fedde Rep. Sp. Nov. Beih. 17:22. 1923.
Large (for genus), caespitose, epiphytic herbs up to about 7 dm. tall бес-
ondary stems up to probably 35 cm. long. Leaves 20—25 cm. long and 7-9 cm.
broad, oblong-elliptic to ovate-lanceolate, obtuse or acute. Inflorescence several-
to many-flowered, exceeding the leaves in length. Sepals all slightly connate at
the base, the laterals connate nearly to their apices; dorsal sepal 12-20 mm. long
and 2-2.5 mm. broad, ligulate or lanceolate, acute or acuminate; lateral sepals
14-20 mm. long and 6-7 mm. broad, oblong-lanceolate to lanceolate-ovate, cymbi-
form, acute. Petals about 6 mm. long and 1.6—2.6 mm. broad, oblanceolate to
oblong-oblanceolate, obtuse, falcate. Lip about 5 mm. long and 2-2,5 mm.
broad, lanceolate-ovate, obtuse, very fleshy, with 2 upright carinate wings above,
one on either side and one dorsally.
Costa Rica (?), and Panama.
CHIRIQUÍ: “Chiriquí,” alt. 1100-1200 m., Powell 146, 3365.
One of the largest species of Pleurothallis in Panama. Known only from one
collection, with field number and garden number—a numbering often practiced
by Mr. Powell, whose field numbers were rarely used on specimens and then always
in conjunction with the garden numbers.
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19461
FLORA OF PANAMA (Orchidaceae) 123
46. PLEUROTHALLIS SEGOVIENSIS Reichb. f. in Bonplandia 3:223. 1855; C.
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 4:96. 1937.
Pleurothallis Wagneri Schltr. in Fedde Rep. Sp. Nov. 17:141. 1921.
Pleurothallis falcatiloba Ames in Proc. Biol. Soc. Wash. 34:152. 1921.
Pleurothallis canae Ames, Sched. Orch. 2:18. 1923; loc. cit. 7: Ё. 13. 1924.
Caespitose epiphytic herbs 3-22 cm. tall, ordinarily 10-15 cm. tall. Secondary
stems about 0.5-6 cm. long, covered with sheaths, much shorter than the leaves.
Leaves (when mature) 4.5-12 cm. long and 0.2-1.2 cm. broad, linear-oblanceolate
to oblanceolate, obtuse or rarely acutish, gradually attenuated to the base. In-
florescence a few- to several-flowered raceme, often unilateral, usually exceeding
the leaves but rarely shorter. Dorsal sepal 5-11 mm. long and 2-3.5 mm. broad,
lanceolate to oblong-lanceolate, acute or acuminate, cucullate, keeled dorsally,
pubescent within (especially laterally) to glabrous. Lateral sepals 5.5-11 mm.
long, oblong-lanceolate to suborbicular-ovate, acute, cucullate to cochleate, con-
nate almost to their apices or rarely connate only 24 their length, pubescent within
(especially laterally) to glabrous, keeled dorsally along the mid-ribs. Petals 2—3.5
mm. long and 1-1.5 mm. broad, elliptic-lanceolate to ovate, obtuse or acute, more
or less arcuate, apex rarely trilobulate, the base auriculate. Lip 3-4 mm. long and
2.5-3 mm. broad, occasionally papilliferous, 3-lobed; lateral lobes basal or sub-
basal, from lanceolate-falcate to subtriangular, usually erect; lamina cblong,
obtuse, with 2 large lamellate or ridge-like calluses on the basal half or two
thirds, and with an elongate bipartite central callus at the base.
Mexico, Guatemala, Costa Rica, and Panama.
CHIRIQUÍ: Bajo Chorro, alt. 1800 m., Davidson 170; Rio Quebrada, alt. 1700 m.,
Killip 3540; “Chiriqui,” alt. 1200-1650 m., Powell 236, 278, 280, 3176; Wagner 25;
Monte Lirio, alt. 1300-1900 m., Seibert 190, 212, 213, 271; Bajo Mona, alt. 1500-2000
m., Woodson, Allen & Seibert 1004. COCLE: vicinity of La Mesa, alt. 1000 m., Allen
2307. DARIEN: Cana and vicinity, Williams 971.
One of the most variable of the species of Pleurothallis in Panama. The
description is taken from all of the specimens available. A critical study may
show some of the forms to be varietally distinct. In addition to the Panamanian
synonyms listed above the following names seem to belong here also: Pleurothallis
Aguilarii Ames, P. amethystina Ames, P. Jobannis Schltr., P. melicoides Schltr.,
P. pompalis Ames, and P. vinacea Ames.
47. PLEUROTHALLIS GHIESBREGHTIANA Rich. & Gal. in Ann. Sci. Nat. III, 3:16.
1845.
Jas racemiflora Lindl. ex Lodd. Bot. Cab. 10: /. 949. 1824, non (Sw.) Lindl.
Pleurothallis os aon pus Folia Orch. Pleuroth. p. 31. re pote 2% in Xenia
Im. s. 6-9. 1867; in Saunders’ Ref. Bot.
Pleurothallis bis Ыы Schltr. in Fedde Rep. Sp. Nov. 8:566. 5;
Pleurothallis Niederleinii Schltr. in Beih. Bot. Centralbl. 36, Abt. i 396. 1918.
Caespitose or repent epiphytic herbs up to about 6 dm. tall, but usually about
3 dm. tall. Secondary stems 3-18 cm. long, usually shorter than the leaves.
(229)
[Vor. 33
124 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
Leaves 4-17 cm. long and 0.9—3 cm. broad, oblanceolate to elliptic-oblong, obtuse
or rarely acute, usually attenuated into a short petiole at the base. Inflorescence
a few- to usually many-flowered raceme, usually much exceeding the subtending
leaves. Dorsal вера! 6-8.5 mm. long and 2.5-4 mm. broad, lanceolate to elliptic-
ovate, aeute. Lateral sepals 6—8.5 mm. long and together 3-5 mm. broad, elliptic-
oblong to oval, acute or obtuse, connate to their apices or nearly so, cucullate.
Petals 6-8.5 mm. long and 1.5-3 mm. broad, elliptic-lanceolate to ovate-lanceolate,
acute or obtuse, oblique. Lip 3.5-6.5 mm. long and 1.5-3 mm. broad, oblong-
pandurate, obtuse, basal portion orbicular, terminal portion oval.
Mexico, British Honduras, Guatemala, Honduras, Costa Rica, Panama, and
the West Indies.
IRIQUÍ: Cerro Vaca, alt. 900-1136 m., РИНег 5320. PANAMA: Rio La Maestra,
alt. 0-25 m., Allen 63. УЕКАСУА8: Bahia Honda, Taylor 1511
48. PLEUROTHALLIs DOLICHOPUS Schltr. in Fedde Rep. Sp. Nov. 10:394. 1912.
Pleurothallis lamprophylla Schltr. loc. cit. 15:205. 1918.
Pleurothallis poasensis Ames, Sched. Orch. 1:10. 1922; loc. cit. 7:/. 12. 1924.
Pleurothallis peregrina Ames, loc. cit. 6:67. 1923.
Caespitose or repent epiphytic herbs up to about 3.5 dm. tall. Secondary
stems up to about 23 cm. long, usually exceeding the leaves. Leaves 5-15 cm.
long and 1.3-3.5 cm. broad, elliptic-oblong, obtuse or rarely acute, contracted
into a short petiole at the base. Inflorescence 1 or more, few- to many-flowered
racemes, subequal to or usually exceeding the leaves. Dorsal вера! 14-23 mm.
long and 2-3.5 mm. broad, linear-lanceolate to lanceolate, ciliate and puberulent
to glabrous within. Lateral sepals 9-22 mm. long and 1.5-3 mm. broad, linear-
lanceolate to lanceolate, acuminate, ciliate and puberulent to glabrous within,
free almost to their bases. Petals 3-5 mm. long and 1-2 mm. broad, from nearly
oblong to oblong-oblanceolate, obtuse. Lip 2.5-5 mm. long and 0.8-2.5 mm.
broad, oblong to oblong-obovate, obtuse, the disc with 3 longitudinal calluses and
the outer 2 serrated in front but erect and forming the side of the lip below,
terminal part of the lip denticulate or entire.
Mexico, Guatemala, Costa Rica, and Panama.
CHIRIQUÍ: Volcan de Chiriquí, alt. 2875 m., Davidson 1030; Volcan de Chiriquí, alt.
3500-4000 m., Woodson & Schery 302.
Some Mexican specimens have the sepals no more than 6 mm. long, which is
unusual. The normal length is about 12 mm.
49. PLEUROTHALLIS Pravu Reichb. f. in Flora 69:555. 1886.
Masdevallia platyracbis Rolfe іп Gard. Chron. III, 4:178. 188
Pleurotballis platyracbis Rolfe іп Jour. Bot. 28:136. 1890; Hook. f. in Bot. Mag. 116:
t. 7120. 1890.
Kraenzlinella platyracbis Rolfe in Orch. Rev. 23:326. 1915.
Caespitose epiphytic herbs up to about 30 cm. tall. Secondary stems up to
about 2 cm. long. Leaves 7-20 cm. long and 1.2-3 cm. broad, ligulate-oblanceo-
late, obtuse, coriaceous, attenuated into a petiole at the base. Inflorescence up to
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19461
FLORA OF PANAMA (Orchidaceae) 125
25 cm. long, exceeding the leaves; peduncle ancipitous and winged, with one or
more bracts; rachis fractiflex, somewhat flattened or nearly terete, bracts large.
Sepals spreading, free nearly to their bases, verrucose within except at the bases,
keeled dorsally toward their apices. Dorsal sepal 13-20 mm. long and 4.5-5.5
mm. broad, lanceolate to ovate-lanceolate, acute, cucullate. Lateral sepals 13-20
mm. long and 2-4 mm. broad, lanceolate, acute, connate nearly to their apices.
Petals 4-5 mm. long and about 1 mm. broad, ligulate, falcate, verrucose externally.
Lip 5-6 mm. long and about 1.5 mm. broad, lanceolate, arcuate, acute, with a
pair of inframarginal, lamellate calluses beginning above the base and extending
nearly to the apex.
Mexico, Costa Rica, and Panama.
CHIRIQUÍ: "Chiriqui," Pfau.
Originally collected in Panama but not reported in recent years.
50. PLEUROTHALLIS SPECTABILIS Ames & Schweinf. Sched. Orch. 8:34. 1925.
Caespitose epiphytic herbs up to about 3 dm. tall. Secondary stems up to 1.5
cm. long, covered with loose, tubular sheaths. Leaves 6-13 cm. long and 0.6-1.5
cm. broad, elliptic-oblanceolate to oblanceolate, obtuse or acute, attenuated into
a petiole at the base. Inflorescence up to 30 cm. long, much overtopping the
leaves; peduncle ancipitous; rachis distichous. Sepals spreading, lanceolate, acute,
densely verrucose within except at the bases. Dorsal вера! 15-21 mm. long an
5.5—7.5 mm. broad, cucullate toward the base. Lateral sepals 15-21 mm. long
and 4—5.5 mm. broad, arcuate, connate ас the base. Petals 3-5 mm. long and
1—1.5 mm. broad, oblong-falcate, slightly broader toward the obtuse or tridentate
thickened apex, slightly verrucose. Lip 4.5—6 mm. long and about 2 mm. broad,
oblong, obtuse or slightly retuse, base of lip thin, upper 25 somewhat fleshy and
with a pair of inframarginal, lamellate calluses which extend nearly to the apex.
Costa Rica and Panama.
VERAGUAS: Santa Fé, alt. 460 m., Powell 382.
51. PLEUROTHALLIS MURICATA Schltr. in Fedde Кер. Sp. Nov. 10:293. 1912.
Pleurotballis sororoa Schltr. loc.
оке diuturna 127 іп Fedde Rep. Sp. Nov. Beih. 17:19. 1922; Ames, Sched.
Orch. 7:
Caespitose or repent, eos herbs up to about 5 dm. tall. Secondary stems
up to 5.5 cm. long, usually angled and sulcate. Leaves 8-17 cm. long and 1.2-4
cm. broad, oblong-ligulate, obtuse, coriaceous, somewhat attenuated to the base.
Inflorescence up to 23 cm. long, peduncle and rachis terete, the rachis somewhat
fractiflex and distichous, much exceeding the leaves. Sepals free nearly to their
bases, fleshy, keeled along the mid-nerve dorsally. Dorsal sepal 6-18 mm. long
and 4-7 mm. broad, broadly lanceolate, acute, cucullate. Lateral sepals 13-19
mm. long and 3-5.5 mm. broad, lanceolate, acute, cucullate. Petals 5-10 mm.
long and 1.2-2.5 mm. broad, linear-elliptic to elliptic-oblanceolate, acute, slightly
arcuate, obscurely auriculate at the base, adnate to the base of the column. Lip
(231)
[Vor. 33
126 ANNALS OF THE MISSOURI BOTANICAL GARDEN
6-10 mm. long and 2-3.2 mm. broad, unguiculate, ligulate, lobulate toward the
base; the claw short, thin; lamina fleshy, the surface slightly muricate, arcuate,
with two small, erect, lateral lobes toward the base. Ovary muricate.
Mexico to Panama, possibly Peru.
cocLÉ: vicinity of El Valle de Antón, alt. 800 m., Allen 2294, Hunter & Allen 361.
VERAGUAS: near Santiago, alt. 150—220 m., Powell 172.
52. PLEUROTHALLIS CRENATA Lindl. in Gard. Chron. 6:207. 1846; Lindl. Folia
Orch. Pleuroth. p. 38. 1859.
Masdevallia aperta Kránzl. in Fedde Rep. Sp. Nov. 17:430. 1.
Pleurothallis Hunteriana Schltr. in Fedde Rep. Sp. Nov. Vera 17:20. 1922.
Pleurothallis hamata Rolfe ex Ames, Sched. Orch. 3:8
Pleurothallis aperta Ames, Sched. Orch. 7:17, fig. 3. T
Small repent or caespitose herbs up to about 15 cm. tall. Secondary stems up
to 1 cm. long. Leaves 4.5—13 cm. long and 1-2.7 cm. broad, broadly to narrowly
oblanceolate, acute or obtuse, attenuated into a petiole at the base. Inflorescence
about half as long as the leaves, 1- to several-flowered, peduncle and rachis terete.
Sepals more or less spreading, minutely papilliferous within, keeled or winged along
the mid-nerve dorsally, the keel or wing extending onto the ovary, the apices sub-
terete; dorsal вера! 11-18 mm. long and about 3-3.5 mm. broad, lanceolate,
cucullate; lateral sepals 12-18 mm. long and 1.5—3 mm. broad, lanceolate, acute,
connate to, or nearly to, the middle. Petals 4-4.5 mm. long and 1.5-2 mm.
broad, oblong-lanceolate, somewhat pandurate, acute, slightly oblique. Lip 4.5-5
mm. long and 1.5-2 mm. broad, ob-
long-lanceolate, obtuse or acutish,
the sides erect and fleshy, disc densely
papillose-verrucose.
Mexico, Costa Rica, and Panama.
CHIRIQUÍ: without locality, alt. 1200
-1500 m., Powell 245, 277, 3436.
53. PLEUROTHALLIS FILAMENTOSA
(A. & S.) L. Wms. in Bot. Mus.
Leafl. Harv. Univ. 8:144. 1940.
poor filamentosa Ames & Schweinf.
d. Orch. 8:19, fig. 3. 1925
Small, caespitose, epiphytic herbs
up to 12 cm. tall. Secondary stems
up to about 6.5 cm. long, slender,
covered with loose, chartaceous
sheaths. Leaves 4-5.5 cm. long and
1-1.7 cm. broad, elliptic or elliptic-
lanceolate, acute, attenuated into a
short petiole at the base. Inflorescence
Fig. 116. Pleurothallis filamentosa shorter than the leaves; peduncle
(232)
1946)
FLORA OF PANAMA (Orchidaceae) 127
slender, 1-flowered, pedicel with a short filamentose appendage on the anterior side.
Dorsal вера! 13-16 mm. long and about 2 mm. broad, linear-lanceolate, the terminal
half subfiliform and the apex thickened and clavate, free nearly to the base. Lateral
sepals 14-16 mm. long and together 4-5 mm. broad, connate nearly to their apices,
puberulent within, the lamina oblong-lanceolate, the free apices acute. Petals
10-12 mm. long, about 0.5 mm. broad, subfiliform, apices thickened and clavate.
Lip about 6-7 mm. long and 2-2.5 mm. broad, 3-lobed, lanceolate-ligulate, obtuse,
short-unguiculate; the basal 14 oval, fleshy, with a small porrect lobe near the
middle on either side; terminal 25 linear-oblong, denticulate, the surface more or
less verrucose.
Panama.
CHIRIQUÍ: Cerro Horqueta, alt. 1500-1700 m., Powell 344.
Very closely allied to P. subserrata (Schltr.) L. Wms.
54. PLEUROTHALLIS SUBSERRATA (Schltr.) L. Wms. in Bot. Mus. Leafl. Harv.
Univ. 8:144. 1940. |
Restrepia subserrata Schltr. in Fedde Rep. Sp. Nov. Већ. 19:291. 1923.
Small, caespitose, epiphytic herbs up to about 15 cm. tall. Secondary stems up
to about 9 cm. long, covered with loose, chartaceous sheaths which disentegrate
with age. Leaves 4-6 cm. long and 1.5-3 cm. broad, elliptic to ovate, obtuse,
contracted into a short petiole at the base. Inflorescence as long as or exceeding
the leaves; peduncle 1 or more from each stem, slender, 1-flowered. Dorsal sepal
1.6-2 cm. long and about 0.2 cm. broad, narrowly lanceolate, with the terminal
half subfiliform and the apex clavellate, free nearly to the base. Lateral sepals
1.6-2 ст. long and together 0.5-0.65 cm. broad, connate nearly to their apices;
the lamina oblong-elliptic, the apices acute. Petals about 1.3 cm. long, 0.75 cm.
broad at the base, with a narrow, entire or denticulate lamina at the base, but the
remainder subfiliform and the apex clavellate. Lip 8-10 mm. long and 2-3 mm.
broad at the base, lanceolate-ligulate, acute or obtuse, short-unguiculate; the basal
lá oval, fleshy, with a small porrect lobe on either side near the middle; terminal
22 linear-oblong, serrulate, the surface slightly verrucose.
Costa Rica and Panama.
IRIQUI: vicinity of "New Switzerland,” central valley of Río Chiriquí Viejo, alt.
1800- “2000 m., Allen 1384; Bajo Chorro, alt. 2000 m., Davidson 116.
55. PLEUROTHALLIS XANTHOPHTHALMA (Reichb. f.) L. Wms. in Bot. Mus.
Leafl. Harv. Univ. 8:144. 1940.
ене to обрач Reichb. f. іп Hamb. Gartenzeit. 21:300.
ichb. f.", sensu Hook. in Bot. Mag. 87: 6 esp. 1861.
сақау Powellii ма а in Fedde Rep M es Beih. 17:25
Restrepia Tonduzii Schltr. loc. cit. 19: 291 3.
Caespitose epiphytic herbs up to 2 dm. tall. Secondary stems up to about 12
cm. long, slender, covered with several loose, ancipitous sheaths. Leaves 3.5—8
(233)
[Vor. 33
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN
cm. long and 0.6—3.5 cm. broad, elliptic to lanceolate to ovate, acute or obtuse,
attenuated into a short petiole at the base. Inflorescence 1- to several-flowered,
very much shorter than the leaves, rarely reaching to the middle of the subtend-
ing leaf. Dorsal sepal 10-12 mm. long and 1-2 mm. broad, linear-lanceolate,
becoming subfiliform above, with the apex clavellate. Lateral sepals 8-12 mm.
long, together 4-6 mm. broad, connate nearly to their apices, the lamina oval to
obovate. Lip 5-6.5 mm. long and 2-2.5 mm. broad, ovate-lanceolate, obtuse,
with two slender lateral lobes near the base, terminal part somewhat narrowed
and ciliate or denticulate.
Mexico, Guatemala, Honduras, Costa Rica, and Panama.
CHIRIQUÍ: “Chiriquí,” alt. 1280 m., Powell 123.
EXCLUDED OR OBSCURE SPECIES
PLEUROTHALLIS BARBAE Schltr. in Fedde Rep. Sp. Nov. Beih. 19:104. 1923.
Specimens so determined belong elsewhere.
PLEUROTHALLIS PERPUSILLA Reichb. f. in Bot. Voy. Herald, 215. 1852.
Obscure, possibly a species of Stelis.
PLEUROTHALIS PHYLLOCARDIA Reichb. f. Beitr. Orch. Centr.-Am. 97. 1866.
The species is obscure. Specimens so determined belong elsewhere.
PLEUROTHALLIS PHYLLOCARDIOIDES Schltr. in Fedde Rep. Sp. Nov. Beih. 19:193.
A plant so determined, from Darién Province, is probably an undescribed
species but the material is not good.
PLEUROTHALLIS PYROsODES Reichb. f. in Gard. Chron. п. s. 6:386. 1876; Schltr.
loc. cit. 17:22. 1922.
The specimen reported by Schlechter is P. Brigbamii S. Wats.
24. MALAXIS Solander ex Swartz
Ma axis Solander ex Sw. Nov. Gen. & Sp. Pl. Prodr. 119. 1788; in Svenska Vet.-
. Handl. 21:233. 1800; Fawc. & E Fl. Jam. 1:41. 1910.
Achroanthes Raf. in ме. Repos. М. Ү. 5:3 180
Microstylis Nutt. Сеп. Am. Pl. 2:196. pes Lindl. Gen. & Sp. Orch. РІ.
Benth. & Hook. ей РІ. 2:494. 1883; Pfitzer i in Engler & m Nu aur P
6:130. 1888; Ridley in Jour. Linn. Soc. 24:308-351.
Small, erect or repent, terrestrial or epiphytic herbs with cormous pseudobulbs
or repent stems. Section EUMALAxIs with cormous, usually hypogaeous, pseudo-
bulbs bearing 1 or 2 leaves. Section BLEPHARIGLOTTIS with more or less elongated
stems and leaves scattered along them. Leaves 1—3, continued at the base into a
sheathing petiole, membranaceous, the lamina usually broad. Scape or peduncle
slender, usually elongated. Inflorescence a spicate or subumbellate raceme; flowers
small. Sepals free or the laterals connate, subequal, spreading. Petals equal to
the sepals or usually shorter, narrow to subfiliform. Lip erect or spreading,
sessile, entire or lobed, usually cordate or auriculate, with the auricles embracing
the column, usually broader than the sepals, concave at the base (or plane in
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19461
FLORA ОЕ PANAMA (Orchidaceae) 129
Malaxis Wendlandii). Column very short, terete, usually at right angles to the
axis of the lip; anther erect or suberect, persistent, 2-celled, the cells often nearly
separate; pollinia 4, 2 in each cell of the anther, waxy, their long axis parallel to
the long axis of the column or nearly so; caudicle none.
About 200 species distributed in the temperate and tropical regions with the
greatest concentration in tropical Asia and Oceania.—A fascinating but difficult
genus, closely allied to Liparis, and with no really good characters separating the
two. The section BELPHARIGLOTTIs is entirely tropical American, and while it
resembles some species of Liparis its characters are more those of Malaxis.
8 4.5-7 “> long 10. M. TIPULOIDES
а: т > nd nee he stems.
Lip 5
bb Lip 3 атаны қ pe 9. M. BLEPHARIGLOTTIS
aa. Plants нан аса bulbous stems "вук аге пос leafy along their length
but bear S or 2 lea
b. Lea 1; j inflorescence a narrow raceme
c. Lip simple in uo not tridentate, tri 2. nor retuseé........,......... 1. M. MAJANTHEMIFOLIA
се Реде not simple іп front, either нее | trilobate, or retuse. 4
d. Lip мінді; ап 11. M. WENDLANDII
dd. Lip not obovate-triangular; anther not rostrate.
e. Lip кк» at the apex; inflorescence a dense spike................ 3. M. SouLer
rilobate in front; inflorescence a кес: raceme 2. M. PITTIERI
Lip tril
bb. basa 2; oun nce a subumbellate racem
c. -Lip сібе ог denticulate За. M. PARTHONI var.
DENTICULATA
сс. Lip neither ciliate nor denticulat
rei ignc or одн; пеаг "did
. Lip uadrat outline, ud Ва Басгојасе basal
2 а” 6. M. Хоорзомп
ee. Lip not subquadrate in outline, without linear-lanceolate
auricles
f. Lip without basal auricles 4. M. SIMILLIMA
PE Lip with E or uncinate basal auricles 5. M. EXCAVATA
dd. Lip entire in
e. Lip s d сун to transversely oblong-oval, 3.5—7 mm.
broad 8.
ee. Lip ovate-triangular to ovate-lanceolate; 1.5—2.5 mm. broad... 7. M. FASTIGIATA
. PARTHONII
1. MALAXIS MAJANTHEMIFOLIA Cham. & Schlecht. in Linnaea 6:59. 1831; Ames
& Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:121. 1935; L. Wms. in Ann.
Missouri Bot. Gard. 26:281. 1939.
Malaxis icbtbiorbynca Rich. & Gal. in Ann. Sci. Nat. ІП, 3:18. 1845.
Malaxis cocbleariaefolia Rich. g^ Gal. loc. cit
Microstylis majantbemifolia Reichb. f. in Linnses 22:834. 1849.
Microstylis cochleartaefolia Reichb. f. loc. cit.
ы ias al ыш н Reichb. f., Beitr. Orch. Centr.-Am. 99. 1866.
ant bemifolia Greene, Pittonia 2:184. 1891.
Microst-ylis eue Schltr. in Fedde Rep. Sp. Nov. 3:17. 1906.
Small, erect, terrestrial herbs from 1 to 4 dm. tall. Stem short, cormous or
flehy, covered with scarious sheaths. Leaves long-petiolate, one to each plant;
lamina 2.5-12 cm. long and 2—9.5 cm. broad, ovate-cordate to broadly ovate-
cordate, acute or obtuse, membranaceous; petiole elongated, sheathing and enclosing
the peduncle for half its length (thus the lamina seemingly borne near the middle
of the otherwise naked part of the peduncle). Inflorescence an elongate, many-
flowered raceme. Dorsal sepal 2.5-3 mm. long and 1-1.5 mm. broad, elliptic-oval
(235)
[Vor. 33
130 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
to elliptic-ovate, obtuse ог acutish. Lateral sepals 2.3-2.8 mm. long and about
1.5 mm. broad, oval, obtuse or acute, slightly arcuate, shortly connate at the
base. Petals 2-3 mm. long and 0.3—0.7 mm. broad, linear-lanceolate, acute,
slightly arcuate. Lip 2-3 mm. long and 1.5-2.5 mm. broad, ovate-cordate or
triangular-cordate, acute or obtuse, somewhat concave. Anther bipartite with 2
fused pollinia in each part of it.
Mexico, Guatemala, ipsus and Panama.
cHIRIQUÍ: Boquete, alt. 1515 m., Davidson 801; Casita Alta, Volcán de Chiriqui, alt.
1500—2000 m., Woodson, Alles. 6 Seibert 830; Potrero Muleto de Volcán de Chiri iquí,
alt. 3500-4000 m., Woodson © Schery 474.
2. Maraxis PITTIERI (Schltr.) Ames in Proc. Biol. Soc. Wash. 35:84. 1922.
Microstylis Pittieri Schltr. in Fedde Rep. Sp. Nov. 12:203. 1913.
Erect terrestrial herbs up to about 2 dm. tall. Stems short, fleshy or cormous,
unifoliate. Leaves long-petiolate; lamina 2-3 cm. long and about 2-2.5 cm.
broad, broadly ovate-cordate, acute or short-acuminate, membranaceous; petiole
sheathing the lower half of the peduncle. Inflorescence a sublax, many-flowered
raceme up to about 9 cm. long. Dorsal вера! 2.2-2.7 mm. long and about 1 mm.
broad, elliptic-oblong, obtuse. Lateral sepals about 2-2.3 mm. long and together
about 1.7 mm. broad, suborbicular-ovate, connate 34 their length. Petals
about 2.5 mm. long and 0.5 mm. broad, linear, acute or obtuse. Lip 2.2-2.5 mm.
long and 2-2.5 mm. broad, 3-lobed, the disc cochleate and with a carinate U-
shaped ridge in front; mid-lobe triangular-lanceolate; lateral lobes oblong-ovate.
Known only from Panama.
cHIRIQUÍ: between Alto de Las Palmas and the top of Cerro de la Horqueta, alt.
2100-2268 m., Pittier 3277.
A rare species, distinctive because of the lobing of the lip.
3. Maraxis Souter L. Wms. іп Ann. Missouri Bot. Gard. 21:343. 1934.
Microstylis montana Rothr. in Wheeler, Rept. О. S. Geog. Surv. W. 100th Меп.
6 (Bot.) :264. 187
Achroanthes montana Greene, Pittonia 2: ee 1891.
Malaxis montana O. Ktze. Rev. Gen. Pl. 2: ye: 1891, non Blume.
Malaxis venerit "(La Llave & Lex ve . Ku ntze,” sensu Ames & Schweinf. in Bot.
Mus. Leafl. = arv. Univ. 3:120. 1935, Һа я of La Llave & aun nor of Lindley
nor of Kuntz
Erect terrestrial herbs up to about 4.5 dm. tall. Stems short, fleshy or cormous,
probably hypogaeous, covered with the scarious bases of the sheaths and petiole.
Leaves 1 on each growth; lamina 2.5-16 cm. long and 1-6 cm. broad, elliptic-
ovate to oblong or oblong-ovate to obovate, variable, acute or obtuse, mem-
branaceous; petiole elongated and enclosing the peduncle for a third to a half of
its length and thus the lamina borne well above the ground. Inflorescence an
elongated, densely flowered spike. Dorsal вера! 1.5-2.5 mm. long and about
0.8—1.2 mm. broad, lanceolate-ovate to elliptic-oblong, acute or obtuse. Lateral
sepals 1.5-2.2 mm. long and 0.8—1.2 mm. broad, oblong-elliptic, obtuse. Petals
(236)
19 46]
Fig.
FLORA OF PANAMA (Orchidaceae)
Ру 252.
Ss
oer,
Lil P
ser:
ene
«210409 т
ANS AT db s".
ЛЗ
D %
ie
V,
Q
117. Malaxis Soulei
(237)
131
[Vor. 33
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1.2-2 mm. long and 0.2-0.6 mm. broad, linear or linear-lanceolate, obtuse. Lip
1.7-2.5 mm. long and 1.4-2.5 mm. broad, ovate-cordate to suborbicular-cordate,
auriculate, cochleate, apex tridentate.
Arizona, Mexico, Guatemala, Costa Rica, and Panama.
CHIRIQUÍ: Volcan de оа alt. about 3025 m., Davidson 000; Llanos del Volcan,
E 1120-1200 m., Seibert 128; Casita Alta, Volcán de Chiriquí, alt. about 1500-2000
‚ Woodson, Allen 9 vi 804, 835; Casita Alta to Cerro Copete, alt. A de m.,
Fio 8 Schery 371
4. MALAXIS SIMILLIMA (Reichb. f.) O. Ktze. Rev. Gen. Pl. 2:673. 1891.
Microstylis simillima Reichb. f. Beitr. Orch. Centr.-Am. 101. 1866.
Erect terrestrial herbs up to about 3.5 dm. tall. Stems short, cormous, hypo-
gaeous, bifoliate, covered with the scarious bases of the sheaths and petiole. Leaves
long-petiolate; lamina 3.5-14 cm. long and 2-7 cm. broad, lanceolate-ovate to
broadly ovate, acuminate, membranaceous, reaching to about the middle of the
peduncle. Inflorescence a short, dense, many-flowered, subumbellate raceme.
Dorsal вера! 3-4 mm. long and 1.2-1.8 mm. broad, broadly lanceolate or ovate-
lanceolate, acute or obtuse. Lateral sepals 3-4 mm. long and 1-1.5 mm. broad,
ligulate-lanceolate to elliptic-lanceolate, acute, connate to the middle or less.
Petals 2-2.5 mm. long and 0.2-0.5 mm. broad, linear, acute or obtuse. Lip
3.5—4.5 mm. long and 1.5—3 mm. broad, ovate-lanceolate, acute, cochleate, with
2 small lateral or inframarginal teeth below the apex.
Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, alt. 1900 m., Woodson & Schery 685.
MaLaxis EXCAVATA (Lindl) O. Ktze. Rev. Gen. Pl. 2:673. 1891; L. Wms.
in Bot. Mus. Leafl. Harv. Univ. 6:75. 1938.
Microstylis excavata Lindl. in Bot. Reg. 24:Misc. p. 51. 1838.
Microstylis bastilabia Reichb. f. Beitr. Orch. Centr. Asp. 101. 1866.
Malaxis bastilabia O. Ktze. loc. cit.
Microstylis Carpinterae Schltr. in Beih. Bot. E epum 36, Abt. 2:381. 1918.
Malaxis Carpinterae Ames, Sched. Orch. 7:157
Malaxis uncinata Ames & ee Sched. ote. 2 15. 1930.
мл
Erect terrestrial herbs from а rhizome, up to about 4.2 dm. tall. Stems short,
cormous, bifoliate, covered with the scarious sheaths of the petioles and bracts.
Leaves long-petiolate; lamina 5-12 cm. long and 2-7 cm. broad, ovate-lanceolate
to ovate, acute or acuminate, membranaceous, paired, reaching to about the middle
of the peduncle; petioles sheathing the lower part of the peduncle. Inflorescence
a short, dense, many-flowered, subumbellate raceme. Dorsal sepal 3.5-4 mm.
long and 1.5-2.7 mm. broad, elliptic-oblong to ovate, obtuse. Lateral sepals 3-4
mm. long and 1.3-2.2 mm. broad, oblong-oval to oblong-ovate, obtuse, free to
the base or nearly so. Petals 2.5—3.5 mm. long and 0.3-0.4 mm. broad, linear (or
linear-lanceolate), obtuse. Lip 3-4 mm. long and 2.5-4 mm. broad, oblong-
cordate to triangular-cordate, auricles at the base erect or uncinate, trilobulate at
the apex, disc excavated and gibbous, the cavity with a longitudinal ridge.
(238)
19461
FLORA OF PANAMA (Orchidaceae) 133
Mexico, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Argentina,
and Brazil.
CHIRIQUÍ: Boquete, alt. about 3025 m., Davidson 1002; vicinity of Monte Lirio, их
1300-1900 m., Seibert 175; Casita Alta to Cerro Copete, alt. 2300-3300 m., Woods
& Schery м. Potiero Muleto to summit of Volcan de Chiriqui, alt. 3500-4000 m., Wood.
son & Scher
Dam drawn from Central American material. Some South American
specimens have smaller flowers. А variable species.
Е Ж E о
= иу
Fig. 118. Malaxis Woodsonii
6. Maraxis Woopsonn Г. Wms. in Ann. Missouri Bot. Gard. 26:281, 2. 21,
figs. 1-2. 1939.
(239)
[Vor. 33
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Small terrestrial herbs up to about 15 cm. tall. Stems short, cormous, covered
with the bases of the petioles and bracts. Leaves long-petiolate; lamina 1.5—5.5
cm. long and 1.3—4.5 cm. broad, broadly ovate, obtuse or acute, crenulate or
obscurely serrate, membranaceous, borne about the middle of the peduncle. In-
florescence many-flowered, subumbellate. Sepals 2.5-4 mm. long and 1.5-2.5 mm.
broad, broadly lanceolate, obtuse. Petals about 2.5-3 mm. long, filiform. Lip
3.5-5 mm. long and 3-3.5 mm. broad, quadrate in outline; apex 3-lobate, the
mid-lobe small and exceeded by the lateral lobes in length, lateral lobes large,
rounded, obtuse, the basal auricles linear-lanceolate, acute, erect; disc with two
shallow cavities.
Panama.
IRIQUÍ: Casita Alta, Volcán de Chiriqui, alt. about 1500-2000 m., Woodson, Allen
8 Seibert 831,
Well distinguished from all other species in Central America by the basal
auricles of the lip.
7. MaLaxis FASTIGIATA (Reichb. f.) О. Ktze. Rev. Gen. Pl. 2:673. 1891; Ames
& Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:116. 1935.
rwr fastigiata Reichb. f. in Linnaea 22:834. 1849.
Microstylis brachyrrhynchos Reichb. f. in Flora 71:152.
Malaxis ries шшш] Si а Ames in Proc. Biol. Soc. Wash. 35: 84. 1922, sphalm.
Malaxis Lankesteri Ames, Sched. Orch. 4:7. 1923
Small erect terrestrial herbs up to about 6 dm. tall. Stems short, cormous,
bifoliate, tunicate. Leaves long-petiolate; lamina 3—16.5 cm. long and 1.5-9.5
cm. broad, oval to broadly ovate, obtuse or acuminate, membranaceous, paired,
reaching to about the middle of the peduncle; petiole sheathing the lower part
of the peduncle. Inflorescence a many-flowered, short or rarely somewhat elon-
gated, subumbellate raceme. Dorsal sepal 3-5 mm. long and 1-1.6 mm. broad,
lanceolate to ovate-lanceolate, obtuse. Lateral sepals 3-4.5 mm. long and 1-1,8
mm. broad, lanceolate to elliptic-ovate, obtuse, free to their bases. Petals 2—3.5
mm. long and 0.3—0.4 mm. broad, linear or linear-ligulate. Lip 2-4 mm. long
and 1.5-2.5 mm. broad, ovate-triangular to ovate-lanceolate, the apex from
rounded to subrostrate, acute or obtuse, disc excavated and gibbous, fleshy, the
cavity with a longitudinal ridge.
Mexico, Guatemala, Honduras, Costa Rica, and Panama.
CHIRIQUÍ: Llanos del Volcán, alt. 1120-1200 m., Seibert 327.
Malaxis fastigiata and M. brachyrrhyncha are often considered to be distinct
species. The first is supposed to have the lip distinctly rostrate; the second is not
supposed to be rostrate (although the type is short-rostrate). This seems to be
the only method of separating the two and, as the two conditions intergrade
completely, the two species should be considered synonymous.
8. Maraxis Равтноми Morr. in Bull. Acad. Roy. Brux. 5:485. 1838; L. Wms.
in Ann. Missouri Bot. Gard. 26:281. 1939.
Microstylis bistionantba Lk., Kl. & Otto, Ic. Pl. Rar. Hort. Berol. 1:11, #. 5. 1841; Hook.
(240)
A ӨЗ
19461
FLORA ОЕ PANAMA (Orchidaceae) 135
in Bot. Mag. 70: f. а 1844.
Microstylis Part bonii R ichb. f. in Walp. Ann. 6:206. 1861.
Microstylis Brenesii hie in Fedde Rep. Sp. Nov. Beih. 19:167. 1923.
Erect terrestrial herbs up to about 6 dm. tall. Stems short, cormous, bifoliate,
covered with the scarious bases of the petioles and bracts. Leaves 8—18 cm. long
and 3-12 cm. broad, ovate-lanceolate to broadly ovate or suborbicular, acute or
acuminate, membranaceous, paired, reaching to about the middle of the scape,
their petioles sheathing the lower part of the peduncle. Inflorescence a short,
many-flowered, subumbellate raceme. Dorsal вера! 4.5-6 mm. long and 1.5-2.5
mm. broad, lanceolate-ligulate to elliptic, obtuse. Lateral sepals 3-5 mm. long
and together up to 6 mm. broad, connate to the middle or beyond or free to the
base, suborbicular in outline or separately lanceolate-ovate, obtuse. Petals 2-4
mm. long, filiform. Lip 3.5-5 mm. long and 3.5-7 mm. broad, orbicular to
transversely oblong-oval, concave toward the base.
Mexico, Salvador, Costa Rica, Panama; reported from Peru but а small-
flowered form if so.
ONE: vicinity of Salamanca Hydrographic Station, alt. about 80 m., Woodson,
Allen 8 Seibert 1581. CHIRIQUÍ: Finca Lérida to Boquete, alt. 1300-1700 m., Woodson,
Allen & Seibert 1172.
8а. Mavaxis РАКТНОМП Morr. var. DENTICULATA (Reichb. f.) Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:40. 1934
шар ш Parthonii Reichb. f. var. denticulata Reichb. f., Beitr. Orch. Centr.-Am. 100.
он Wercklei Schltr. іп Beih. Bot. Centralbl. 36, Abt. 2:382. 1918.
Malaxis Wercklei Ames in Proc. Biol. Soc. Wash. 35:85. 1922
Much like the species except the floral parts smaller, the lip denticulate or
ciliate and the cavity at the base of the lip definitely bipartite.
Costa Rica and Panama.
CHIRIQUÍ: vicinity of upper Río Chiriquí, Monte Lirio, alt. 1300—1900 m., Seibert
170, 221; valley of upper Rio Chiriqui, alt. 1300-1900 m., White 9 White 17; vicinity
of Casita Alta, alt. 1500-2000 m., Woodson, Allen 8 Seibert 832bis.
9. MALAXIs BLEPHARIGLOTTIS (Schltr.) Ames in Proc. Biol. Soc. Wash. 35:84.
T1922;
Microstylis blepbariglottis Schltr. in Fedde Rep. Sp. Nov. 12:202. 1913.
Small, erect, epiphytic (or terrestrial?) herbs up to about 4.5 dm. tall. Stems
up to about 2 cm. long, densely leaved. Leaves 4.5—12 cm. long and 0.8—1.5 cm.
broad, elliptic to oblanceolate-ligulate, acute or acuminate, membranaceous. In-
florescence densely flowered, elongated; bracts up to about 8 mm. long, lanceolate,
acuminate, entire or lacerate, cucullate. Sepals 2.5-3 mm. long and 1-1.5 mm.
broad, elliptic-lanceolate to elliptic-oblong, obtuse, slightly cucullate. Petals 3-3.5
mm. long and 0.4—0.6 mm. broad, linear-ligulate, obtuse. Lip 3—4 mm. long and
2-3 mm. broad, oblong-obovate, obtuse or acute, slightly constricted above the
base, the base emarginate and callus thickened, margins ciliate or ciliate-lacerate.
Costa Rica and Panama.
(241)
[Vorn 33
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN
cHIRIQUÍ: above El Boquete, alt. 1650-1925 m., Maxon 5667.
Known in Panama only from the original specimen although the locality has
been often collected. This species, along with Malaxis tipuloides (Lindl.) O.
Ktze., belongs to Schlechter’s section BLEPHARIGLOTTIS of Microstylis (Schlechter,
loc. cit. 203). The species of the section (perhaps a dozen) resemble some species
of Liparis rather closely. The anthers are neither completely erect, as they are
supposed to be in Malaxis, nor are they incumbent as in Liparis. The position of
the anther in these genera is controlled by the length of the rostellum, and this
species has a relatively long one. It is perhaps important that the pollinia of
these two species have their long axis parallel to the long axis of the column,
which doubtless indicates an anther nearly erect. The position of the anther has
been used in separation because there is hardly a way in which the two genera
may be distinguished. Perhaps the relative length of the column would afford a
better criterion.
10. Maraxis TIPULOIDES (Lindl.) O. Ktze. Rev. Gen. Pl. 2:673. 1891, as Malaxis
tipulodea.
Microstylis tipuloides Lindl. in Ann. & Mag. Nat. Hist. ~, 256. 1845.
Liparis eustachys Schltr. іп Fedde Rep. Sp. Nov. Beih. 19:91. 1923.
Liparis fratrum Schltr. loc. cit. 92; L. Wms. in Ann. Missouri Bot. Gard. 27:282. 1940.
Liparis pee Schltr. loc. cit. 91; C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 6:2.
1938
Repent or ascending terrestrial herbs often with the stems rooting at the
nodes, up to about 7 dm. long. Stems up to about 10 cm. long, densely leaved,
slender. Leaves 5-17 cm. long and 0.9—3.5 cm. broad, elliptic to elliptic-oblance-
olate, petiolate, acute, membranaceous. Inflorescence a loose or dense raceme up
to about 5-6 dm. long, much exceeding the stems and the leaves. Dorsal вера!
5.5-6 mm. long and 1-2 mm. broad, ligulate, lanceolate, obtuse. Lateral sepals
similar to the dorsal but somewhat arcuate. Petals 4.5-7 mm. long, linear or linear-
lanceolate, obtuse or acute. Lip 5.5-9.5 mm. long and 3-4.5 mm. broad, elliptic-
obovate, acuminate, auriculate, ciliolate, somewhat constricted below the middle.
Costa Rica, Panama, and Colombia.
CHIRIQUÍ: Cerro Punta, alt. 2000 m., Allen 1528.
See note on preceding species.
11. Maraxis Wendlandii (Reichb. f.) L. Wms., comb. nov.
Liparis Wendlandii Reichb. f. Beitr. Orch. Centr.-Am. 98. 1866.
Small, caespitose, epiphytic herbs up to about 6 dm. tall. Stem a short cormous
pseudobulb covered with fibrous sheaths and the bases of the petioles and bracts.
Leaf 1, petiolate; lamina 0.9—2 cm. long and 0.5-1.2 cm. broad, oval to sub-
orbicular, acute, membranaceous. Inflorescence a few-flowered raceme, the base
enfolded by the petiole and the bracts. Dorsal sepal about 3—3.5 mm. long and
1.5-1.7 тт. broad, ovate-lanceolate, obtuse or acutish. Lateral sepals 2.7-3.2
(242)
1946]
FLORA OF PANAMA (Orchidaceae) 137
mm. long and about 0.8—1.2 mm. broad, lanceolate to ovate-lanceolate, obtuse or
acutish, somewhat arcuate. Petals about 3 mm. long and 0.6-0.7 mm. broad,
linear-lanceolate, acute or acuminate. Lip 3—3.5 mm. long and 2-2.6 mm. broad,
obovate-triangular, retuse, denticulate, with an umbonate callus toward the base.
Column about 0.75 mm. long, at right angles to the lip; anther about 0.75 mm.
long, erect or semi-erect, long-rostrate, the rostrum about 0.5 mm. of the length
of the anther; rostellum very long.
Costa Rica and Panama.
CHIRIQUÍ: valley of the upper Río Chiriquí Viejo, vicinity of Monte Lirio, alt.
1300—1900 m., Seibert 195, 216.
Malaxis Wendlandii is a very curious plant which seems to have most of the
technical characters of Malaxis but a lip like that of Liparis, in which genus it
was originally placed. Тһе short column, erect or semi-erect anther with sep-
arated cells and the long rostellum point to Malaxis. The rostrate anther is
unusual. The shape of the lip certainly reminds one of the genus Liparis.
25. LIPARIS L. C. Rich.
Lrearis L. С. Rich. in Мет. Mus. Par. 4:52, t. 5, fig. 10. 1818; Lindl. Gen &
Sp. Orch. Pl. 26. 1830; Benth. & Hook. Gen. Pl. 2:495. 1883; Ridley in
Jour. Linn. Soc. 22:252-297. 1886.
Small terrestrial or epiphytic herbs with pseudobulbous stems. Leaves few
(1-4), petiolate, the base of the petiole enclosing the pseudobulb; lamina mem-
branaceous (in Panama), usually broad. Inflorescence a few- to several-flowered,
terminal raceme. Flowers small. Sepals free, subequal, spreading. Petals sub-
equal to the sepals or usually smaller and narrower. Lip erect or spreading,
shortly adnate to the base of the column or free, sessile or usually shortly clawed,
entire or usually emarginate, 1- to 2-tuberculate at the base, broader than the
sepals. Column elongated (compared to that of Malaxis), arcuate, semi-terete,
footless, usually parallel to the axis of the lip; anther terminal, operculate, in-
cumbent, usually soon deciduous and versatile; pollinia 4, waxy, their long axis
at right angles to the long axis of the column or nearly so.
About 200 species in tropical and temperate regions with the greatest con-
centration in tropical Asia and Oceania. Closely allied to Malaxis.
1. LIPARIS ELATA Lindl. in Bot. Reg. 14: /. 1175. 1828; Г. Wms. in Ann. Mis-
souri Bot. Gard. 26:282. 1939.
Terrestrial herbs from 13 to 40 cm. tall. Stems short, becoming fleshy or
cormous, sheathed with the bases of the petiole. Leaves petiolate, usually 3 or 4
from each stem, rosulate; lamina 5-30 cm. long and 2.5-11 cm. broad, elliptic-
oval to ovate, acute or acuminate, membranaceous. Inflorescence a more or less
lax, few- to many-flowered raceme. Dorsal вера! 6-7 mm. long and 2-2.2 mm.
broad, oblong-lanceolate, obtuse. Lateral sepals 4-6 mm. long and 2-3 mm.
broad, oval to ovate, obtuse, arcuate. Petals 5-5.7 mm. long and 1-1.5 mm.
(243)
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
138
Liparis elata
(244)
Fig. 119.
19461
FLORA OF PANAMA (Orchidaceae) 139
broad, linear to linear-oblanceolate, obtuse, arcuate. Lip 4-5.5 mm. long and
4-5 mm. broad, obovate to flabellate, truncate or emarginate, fleshy, bituberculate
at the base. Column long, arcuate.
Florida, Mexico, Guatemala, Honduras, Costa Rica, Panama, the West Indies,
Brazil, Colombia, and Peru.
CANAL ZONE: Quebrada Salamanca, alt. 70 m., bobo: Steyermark & Allen 16988;
Salamanca Hydrographic Station, Río Pequeñí, alt. роне 80 m., Woodson, Allen & Seibert
1580. PANAMA: Cerro Campana, alt. 600-800 m., Allen 2677; foothills east of city,
Powell 243.
A widespread and somewhat variable species.
26. HEXISEA Lindl.
HexisEa Lindl. in Hook. Jour. Bot. 1:7. 1834.
Eutbonaea Reichb. f. in Bot. Zeit. 10:772. 1852.
Caespitose epiphytic herbs. Stems simple or branched, terete or angled, seg-
mented, thickened or pseudobulbous. Leaves few, usually one or two from the
apex of each segment, usually deciduous annually. Inflorescence from the apex
of the growths, few-flowered, short; flowers small but highly colored. Sepals
similar, erect or spreading, narrow, the laterals often shortly connate at the base.
Petals similar to the sepals. Lip simple or lobulate, adnate at the base with the
column, base of the lamina geniculate and usually callus-thickened. Column
short, adnate to the lip below the middle; clinandrium 3-lobed; anther operculate,
incumbent; pollinia 4, collateral, waxy.
About a half dozen species in Central and South America. Living material
should prove instructive as to the origin of the inflorescence and new growth, for
both apparently come from the apex of the preceding growth, which is somewhat
unusual. The monotypic genus Alemania Llave & Lex. (The only one of the Llave
& Lexarza orchids for which a type is known to exist) should possibly include
Hexisea. Florally they are not to be distinguished but the vegetative growth may
keep them separate until more is known about them.
1. НЕХБЕА BIDENTATA Lindl. in Hook., Jour. Bot. 1:8. 1834.
Diotbonaea imbricata Lindl. Sert. Orch. 7. 40, f. I
Epidendrum oppositifolium Rich. & Gal. in Ann. Sci, ‘Nat E^ 3:21. 1845.
Diotbonaea oppositifolia Reichb. f. in Linnaea 22:842. 18
at е» Reichb. f. in = Zeit. 10:772. Коу
mbricata Reichb. f. loc.
Heer oppositifolia Reichb. f. in Walp. Ann. 6:470. 1862.
Hexisea imbricata Reichb. f. loc. cit.
Caespitose epiphytic herbs up to about 4 dm. tall. Stems segmented, the
terminal growth producing flowers and then a new growth and leaves from its
apex, simple or branched, the segments subcylindric or fusiform. Leaves 2.5-16
cm. long and 0.3-1 cm. broad, linear to linear-ligulate, obtuse or retuse, cori-
aceous, borne from the apex of the segment of the stem. Inflorescence a short,
(245)
[Vor. 33
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN
few-flowered raceme. Flowers usually bright red. Sepals 10-14 mm. long and
2.5—4 mm. broad, lanceolate or elliptic-lanceolate, acute, the laterals somewhat
oblique. Petals 10-13 mm. long and 2-3 mm. broad, linear-lanceolate to linear-
oblong, acute. Lip 8—13 mm. long and 2.5—4 mm. broad, oblong-oblanceolate,
obtuse or acute, base of the lamina geniculate and the angle or "knee" usually
callus-thickened, claw adnate with the basal half of the column.
Mexico, Honduras, Nicaragua, Costa Rica, Panama, Colombia, and British
Guiana.
CHIRIQUÍ: “Chiriqui,” alt. 1200 m., Powell 64, 3113, 3156, 3157, 3161, 3100, 3102,
3214, 3504. PANAM MA: San Juan Hills, е ast of Panama ea Powell 333, 3146, 3420;
"Panama": Panama et Colombia occidentalis, in 1831, Cuming 1207. ҮЕКАСОАЅ: head-
waters of the Río Cafazas, alt. 300-600 m., Allen 195; Bahia "Honda, Taylor 1509.
The genus should possibly be referred to Alemania Llave & Lex.
(To be concluded in Part III, Fasc. 3)
(246)
Annals
of the
Missouri Botanical Garden
Vol. 33 APRIL, 1946 No. 2
CONTRIBUTIONS TO OUR KNOWLEDGE OF AMERICAN
CARBONIFEROUS FLORAS
ҮШ. ANOTHER MEDULLOSA FROM IOWA
HENRY N. ANDREWS
AND JULES A. KERNEN?
In the last number of this ‘Carboniferous Flora" series (Andrews, 745) it was
mentioned that xylary fragments of a rather large Medullosan stem were present
in a collection of coal balls received from Mr. Frederick O. Thompson, of Des
Moines, Iowa; and that while they apparently represented a distinct species they
were not sufficiently complete to warrant description. А few weeks after this
was published Mr. Thompson sent us another collection of coal balls which in-
cluded an interesting Medullosa, and this specimen consists of a stelar system that
appears worthy of its own alcove in the archives of pteridosperm stem remains.
Perhaps the most disturbing feature connected with coal ball studies is the
necessity of casting a decision occasionally as to when to stop and record newly
acquired information. One must necessarily draw a line somewhere between the
extremes of describing each fragment as it turns up or of waiting indefinitely for
a more or less complete set of fossils to accumulate. Unfortunately, the latter
may never happen. А statement of Dr. Cannon's in his recent book (Cannon,
'45), that "Since no research can be final, every publication is a progress report,"
seems most appropriate to coal-ball investigations.
The Iowa coal-ball plants previously described (Andrews, '45) were obtained
from the Urbandale mine, the location of which is noted on the accompanying
map (text-fig. 1). The present specimen, however, was collected from the Shuler
coal mine a few miles to the west of the Urbandale mine. The exact location of
the Shuler shaft is: 8 miles west of 63rd Street (Des Moines) and 1.2 miles north
of U. S. Highway No. 6. The coal vein lies 610 feet above sca level, and 387
feet beneath the surface of the ground which has an elevation of 997 feet.
! Student at Southwest High School, St. Louis.
Issued April 30, 1946.
(141)
[Vor. 33
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Some question has prevailed as to whether the veins worked by the Shuler and
Urbandale mines are continuous. Recently we have been informed by Mr. Charles
Bendixen, an official of the Shuler Mine, that drill holes show that the Shuler coal
disappears entirely to the east toward the Urbandale mine. There is also some
evidence to indicate that the Des Moines Ice and Fuel Co. coal does not extend to
that worked by the Shuler mine. However, in view of the general similarity of
the fossil floras from the two mines (Shuler and Urbandale), the fact that the
shafts are but six and one-half miles apart and that there is a difference of only
ten feet in the elevation of the veins at the position of the respective shafts, it
seems likely that they represent contemporaneous Pennsylvanian forests. А more
precise statement must await the publication of recent stratigraphic studies.
Inasmuch as we may reasonably expect more paleobotanical contributions from
this region the accompanying map is presented to show the exact geographical
position of the productive mines.
Medullosa anglica var. ioensis, var. nov.
The single specimen available consists of a stelar system composed of three
steles (pl. 1, fig. 1). Although slightly crushed, they were all of approximately
the same size in life. The best-preserved one (fig. 2) measures 18 x 10 mm. in
transverse section. "These measurements are limited by the secondary xylem, the
tissues external to which (except for the partly preserved secondary phloem) being
absent. The primary xylary body of this stele is somewhat crushed and distorted
but apparently measured about 2 mm. in diameter in life. The radial development
of the secondary xylem in two of the steles is fairly uniform, while the third
(fig. 1c) displays pronounced endocentricity. The rather weak development
and loose organization of the xylem on one side of stele a (fig. 1) is apparently
the result of stelar branching.
Although the endocentric stele is crushed, the primary xylary body is, for the
most part, intact (fig. 3). Тһе relative amount of parenchyma associated with
the primary tracheids is markedly greater than that in either M. anglica or M.
Thom psonii. Yn the latter the primary body consists almost entirely of tracheids,
while in M. anglica there is a somewhat greater admixture of parenchyma. Ас-
cording to Scott's account of the English species: ‘Тһе whole interior of each
stele . . . . is occupied by the primary wood, which consists of elements of two
kinds—tracheides and parenchymatous cells. The arrangement and relative pro-
portions of the two constituents vary much in different steles, and in different
parts of the same. At some places, the tracheides form a continuous system over
a considerable area, only interrupted by scattered strands of parenchyma... . In
other cases the tracheides are arranged in definite groups, separated from one an-
other by a network of parenchyma... ., while sometimes the cellular tissue pre-
dominates, and is traversed by comparatively few tracheides, isolated, or in small
groups.... In no case is there any differentiation of a true ‘partial pith,’ or even
any marked increase in the amount of parenchyma towards the middle of the
stele.” [Scott, 799, p. 88].
1946)
143
ANDREWS & KERNEN—AMERICAN CARBONIFEROUS FLORAS. VIII
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Text-fig. 1.
Map showing coal mines near Des Moines, Iowa, where coal balls have been found.
By Frederick O. Thompson.
[Vor. 33
144 ANNALS OF THE MISSOURI BOTANICAL GARDEN
It is evident from Scott’s description that the primary xylem in the steles of
M. anglica presents appreciable variation with respect to the tracheid-parenchyma
ratio. In view of the size variation in the steles this might be expected according
to Bower’s (730) size and form principles. However, іп the three major steles
composing the stem of М. anglica, the primary bodies appear (Scott, 799, pl. 6,
photo 5) to be of about the same areal size and the tracheid-parenchyma ratio
seems to be uniform in all. Comparing that figure with a corresponding one of
М. anglica var. ioensis, two features are noticeable: first, the primary body in
the steles of the Iowa specimen is much crushed which in itself would suggest
relatively more of the delicate thin-walled conjunctive parenchyma, and second,
where the primary wood is well preserved (fig. 3) the relative paucity of tracheids
is conspicuous. The crushed nature of the primary tissues of M. anglica var.
іоепзік makes it impossible to make an accurate size comparison with М. anglica.
The radial walls of the secondary tracheids are composed of closely compacted
and, for the most part, irregularly arranged bordered pits (fig. 5), an organization
so characteristic of many supposed pteridosperms. Individually the pits vary
from oval to an irregularly angular outline where they are most closely compacted.
Occasionally they are nearly hexagonal, tending to be arranged in vertical rows.
In a few cells the pit orifices are remarkably well preserved, being slender, hori-
zontally orientated slits (fig. 4).
The wood rays, which may be classified in group IIB (Andrews, '40), are very
narrow, being one to three cells wide, and of great height, exceeding 6. mm.
Associated with the steles, although not organically connected, are two small
segments of the outer cortical rind of a Medullosan stem or petiole. While the epi-
dermis of these fragments is missing, the general organization of the fibrous strands
and of the enclosing parenchyma resembles that of M. anglica. In one of the two
fragments secretory canals are especially abundant, abutting on nearly every
fibrous strand, and present an appearance not unlike that of Myeloxylon Bendixenii
(Andrews, '45, pl. 7).
Locality: Shuler Coal Mine, Des Moines, Iowa.
Horizon: Des Moines Series, Pennsylvanian.
Type specimen: No. WCB425, Henry Shaw School of Botany, paleobotanical
collections.
Discussion.—
There are now some eight species and varieties assigned to the Anglorota section
of Medullosa. Whether or not these will all retain their present taxonomic dis-
tinction in the light of a future monographic study is a decision that we are not
obliged to meet at the moment. There already exists a most interesting series of
stems displaying significant, although more or less intergrading, anatomical varia-
tions. Some of these are perhaps only physiological variations while others appear
to carry more taxonomic weight. Until more information has accumulated con-
19461
ANDREWS & KERNEN—AMERICAN СА JUS PLORAS: ҮШ 145
= = —
798 nm
— .- --- —
Text-fig 2. А diagram showing the (somewhat restored) stelar system of Мей оза
nglica v е > primary xylem is indicated in solid black, the secondary xylem
by radiating lines, mi ine — position of the stelar periderm by the dotted line. This
diagram has been red at a magnification of x 2.8 to facilitate comparison with other
ibe of the PL a group shown in pl. 12 of Part VII of this series (Andrews, '45).
cerning this most important Pennsylvanian genus it seems most expedient to take
advantage of even relatively slight structural differences.
In the account of Medullosa Tbompsonii considerable weight was placed on
the anatomy of the cortical tissues, particularly the fibrous strand-secretory canal
relationship. However, since this tissue is lacking in the present specimen we are
[ Vor. 33, 1946]
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN
handicapped in making a precise comparison. If the two associated cortical
fragments belong with the stelar system they present characters, correlating with
those of the primary wood, that clearly separate them from Medullosa Thom psonii.
In both of these characters the new stem lies closest to the English M. anglica.
The qualitative differences separating the species of this group as а whole are
by no means striking yet there are good reasons for believing that, such as they
are, they may be of sound taxonomic validity. First, it is well known that stem
anatomy in general is of а more conservative nature than that of other plant
organs such as foliage or reproductive structures. Second, the Iowa coal balls
contain a considerable variety of seeds, a number of which are, in all probability,
those of pteridosperms.
It is a trifle embarrassing that the present report had to be given separately so
soon after the previous account of Iowa Medullosas. On the other hand, we are
not averse to hoping that future discoveries will render possible a continuation of
these reports regardless of how long or short the intervals may be.
Acknowled gment.—
Thanks are again due to Mr. Frederick О. Thompson for his vital and con-
tinued interest in furthering these investigations.
LITERATURE CITED
ide a = N. 2. hes the stelar 2. > the pteridosperms with particular reference to
condary wood. . Mo. Bot. Gard. 1-118.
5 5-2 to our пи of American carboniferous floras. УП. Some
eridosperm stems from Тома. Ann. Mo. Bot. Gard. 32:323-360
1 . Size i
s.
Cannon, W. B. (1945). The way of an investigator. orton & Co.
Scott, D. H. (1899). Оп the structure and affinities of fossil plants from the Palaeozoic rocks.
Ш. On и ee а new representative of the Cycadofilices. Roy. Soc. Lond. Phil.
Trans. В, 191:8
EXPLANATION OF PLATE
PLATE 1
Medullosa anglica var. ioense
Fig. 1. The stelar system in transverse section. Slide 1391, x 2. in
Fig. 2. Stele а shown at a higher magnification. Slide 1391,
Fig. 3. А portion of the endocentric stele c showing the е РЕЙ and part of
the Eon tissues. Slide 2.
Fi Radial view of ni a l secondary tracheid showing the slit-like, nearly
ee orifice of the pits. “Slide 1392, x 12
Fig Radial view of part of three ian tracheids showing the arrangement
of the ite: Slide 1392, x 66.
Рт.АтЕ 1
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ORAS. VIII
L
ANDREWS & KERNEN—AMERICAN CARBONIFEROUS F
MAIZE IN MEXICO
A PRELIMINARY SURVEY!
EDGAR ANDERSON
Mexico, even more than any other country of the New World, is the land of
maize. There it is the actual staff of life, directly, as well as indirectly, for the
vast majority of the inhabitants. Today, as in pre-historic times, the state of the
maize crop is the commonest subject for conversation throughout the Republic.
Maize is so thoroughly identified with Mexico that a survey of the varieties grown
there might be useful to historians, geographers, and anthropologists, as well as
to agronomists and geneticists. To students of maize in the United States the
Mexican varieties have a special significance. Nearly all of the maize now being
grown here must trace back, though often by very complex routes, to varieties
once grown in Mexico. Some of the problems of commercial maize breeding in
the United States and some of our archaeological problems cannot be solved until
we have a more complete understanding of the maize of Mexico.
Any attempt to get a general over-all picture of Mexican maize is a difficult
problem. Maize in Mexico is extremely variable; it not only varies in the same
way that it does in the United States and with greater magnitude; in Mexico
there are further patterns of variation. In the corncrib or the field it varies from
plant to plant as do our open-pollinated (1.е., non-hybrid) varieties, but the
variation is nearly always greater than in an American cornfield or corncrib. In
one Mexican village it often varies widely from field to field for the same variety.
Unlike American maize, there are, in addition, great differences between varieties
and from region to region. There are frequently as many major kinds of maize
in one Mexican village as in the entire United States, yet when one goes to a village
in another part of Mexico he may find quite another set of varieties. For instance,
there are shown in figure 1 the varieties grown by two Mexican families; one
(above), in Toluca near Mexico City; the other (below), from west of Autlan in
Jalisco. It is unfortunate that photographs of the plants could not have been
included in the picture since variation in plant type was even more extreme than
that in ear type. A careful comparison of these two pictures will show that
several quite distinct varieties are being grown by each family. Yet none of
those grown in Toluca are found in Autlan or vice versa. As a matter of fact,
progressive farmers and agronomists in either of these regions know little or
nothing about the types of maize grown in the other. The ear at the left in fig. 1
(No. 1 below) belongs to an ancient, well-established type of maize widely grown
ШЕТ нн dedicated to the maize growers of Mexico who received me with uniform courtesy
and furthe de these dowd in their Арт: т and store- 34 Much of the work reported herein
was carried o M the author was a Fellow of the Guggenheim Foundation. Acknowledgments
are Pw due arl Sauer t Ralph Beals Isabel Kelly, R. H. Barlow, and E. J. Wellhausen for large
and ётё айн msc of Mexican maize turned over to me for st ne
(147)
[Vor. 33
148 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
in western Mexico (Anderson, 44a). When studying the maize of the Toluca-
Mexico City region, ears of this variety were carried about and shown to farmers,
plant breeders, and other agriculturists. All were much interested, but none
recognized it or knew anything about the general type of maize to which it
belonged. Similarly, the small rice popcorns of fig. 1 (above) were carried back
from Toluca to Jalisco where they created general amazement when shown to the
maize growers of that region, yet the air-line distance between Autlan and
Toluca is only 300 miles.
With regional variation piled upon local variation in this fashion, cataloguing
the maize of Mexico and discussing the results in general terms is a research
problem of no mean dimensions. It is indeed as if one were called upon to discuss
the physical anthropology of Europe before any of the technical papers upon that
subject had been published (Anderson and Cutler, 742). The present paper is
little more than a first approximation. It is based upon six months of residence
and travel in Guadalajara and Mexico City and the regions between them, plus
five seasons of study in the laboratory and the experimental field of collections
made by various collaborators. It scarcely even touches upon such important
centers as Oaxaca, Chiapas, and the barrancas of western Mexico.
In studying Mexican maize it becomes immediately apparent that the cus-
tomary pigeon-holes of maize agronomists (popcorn, sweet corn, dent corn, pod
corn, flour corn, flint corn) are worse than useless. When these names were set
up by E. Lewis Sturtevant in 1884 he dignified them with Latin equivalents, and
they have therefore been taken more seriously as outlining real natural entities
than he himself ever intended. They will serve as pigeon-holes in most of the
United States, though they are artificial and do not divide Zea Mays into natural
subdivisions. In Mexico they are bad even as pigeon-holes, and furthermore the
different ones are bad in different ways. Flint and flour are single gene differences.
In regions where both kinds are being grown (Ameca, Jalisco, for example) this
single difference may occur in the same field and on the same ear so that shelled
kernels from a single ear would have to be classified in two different groups by
this system. Pod corn is likewise a single gene difference and, though seldom met
with in Mexico, it might be expected to turn up in any kind of maize which was
being grown. Sweet corn too is a single gene difference. In the case of
Mexican sweet corn this character becomes still less reliable, since Mexican dents
carry suppressors which may prevent the sweet character from showing even
when it has been inherited.
Dent corns and popcorns however, differ from non-dents and non-pops by a
considerable number of genes, and the names delimit more or less natural entities.
Yet neither one will serve efficiently in cataloguing Mexican maize. The dent
corns vary among themselves in the amount of denting, and they have been so
hybridized with non-dented sorts that the classification into dents and non-dents
is difficult and subjective in many parts of Mexico. Popcorns are more a group
by themselves, or rather two groups. The two most different varieties of corn in
1946]
149
ANDERSON—MAIZE IN MEXICO
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150
ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
22
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Fig. 2. Grid used жыны аы this paper and the Appendix for diagram-
ming variation in Mexican maize. Vertical С row number; осе нар
width of the kernel in mm. T o diagrams illustrate how in
e Six
denting and in pointing of the kernel is represented.
e tex endix.
t and in the foreword to the Appe
Further са іп
[Vor. 33
1946]
ANDERSON—MAIZE IN MEXICO 151
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Бір. 3. aae of all the collections made in Mexico on the same grid
illustrated in fig. 2. Each circle represents the average of a single po
fig
t the right the average degree of pointing has been ке шіге d for each row
number (row numbers run from 22 at the top to the төй np 2.00,
strongly pointed; 1.00, slightly pointed; 0, no аб
all of Mexico are both popcorns. They grow in different regions and on very
different-looking plants. When examined cytologically one has a maximum num-
ber of chromosome knobs, the other few or none. All they have in common is the
ability to pop when heat is applied. Furthermore, one of them (the popcorn of
Toluca) is so thoroughly amalgamated with the dent corns of that region that
one cannot tell where dent corns leave off and popcorns begin (see below).
For the above reasons the maize of Mexico could not effectively be catalogued
in these formal categories. It therefore became necessary to find a more useful
classification. А fine preliminary report was published by the Russian Expedition
(Kuleshov, 730) and with that as a basis it was possible from the beginning to
work towards a natural system of classification for the maize of Mexico. (For
a discussion of natural versus artificial classification see Anderson and Cutler, 742).
[Vor. 33
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN
METHODS
In beginning the work, collections of maize on the ear were received from H.
C. Cutler, Ralph Beals, and Isabel Kelly. It was soon found that in a country
with an ancient, indigenous civilization like Mexico, archaeologists, geographers,
or even historians, were better collaborators than agronomists and botanists. Ма/2
reventador (Anderson, 744) is an example іп point. Dr. Isabel Kelly collected
numerous examples of this interesting variety from various localities in Jalisco and
Zacatecas, though it was not represented in collections made by agronomists,
either Mexican or American.
A variable and cross-pollinated crop such as maize cannot be effectively studied
as individuals; more significant than any one plant or any one ear is the variation
in the entire field or in the entire corncrib. To study the problem effectively it
was therefore necessary to work out techniques for recording and analyzing varia-
tions in populations. Seed from samples from various parts of Mexico was
grown and studied throughout the growing season. Ву trial and error, methods
were worked out for recording the variation between plants, most particularly
the differences between varieties from different parts of Mexico. Herbarium
specimens of tassels and leaves were made and assembled with photographs of the
original ears, diagrams of the internode patterns, samples of the kernels, and in-
formation about the chromosome knobs (in the few cases where that had been
determined).
From these preliminary studies characters were chosen according to the fol-
lowing criteria:
1. The characters used should be objective and, if possible, capable of being
measured or scored in an exact, objective fashion.
2. Their genetic basis should be broad (i.e., they must depend upon a large
number of genes), so that their use will tend to bring genetically similar
individuals together.
3. They should tend to characterize the varieties of any one area and to dis-
tinguish between the varieties of different areas.
The following characters were finally chosen:
Mid-ear width—The width of the mature ear with its kernels attached, meas-
ured to the nearest millimeter with sliding metal calipers.
Shank. diameter-—The diameter of the shank which supports the ear, measured
immediately below the ear with sliding metal calipers. Unfortunately, in most
Mexican maize this has to be estimated from such portions of the shank as adhere
to the ear when it is picked.
Kernel width.—The width of a row of kernels, measured on the ear.
Kernel thickness-—Measured at right angles to the above. In practice the
thickness of ten consecutive kernels was measured with the calipers and the result
was divided by ten.
1946]
ANDERSON—MAIZE IN MEXICO 153
Tesselation—The degree to which the kernels аге tesselated (i. e., like tiles in
a pavement). This is an important difference but needs to be broken down into
such components as variation in kernel size and shape, 6- vs. 5- or 4-sided kernels,
etc. In this material it was scored in three grades.
Denting of tbe kernel.—Scored as absent, slightly developed, strongly devel-
oped. А more objective way of scoring since worked out in North American
material is: 1, kernel with no denting or capping of soft starch; 2, kernel
markedly capped with soft starch but not dented; 3, kernel capped and lightly
dented; 4, kernel capped and dented but surface of kernel not conspicuously
wrinkled; 5, kernel capped, dented, and wrinkled.
Husk striation.—Longitudinal striations across the face of the kernels due to
tight husks. Scored in three grades: 0, absent; 1, striations present but not
making grooves across several consecutive kernels; 2, deep striations across the
face of consecutive kernels.
Base compression.—Ears tapering slightly toward the base due primarily to
tight husks. Scored in three grades.
Row number.—The number of rows, counted about a quarter of the way up
the ear from the base, since many varieties have irregular rowing and/or in-
creased row numbers immediately above the base, and many varieties drop out
rows progressively between the middle of the ear and the apex.
Pointing of tbe kernel.—Scored in three grades.
Enlarged base.—Base of the ear, perceptibly wider than the rest. Scored in
three grades.
Using these characters, variation in populations was studied from one field to
another, largely in Jalisco, Michoacán, and the state of Mexico. In making the
actual collections, 25 ears were selected at random from the field or corncrib
under study. Except in a few instances, the corn in the crib represented the
harvest from a single field. Obvious nubbins and ears on tillers were rejected;
otherwise the selection was completely at random. Іп the corn cribs the microm-
eter was tossed into the crib and the ear nearest its point was selected for measur-
ing. In fields the fourth plant in the row was taken, then the eighth in the
next row, the twelfth in the next, and so on. Whenever possible Leica photo-
graphs were made of a portion of the collection. Due to the great scarcity of
corn in Mexico, the minimum number of ears was retained as samples; it was
usually easy to obtain permission to study the corn if the point was stressed that
we did not care to buy it, but merely wanted to study it for an hour or so.
As soon as a few cornfields and corncribs had been sampled it was apparent
that an efficient method of analyzing and summarizing the data was of first
importance. The variation was too extreme and too manifold to carry in the
mind or even to interpret from a series of averages and ranges of variation. What
was needed was some method of turning the data on each cornfield into а general
[Vor. 33
154 ANNALS OF THE MISSOURI] BOTANICAL GARDEN
over-all picture of that particular field. The graphical method used in Carter
and Anderson’s study of the maize of the Southwest (745) had certain advantages,
but the indices were somewhat subjective, the computations involved and there
was no transparent relation between the spots on the comparison grid and the
actual data. After considerable experimentation the method illustrated in fig. 2
was eventually adopted. It uses the two characters (row number and kernel
width) which best fulfil all the criteria mentioned above. They are diagrammed
at right angles to each other on Cartesian coordinates, the former being plotted
on the vertical axis, the latter on the horizontal. The next two most important
characters in studying the maize of Mexico are the denting of the kernel and the
degree to which it is pointed. Both of these characters are easy to observe but
dificult to score exactly in objective grades. Each was merely recorded as
, "weakly developed", or "strongly developed”. They are, however, so
important in characterizing the maize of different regions and of different varieties
in Mexico that they were included in the diagram. This was done graphically as
shown in fig. 2. The shape of each individual spot on the diagram shows whether
that individual had unpointed, slightly pointed, or sharply pointed kernels. The
degree of darkening of the spots shows the amount of denting on the kernels.
The method outlined above makes it possible to "see the woods in spite of the
trees" and to compare one field of maize in its entirety with another field measured
weeks, months, or even years later. In a variable and cross-pollinated organism
such as maize, populations are more significant than individuals. Each individual
combination of characters is more or less an accident; what is important is the
general complexion of the whole population. If the diagrams in the appendix
are examined it will be seen that each population is a more or less coherent whole
and that the spots are not spattered at random over the diagram; for each sample
they tend to cluster about a particular average. Іп some cases they are strongly
aggregated, in others more dispersed, but even where they are widely scattered
they are coherent. In other words, the field from which they came had a sig-
nificant combination of gene frequencies and the diagram of 25 spots, all seen at
once with a glance of the eye, is a fairly efficient reflection of that population. It
is at least far more significant than isolated averages or frequency distributions of
the four characters taken separately. As an example of how much can be analyzed
and demonstrated by these diagrams, compare the two fields of Maz criollo studied
at Tlaquepaque with the two studied a few miles to the southwest at El Grullo
and Autlán. At Tlaquepaque Maíz chino and similar varieties (with higher row
numbers and more denting and pointing) are growing, and have been grown for
at least a generation, in adjacent fields to the Maz criollo. At Autlán and El
Grullo these other varieties are not grown. It will be seen that, although all four
diagrams present the same general picture, the spots in the Tlaquepaque diagrams
show a tendency to drift up towards higher row numbers and narrower kernels
and that in one of them is strong indication of mixture with a highly dented,
more pointed-kernelled variety of higher row numbers (Appendix Nos. 1 & 2
vs. 15 & 20).
1946]
ANDERSON—MAIZE IN MEXICO 155
The results of the Mexican collections are summarized in Table I and the Ap-
pendix. In Table I the collections are assembled by communities and according to
states, the latter arranged roughly from west to east and from north to south.
Averages for ten of the characters are presented for each collection, together with
the local name of the variety (as well as the name of the grower, or the location
of the field in those cases where several collections in one locality make it necessary
for precise identification).
Table I deals only with averages of populations. Records of individuals are
presented in Appendix I as photographs of representative ears and diagrams of the
type described above. It is apparent from even a casual inspection of Table I and
the Appendix that the ten characters presented in the table are not distributed at
random. On the whole, the characters tend to be associated in complexes, and
these complexes show strong geographical trends. High row number, narrow
kernels, and pointing of the kernel tend to go together as one complex which is
most frequent in the state of Mexico and becomes increasingly less frequent as
one goes westward. These facts are shown more precisely in fig. 3. In that figure
each spot represents the average of a collection rather than individual ears.
It will be seen that, as a whole, the collections run from narrow-kernelled and
many-rowed varieties to wide-kernelled and few-rowed varieties. Furthermore,
pointing of the kernel shows a very definite tendency to be associated with narrow
kernels and many rows. This is demonstrated statistically at the right of fig. 3.
Assigning grades of 2.0 for extreme pointing of the kernel, 1.0 for intermediate
pointing, and 0.0 for no pointing, the averages have been calculated for the
average (median) ears of each row number. It will be seen that the degree of
pointing decreases regularly from 2.0 for 22-rowed ears to 0.0 for ears of 8 and
10 rows.
Table I also shows that this complex of high row number, narrow kernels, and
pointed kernels follows a fairly definite trend from western Mexico to central
Mexico. In the state of Jalisco, the bulk of the ears measured are 8- to 10-rowed,
and population averages never go above 14 rows. While pointed kernels are
fcund in certain varieties, they always represent less than half of the sample. In
Michoacan, the next state to the east, there were no collections which averaged
less than 12 rows and one which averaged as many as 18, and for some of the
collections more than 50 per cent had pointed kernels. Still further east in the
state of Mexico, there were no collections with average row numbers below 14,
and half of the collections had a majority of pointed kernels.
The geographical trend in pointing and high row numbers is demonstrated
more precisely in fig. 4, which presents collection averages for all those communi-
ties from which five or more collections had been made. The location of these
four communities is shown in fig. 5. It will be seen that in the neighborhood
оғ Autlan, in western Jalisco, all five collections had very similar averages in spite
сі the fact that the Autlan collections were actually made over a much wider
area (San Gabriel to Autlán) than those in the other three communities. At S. Р.
[ Мог. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
156
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TABLE I
19461
ANDERSON—MAIZE IN MEXICO 157
| = 5 E
5 > | Вы! Я „| Mg
z = 29| ба | м; | 55 = e xu 2 | 225| Е |+
= 5 Б= | 25 | 55 | 55 | Eg] 8 | 25| ЗЕ | 52 | 8 |4;
% 3 |25| => | we | м | ve | A | 25) 28| 2 | 2 |58
MICHOACAN
ЕУ. Г. Coalcoman area | 16 3.9 1.0 29 32 = + 0 10 0 0
Е. МЕ. Y 15 4.1 ESI Lu 227 = + = 10 0 0
У. № х 14 9:9 LI 1.0 33 за re = = 14 0 0
Е. А. Apatzingan 25 4.6 1.4 ы 529 гір + = 10 0 0
ЕАС г 5 4.2 1-9 1.3 9 = + = 10 0 0
Кел, Sahuayo 3 4.3 1.6 1.4 44 Е E a 10 0 0
E. А. Jiquilpan vi 4.6 1:5 1.0 41 + 0 0 14 0 0
EvA: г 25 4.5 152 .8 .34 Ls == 0 18 um 0
ЕА; E 25 4.7 1.4 ы 41 Ste = 0 12 0 0
E. А. ја 25 251 9 .8 .34 0 E -- 14 0 0
ЕГА ” 25 4.8 3 1:2 .40 + + = 12 zs 0
EA. id 25 4.5 1.6 1.3 45 0 -- T 12 0 0
Е. А. Uruapan 25 4.2 122 LI .40 pm = 0 12 0 0
Е. А. Я 25 4.4 1.7 1.4 .48 ЕЕ + ZEE 10 0 0
Boas San Lorenzo 25 4.4 1.4 1d .45 + 0 0 12 ZI 0
E. A. Cheran 25 3.7 1:5 .8 44 Бі 0 0 14 = 0
Е. А. 3 25 4.4 1.3 1.0 „30 тн 0 = 14 0 0
E. А. ч 22 4.8 1.5 Я 43 Et: 0 0 16 = СЕ
Е. А. Patzcuaro 25 2:7 11 19 38 БІ 0 0 12 0 0
Е. А. й 5 4.4 1,3 42 42 =m m 0 14 0 a
E. А. Lake Patzcuaro | 25 4.2 1.4 3 41 Е 0 0 14 0 0
Е. А. + в 5 4.5 1.4 121 .45 ЕЕ ал 2 12 0 0
E.J. W. Zitácuaro 25 3.6 9 IN ат 0 0 12 0 ШЕ
STATE OF
MEXICO
1 el a A Toluca-
km. 60 25 5.0 .8 152 1 0 zr = 12 0 0
E.J.W. ОО 85570 | 25 3.4 .8 4-28 0 0 0 22 + 0
ЕЛЕУ ” Кт.17 |25 | 4.6 1.0 8 |1139 mm 0 0 16 0 0
ECT WA та 25 3.9 .9 .8 57 0 0 0 14 0 0
E.J.W. КААЛ 4:25 3.9 .8 7 597 ЕЕ 0 0 16 A 0
E.J.W.| Chal 25 | 49 | зр е ЕТЕ о | о | 14 | || о
E: Т.М: Metepec 5 4.7 17] .8 41 + 0 0 16 zm 0
E. A Tepotzotlán 24 p S 4 8 |.40 | + | о 0 16 0 0
E. A. Huehuetoca 25 4.4 .9 .8 41 = 0 гіз 16 0 0
Е.А. Асирша 25 3.9 m 2 132 0 =e Ж 16 0 0
Е. А. Huehuetoca 22 4.2 .8 .8 .42 £ zi 6 14 0 0
GUANAJUATO
Е. Л 25 4.6 1 8 39 xum 0 0 16 0
Е » 25 4.2 1.3 9 215 zum 0 0 16 0 0
MORÉLOS
ЕА; Tlaltitzapan 25 5.3 1.1 .8 102 ЕЕ T 0 16 = 0
Е. А. 4 25 | 4.4 юры] 12 | 0 0
| PUE | | |
R.H.B. Otlatlán 25 4.5 .8 8 .43 cUm 0 16 0 0
TABLE I (Continued)
[Vor. 33
158 ANNALS OF THE MISSOURI BOTANICAL GARDEN
б
4
~
ко
MVO
79 192%
1 1 4 А 197 02
Fig. 4. eee of 5 varieties of maize grown at each of the following towns: 1, Autlán in
western Jalisco; 2, S. P. Tlaquepaque in centra | Jalisco; 3, Patzcuaro, Michoacán; 4, "To luca, "Mexico
Each circle represen nts a collection of 25 ears averaged for row number, kernel width, denting of
the kernel, and pointing of the kernel.
Tlaquepaque, a suburb of Guadalajara, some of the five collections are from ad-
jacent fields and none is more than a mile distant from any of the others. These
Tlaquepaque averages are particularly significant because although two of them
are quite as extreme as those from Autlan, the other three are not, and one of the
collections averages exactly the same as one from Patzcuaro, Michoacan. The
Toluca collections present the other extreme. It will be noted that three of the
five have a majority of pointed kernels and that all of them average 14 or more
rows. Here again it should be pointed out that while these collections were made
within a few miles of each other and some of them from adjacent fields, one of
the collections is no more extreme on the average than are those from Patzcuaro.
Figure 4 therefore demonstrates a fact which is borne out repeatedly in the de-
tailed observations reported in the Appendix: The pointed-kernel, high row-
number complex is not a direct effect of the environment. It is possible to grow
19461
ANDERSON—MAIZE IN MEXICO 159
unpointed, few-kernelled varieties in central Mexico or many-rowed and pointed-
kernelled ones in Jalisco. On the average, however, this is not done, and there is
a strong and surprisingly regular gradient in these characters between Mexico
City and western Jalisco.
64;0 5632 40;
56;0 48; 2846
52,0 4424 20; 0
14;0 6:84
о 8; в
==
TA PAL РА
СНЕ RAN
PATZCUARO
ZI ТАСЈАВО
TOLUCA
g. 5. Approximate cross-section from west to east icis ugh the area in central Mexico where
most of these pue were made. Figures at the extreme right, elevation in feet ab sea-
iie For three mountain towns t of ea riety with ears enlarged at the base is
above at left of colon, the per cent of pointed kernels at right of n. It will be seen that
col
Шы. of ride Sn increase from east to west and percentages of pointed kernels from
st to east, on the av
While the discussion has so far been limited to the evidence on kernel size and
shape and on row number, many other characters are associated with this same
gradient. The photographs of the Appendix show that the western Mexican
extreme types have narrow, irregularly tapering ears which taper slightly towards
the base as well as towards the tip. Тһе varieties from central Mexico tend to
have much shorter ears, many of which taper sharply to the apex and in an exact,
almost mathematical fashion. From observations and collections in Mexico, as
well as from progeny tests made at various points in the United States ( Arcadia,
Calif.; College Station, Tex.; St. Louis, Mo.; Johnston, Iowa; Blandy Farm, Va.),
we know that there are correlated tendencies in the rest of the plant.
JALISCO CENTRAL MEXICO
Long, narrow leaves with tough veins Broad leaves with weak veins
More or less glabrous leaf sheaths Very hairy leaf sheaths
Long, wiry tassel branches Short о branches with densely arranged
Tassel branches, several to many spikel
Tassel cé SN several to few «
Plant color, when co ii is present, Plant color, when conspicuously present,
strongest along the veins and towards the EE between the veins and above the
base of the plant (strong R, a shoot (strong B, weak r)
Strong root system Shallow, weak root system
In other words, we have two extreme types of maize, one characteristic of
western Mexico, the other of central Mexico. While the two extremes are so dif-
ferent as to be instantly recognized even as single individuals, they are connected
by series of various intermediates. On the whole, the average intermediacy of a
[Vor. 33
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN
population is proportional to its geographical position between the two different
centers. Such geographically centered groups of correlated tendencies are char-
acteristic of Zea Mays, and we have chosen to refer to them as "races" (Anderson
and Cutler, '42), defining that term as “а group of related individuals with enough
characteristics in common to permit their recognition as a group." The two
races described above, while not the only major races in Mexico (see below), are
by far the commonest. We have already designated the one from central Mexico
s "Mexican Pyramidal” in reference to the short, and regularly tapering ears. It
has also been recognized by Kuleshov (730) in the report of the Russian expedition
to Mexico as "Central Mexican Type." Kuleshov also referred to the other race
described above as the "Narrow, long-leaved type." Ав a major race it probably
extends all the way from the southwestern United States to the lowlands of South
America. Until it has been more widely studied and more accurately measured
it seems best to designate by name only that portion of it which has been accurate-
ly studied in western Mexico as "Mexican Narrow Ear."
The demonstration of a geographically oriented difference of this magnitude
presents two questions: first, the theoretical one of how it originated and how it
is maintained; second, the practical one of how to use it in making a natural
classification (Anderson and Cutler, '42) of Mexican maize. А detailed discussion
of the latter question will be deferred until some of the special varieties of Mexican
maize have been described. For the first question there is fairly strong circum-
stantial evidence that the ultimate reason is historical; that these two races of
corn were associated with different peoples and different cultures and that they
once were even more distinct than they are now. Ultimately, of course, they
must have had a common origin, but that problem is beyond the scope of this
paper. However, even though there do seem to be historical reasons for the begin-
nings of these two races, their persistence must be due in part to differences іп such
factors as rainfall and altitude. The whole problem is much too complicated to be
anything more than outlined here. Some of the most significant pieces of evidence
are as follows:
The fact reported above, that varieties of very different positions on the
gradient were sometimes grown in adjacent fields.
2. Ina good part of Jalisco (see Appendix) the Mexican Narrow Ear varieties
are referred to by a variety of names but are very commonly called “maíz criollo.”
Criollo, literally "Creole," used in this sense, means native or local, “old stuff be-
longing to this part of the country." Іп this same area intermediate varieties,
however, are most commonly referred to as maíz chino. The word chino, literally
"Chinese", is frequently used to indicate something foreign or peculiar. The
common use of these two terms indicates that the 8-rowed varieties are the older
sort and that the many-rowed, pointed-kernelled types are a later introduction.
Though the latter are still much in the minority, their introduction is not a matter
of just a few years. All the farmers with whom I talked assured me that "maiz
chino" varieties were being raised in the locality when they were small boys (1890
to 1910).
1946)
ANDERSON—MAIZE ІМ MEXICO 161
3. There are various theoretical reasons for believing that when native var-
ieties of popcorn are present they may represent the most primitive types of
maize in that area. The native popcorn of western Mexico, maiz reventador, is
like Mexican Narrow Ear varieties in everything but kernel size and its slightly
higher row number. It has 12 to 14 rows but never the high row numbers of
central Mexico. The native popcorn of the Mexico City-Toluca region, however,
is vastly different and a typical “Mexican Pyramidal” variety. It is exactly like
the small white dents of that region in every character except kernel size and
texture. There is archaeological evidence to indicate that these two popcorns
have been in their two respective areas for a considerable time. As has already
been reported, Dr. Isabel Kelly has found charred remains of a corn at least very
similar to maiz reventador in two excavations in western Mexico (Anderson,
44a). In Gamio’s excavations at Teotihuacan (roughly from 1200 A. D.), just
north of Mexico City, he found large masses of carbonized corn cobs and kernels
which are on display in the local museum and which I was allowed to examine
through the courtesy of the custodian. Measurements for the material are as
follows:
Row number: 18, 18, 20
Kernel width in mm.: 4-7
Kernel thickness in mm.: 4
Kernel length in mm.: 8
Allowing for a little shrinkage in carbonizing they are exactly the same size and
have the same shapes and row numbers as the popcorns and small pop-dents which
are now being grown in the same region.
4. Among the most conservative and primitive Indian tribes of western
Mexico are the Huichol who live in the steep-sided cañon region of Nayarit and
adjacent states. The ethnologist Carl Lumholtz visited them in 1890 and collected
a fine series of the varieties of maize used in their ceremonies, which are now in
the American Museum of Natural History in New York. Three of the varieties
are identical with those still being grown in the neighborhood of Autlán, the
other differs only іп its color (stippled aleurone). All are typical "Mexican
Narrow Ear.” Through the courtesy of the department of Archaeology I was
able to measure these and the results are presented in fig. 6.
5. The differences between these races are inherent. When grown at various
experimental fields in this country, these two extreme types were quite as con-
spicuously different as they are in Mexico. Both forms were somewhat abnormal
when grown in the north, though mostly in different ways. The Mexican
Pyramidal varieties often developed long ear branches. The Mexican Narrow Ear
extremes have the capacity to develop tillers equalling the main stalk. In western
Mexico these varieties are usually planted in the summer on account of the spring
dry season, and only а few tillers develop. (In S. P. Tlaquepaque, Jalisco, I made
a small experimental planting in the early spring and ascertained that Mexican
[Vor. 33
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN
|
5
оо
-
оо ө
А о
оо о
|
o0 O
о
>. 6. сое pd 5224 m Me Huichol Indians by Carl Lumholtz and now in
the American Museum of ral His New York. — circle represents a single ear
(collection пов. 1889, 1892: 5. 1890). pos "of grid as in fig.
Narrow Ear varieties would also develop many tillers if planted there at that season,
though Mexican Pyramidal varieties did not tiller.) When planted in the north
in the springtime, these western Jaliscan varieties tiller profusely. As a result of
these two tendencies, these two races of Mexican corn look even more extreme
when grown in the United States than they do in Mexico.
6. All the above evidence shows that the two extreme types have been in
their areas for a long time and suggests that anciently the boundary between the
two may have been sharper than it is to-day. There is some evidence, however, to
show that ecological factors also play a role in keeping up the partial barrier
between the two sorts. Between Autlan and Mexico City there is a general
change in altitude and in climate pattern. The differences in amount and distri-
bution of rainfall are probably a controlling factor. All of the large collections
of corn which I measured in the plain and valleys around Autlán (Autlan,
1946]
ANDERSON—MAIZE IN MEXICO 163
Chachahuatlan, Tuxcacuesco, San Gabriel, El Grullo) were either specialty crops
like maiz dulce (see below) or were typical Mexican Narrow Ear varieties. This
is a region in which the scarcity of water has been a problem for a long time
(Kelly, 745). Near by, in small clearings in the forested hills between Tuxcacu-
esco and El Limon, however, I made small collections of much superior, large-
cobbed, many-rowed, deeply dented varieties (see Palmar in Table I and the
Appendix). I was told locally that these varieties were grown back in the hills
because there was more water available there.
MOUNTAIN YELLOW
While the maize of the Mexican plateaus is varied as compared with that
of the American Corn Belt, the variation becomes even greater as one ascends the
mountains which rise above the plateaus. This maize of the mountains is so ex-
tremely variable from plant to plant and from field to field that it is difficult to
discuss its general over-all tendencies. Collections were made in three such
mountain regions and at about the same elevation in all three cases (fig. 5):
(1) Toluca, just to the west of Mexico City; (2) Cheran in Michoacan; and
(3) Tapalpa in Jalisco, on the northern flank of the great Volcan de Colima. In
all three of these regions the corn was smaller-eared and shorter-statured than on
the plateaus below. It was also more variable in pericarp color and with more red
in the pericarp on the average. The percentages of pointed kernels were also
somewhat higher. In the mountains one also finds a character which is very
common in Guatemala though rare in most of Mexico: ears which are conspicu-
ously larger at the base, giving them the over-all appearance of slightly tapering
cylinders emerging from flattened spheres. Such ears are seldom seen on the
plateaus. Around Guadalajara, Jalisco, for instance, at a general altitude of
5200-5400 feet there were twelve collections made. In eight of these there was
not a single ear with a perceptibly enlarged basal region; there were two collec-
tions with 4 per cent and two with 16 per cent. In the mountains of Michoacan
nine collections were made at elevations over 7000 feet with percentages from
32 to 80.
In spite of the variability of mountain corn and in spite of a general tendency
for increased basal enlargement with altitude there is a very perceptible geograph-
ical trend in mountain maize from east to west. In central Mexico the mountain
corns tend to be white, many-rowed, with pointed kernels. As one goes westward
at equivalent elevations there is an increasingly great tendency for them to be
yellow, few-rowed, and with an ear perceptibly enlarged at the base. This con-
clusion is supported not only by the detailed collections from the three areas
referred to above but by much more maize which was seen but not measured, and
by the large collections made by Beals in Michoacan and by Kelly in Jalisco. For
Tapalpa, Cheran, and Toluca the measured collections yield the following infor-
mation:
[Vor. 33
164 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Prevailing
Locality E State Per cent enlarged Базе | Per cent pointed color
40 4 Bright yellow
28 16 Light yellow
Toluca Mexico 20 0 White
8 84 White
| ___|- 8 _|__ 8 ____| White-yelow —
56 32 Light yellow
EIL А у 48 0 White
Cheran Michoacan 44 24 qoe m
- О __|_____4 |. 32 _| Light yellow —
64 | 0 Yellow
56 0 | Yellow
Tapalpa Jalisco 52 0 White
4 0 Light yellow
| 200 | 0 Light yellow
TABLE П
This tendency to yellow-endospermed varieties with fewer rows and enlarged
bases seems the more significant because this combination of characters is met with
in Guatemala to an exaggerated degree. It seems likely that, in addition to the
Mexican Narrow Ear and Mexican Pyramidal complexes of characters, there is a
third complex much less strongly represented, which we may call "Mountain
Yellow." Much more study will be required to demonstrate its presence and to
analyze its historical and biological basis. From the facts at hand it seems to be
connected with Guatemala in some way and in central Mexico to have spread from
the west eastwards, at high elevations.
VARIETIES FOR SPECIAL PURPOSES
“Elote” Varieties (see pl. 7).—
In Mexico, as in the United States, much corn is used before it is fully ripe,
either as corn-on-the-cob (elotes) ог in various special dishes. Though native
sweet corns are widely distributed in western Mexico (Kelly and Anderson, 743),
they are not used for green corn since they are too gummy. Most of the common
field corns are so used but there is also a widespread tendency to grow special
varieties with colored (blue or red) aleurone for that purpose. This tendency 18
more clearly marked, and the varieties are more clearly differentiated morpho-
logically from other local varieties, among the Tarascans of Michoacan (as has
been reported by Beals, 746 [in press]). The elote varieties of this region are dark
blue or black and are strikingly different in kernel size and ear shape from the
field varieties there. They are not grown in the regular corn fields (mil pas) but
in small fenced enclosures close to the houses. In Jalisco the differentiation of
elote varieties is not so extreme but it exists. In the town of S. P. Tlaquepaque,
for instance, field corns are mostly white or yellow and are planted quite thickly
in heavily ridged rows with an underplanting of squashes. The elote varieties are
red (less frequently blue) and are ordinarily planted widely spaced in fields of
peanuts. They are early-maturing and are harvested before the regular field
varieties. In Tapalpa, Jalisco, on the northern flank of the great volcano of
1946)
ANDERSON—MAIZE IN MEXICO 165
X
ig. 7. Relation between the pe Of purple Es corns and the common varieties of maize
grown in seven A M communities in various parts of Mexico. Grid
kernel width as in fig. 2. In each case the circle with a number shows the average ro
T
Tlaltizapán, ar élos; 4, Uruapan, wer oacán; T Jiquilpan, оф MERE a
, Huehue , Mexico. Number or instance, indicates that "elote" v
er of 8 n m
grown by the same family had an average row number of 14 and an average [фар width of
10-11 mm. Further discussion in the text
Colima, I was at first told that no special varieties were used for elotes. Descrip-
tion of the blue and red corns I had seen elsewhere brought out the fact that they
were also grown in Tapalpa but on a very small scale. The local corn merchant
himself showed me seed ears for his next year’s crop of elofe corn hanging in his
own patio, though no stocks of this variety were stored in his big commercial
granary.
Where they were grown these blue and red varieties were reported to be super-
ior for green corn-on-the-cob, and for the production of various other maize
foods such as pinole (Kelly and Anderson, 743). Inquiry as to why they were
[Vor. 33
166 ANNALS OF THE MISSOURI BOTANICAL GARDEN
grown usually brought out that they were sweeter or of a smoother consistency
("son mas suave"), and sometimes that they were earlier to ripen.
In the Tarascan country (Cheran and other villages near the new volcano)
the elote varieties were so strikingly different morphologically from the other corn
of that region that it seems as if they must have been introduced at another time
than the field varieties or from a different region. In other parts of Mexico the
difference is not so striking but there is the same general tendency for these
varieties to have broader kernels and fewer rows than the ordinary field varieties.
It would be difficult to be certain on this point from mere inspection, due to the
great variability of Mexican maize. However, the method of recording the varia-
tion in an entire corn field, illustrated in fig. 2, allows us to make exact compari-
sons between elote varieties and field corns from the same localities. Figure 7
shows comparisons of the averages of the blue or red elote varieties in seven dif-
ferent communities with the white or yellow field varieties of the same com-
munities. In cases where more than one field variety had been examined the one
chosen was from the nearest field or was one grown by the same farmer. It will
be seen that in each of the seven cases the elote varieties are more like Mexican
Narrow Ear varieties. They are either fewer-rowed or broader-seeded or both.
To summarize: In Mexico special varieties of maize with colored endosperms
are grown for use as green corn. These varieties are generally broader-kernelled
and have fewer rows than the common varieties with which they are grown. There
is a tendency to grow them in special plots. All these facts suggest that after
maize growing was already established, there was the introduction of Mexican
Narrow Ear varieties with colored aleurone for use as green corn. There is no
evidence that this introduction was a recent matter and there are some indications
that it may have been pre-Columbian. The habit is widespread; it is highly
developed in little out-of-the-way towns with conservative habits, and similar
varieties are used Бу the Huichol Indians in their ceremonies. This last fact is
particularly significant since the Huichol are a conservative group living in a
region isolated by high mountains and deep canons. The collections of ceremonial
corns described above were still being used when Lumholtz collected them, and
they probably represent varieties which are very ancient in that part of Mexico.
Two of the varieties collected by him, one a blue and one a red, are so similar to
the elofe varieties now being grown in Jalisco that they could not be separated
out if the ears were mingled. Since the Huichol are in western Mexico and since
the general trend in each locality is for the elote varieties to be more like extreme
examples of Mexican Narrow Ear, it seems highly probable that these elote corns
spread across Mexico long ago from the west and that they have since become
more or less contaminated in each locality by the common maize of each region.
Cacahuazintle.—
One of the most distinctive varieties grown in Mexico is the kind called
Cacabuazintle (literally, "ear of corn with a tough skin") in the region around
1946]
ANDERSON—MAIZE IN MEXICO 167
Mexico City. We have two collections, one from the valley of Toluca and another
from the little Nahua-speaking village of Otlatlán in Puebla. In both ear and
plant characteristics this variety is enough different from the other varieties of this
region to be an obvious introduction. On the other hand, its Nahua name and
its presence in this remote village suggest that its introduction is not a recent
matter (see fig. 1, lower right two ears, and Appendix No. 71).
Cacabuazintle kernels are filled almost completely with soft starch, though
some of the ears from Toluca and a majority of those from Otlatlán show indica-
tions of a more or less well-developed dent. Their kernels are large, nearly as
thick as they are wide, and tend to be well distributed over the apex of the ear.
I did not see the plants but was told in Toluca that they are of a different color
from the common corn of that region and that they are taller and later to mature.
The ears of Cacahuazintle are so similar as to be almost identical with the
Guatemalan variety known there as "Salpor", which in turn differs but slightly
from some of the large-kernelled white flour corns of highland South America.
Cacahuazintle is reported to be used commercially in the neighborhood of Mexico
City for the manufacture of starch. In that region it is commonly prepared by
swelling it in rapidly boiling water, and when so treated is considered as a kind of
popcorn. While I never saw any specimens of it in western Mexico, its reputation
had preceded it there, and I was several times told about the wonderful big pop-
corn of Mexico City which was cooked in boiling water.
The facts reported above are all in harmony with the hypothesis that Caca-
buazintle spread into central Mexico from Guatemala or southern Mexico in
pre-Columbian times.
The maize varieties of several Spanish-American communities in the American
Southwest bear a suspicious resemblance to Cacahuazintle in their large white
kernels. This is particularly true of some obviously mixed commercial varieties
from Lower California and in the "maíz del pais", which has been previously
described (Anderson, '44b), from San Luis Obispo, California. It seems quite
possible that a variety with kernels of this size and quality might have been widely
scattered in colonial as well as pre-colonial times. А careful comparative study
of Cacabuazintle and similar varieties should give us a more precise understanding
of these relationships.
Maíz dulce (see pl. 8).—
These distinctive varieties, carrying the recessive gene for sweet, have been
the subject of a special monograph (Kelly and Anderson, '43). It remains to
point out here that although collections made in Jalisco show certain general re-
semblances to the other maize of that region, they also show a number of differ-
ences. They have more rows (the rowing is more irregular), the ears are shorter
and are much more smoothly rounded at the butt. The kernels are smaller and
the red color of the pericarp (due to various crown alleles of P) is not common
in that region. А detailed discussion of these differences must be postponed until
[Vor. 33
168 ANNALS OF THE MISSOURI BOTANICAL GARDEN
such a time as the origin and dispersal of sweet corn in the New World is con-
sidered in detail. То clarify this survey of Mexican maize the following summary
of the evidence may be inserted parenthetically. Recessive genes for sweet are
known to have appeared as mutations at various times and places, and may well
have occurred repeatedly in various parts of North and South America. The
origin of the use of these distinctive mutations, however, follows a fairly simple
pattern. [t apparently took place in pre-Columbian South America as a source
of sugar, before the introduction of sugar cane, sorghum, etc. Like the varieties
of the region where they originated these primitive sweet varieties had "hand-
grenade" ears, crowned pericarp, and many and irregular rows. They spread
northward, gradually mixing with other kinds of maize, but protected somewhat
by being recessives so that when careful seed selection was practiced, outcrossed
kernels were discarded. In combination with the characteristics of the maize in
which they originated, these sweet mutants were too gummy to be eaten as green
corn. As the sweet varieties were gradually modified in their spread northward
they eventually became watery enough to use as green corn, and were developed
in this way among the North American Indians. The post-Columbian introduc-
tion of sugar cane reduced their importance as a sugar source and only in a few
conservative areas (as in small towns in Jalisco or among the Hopi) have they
been retained for their original purpose.
Popcorns.—
In addition to cacabuazintle (which is scarcely а popcorn by American
standards) there are at least two popcorns in Mexico. They are grown in different
areas and are associated with different races of maize. Maíz reventador (pl. 9), of
the west of Mexico, is similar to Mexican Narrow Ear varieties in everything but
kernel size and row number and is probably one of the ancestral sources of those
varieties. It has been the subject of a special monograph (Anderson, '44a). The
popcorns of Toluca are typical Mexican Pyramidal maize and are thoroughly
amalgamated with the small white dents of that region (pl. 5). No natural bound-
ary can be laid down between these small white, many-rowed dents and the small
rice popcorns of the same regions. They are identical in plant type and they fade
into each other from plant to plant and from field to field. A little denting is
to be found in every crib of popcorn which I have examined, and hard, flinty,
pointed kernels are a commonplace іп the white dent corns of that region.
Archaeologically these Toluca popcorns and pop-dents are interesting because they
are identical in size and row number with the corn excavated by Gamio at the
ancient pyramids constructed by the Toltecs in the valley of Mexico. Agronom-
ically they are noteworthy as the probable source of the so-called Jap Hull.less
popcorns to which they bear an extremely close resemblance in plant, kernel, and
ear characteristics.
1946]
ANDERSON—MAIZE IN MEXICO 169
DISCUSSION
There remains the discussion of certain problems in the light of this general
survey of maize in central Mexico: (1) the practical question of how to describe
and catalogue Mexican varieties of maize, (2) the origin of denting, (3) the
genetics of multiple factor characters in modern maize.
(1) The classification of Mexican maize.—lhe question of artificial vs.
natural systems of classification was discussed in extenso in the first paper of this
series (Anderson and Cutler, '42) and need not be repeated here. It was pointed out
there that the system usually followed (Sturtevant's) was almost purely artificial
and that it was not very practical outside of the United States. A wholly natural
system, however, if indeed it ever can be attained, is a goal to work towards rather
than something which can be immediately set up in its complete form. We have
therefore studiously avoided the use of Latin names and descriptions which would
give our tentative and fragmentary attempts a more official status than they
deserve. In the survey reported above we have attempted to demonstrate that
the characters of Mexican maize are not distributed at random but are loosely
associated in a few great complexes with marked geographical and altitudinal
trends. The groups of varieties bearing these complexes of characters we have
termed races, and while some fields show strong evidence of only one race, much
of the maize of Mexico shows two or more in a single field.
For the practical business of describing and cataloguing Mexican varieties it
would seem quite feasible to describe the typical "nucleus of common features"
which typifies each race and then to indicate the preponderance of the various
races in any one variety. When more than one race is evident in a variety, they
should be cited in the order of their predominance. Races evident only upon close
inspection should be cited within brackets. The varieties grown around Guadala-
jara, for instance, would mostly fall into Mexican Narrow Ear or Mexican Narrow
Ear (Mexican Pyramidal). The varieties grown in Tapalpa would be classified
as Mountain Yellow, Mountain Yellow (Mexican Narrow Ear), and Mountain
Yellow (Mexican Pyramidal and Mexican Narrow Ear). When all the main races
of maize in Mexico have been recognized and described it should then be possible
to work out exact statistical techniques for assigning numerical values to the
various racial components of any particular variety.
It is still premature to do anything more than suggest such a system. In ad-
dition to the races described above there are certainly others in Mexico. Тһе
varieties of the Tarahumare Indians need intensive study, so do the varieties of
Oaxaca, Chiapas, and the east coast. Through the kindness of the Dirección
de Agricultura y Fomentá and of Dr. E. J. Wellhausen of the Rockefeller
Foundation I was able to measure and photograph collections from other
parts of Mexico. From these collections it is clear that the highly derived
varieties of the Pueblo Indians have spread down to northern Chihuahua and
that some of the maize of the Mexican state of Chiapas is similar to that of
33
[ Vor.
ANNALS OF THE MISSOURI BOTANICAL GARDEN
170
еі
НА ПР НИТИ а
РА А НИНА ETT DP ТЕЛДЕ
эне ТА ТІ ЕНІП»
i
«x
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РНИИ» |
Пира фое So
: jer
san
x LEE ВР ОНУ Ае уло (2; '
ч
8.
Бір.
among the collections made in Mexico:
Drawings showing racial extremes
sc 2
“4.9 5
= о
wos
EP?
S =
~ ©
520
"e
—
ша A „щ
cU a
Mountain Yellow
-—
=
с
2
c
о
~ Бе:
D 792
w с
с
=
~
e 4
5 о
= ~
E
c =
< [s]
е ж
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= 4 E
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309 =
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representing an even more extreme type than
a,
maíz chapolote from Sinale
Lower right,
“Mexican Narrow Ear.
Scale in ст.
1946)
ANDERSON—MAIZE IN MEXICO 171
adjacent Guatemala. The large shanks, enlarged base to the ear, and long ears
of these varieties from Chiapas are a complex of characters found іп some-
what diluted form in Mountain Yellow but of particular importance in Guatemala.
Just as Mexican Pyramidal maize is tied up with the history of the Aztecs and
their predecessors in the Valley of Mexico, so this Guatemalan complex is asso-
ciated with the Mayas and their forerunners. Ав a convenience to students of
Central American maize, the characteristics of these Mexican races are listed
below in condensed form:
Mexican. Pyramidal.—Found in purest form in the Mexico City-Toluca neighborhood. Plants
short, highly colored, color characteristically interveinal (strong B, weak r).
root systems, leaves broad, with hairy sheaths. Tassels with few branches or none, condensation
index very high. Ears short, tapering regularly, row number high to very high. Ke
less pointed, more or less dented. Endosperm prevailingly white, and yellow, usually pale, even
when present (pl. 4).
Mexican Narrow Ear—Apparently the common гасе of western Mexico, though variously
modified locally by introduction and selection. The prevailing type in Jalisco up to 7,000 feet
Allied to such ancient varieties as maíz reventador. Plants tall, slender, from slightly to highly
colored, color veinal (strong R, weak b). Stalks stiff, sheaths only slightly hairy. Tassel branches
several to many, long and wiry, condensation index low. Ears narrow with small cob ir
ularly long-tapered, compressed at butt and with long striation lines from tight husks. R -
ber characteristically 8 to 14. Kernels unpointed, slightly dented, if at all. Endosperm prevailingly
white (all those tested have been ccrr [pl. 2
ый
8
Ze
5
о
Ud
Mountain Yellow.—At high elevations. Found in purest form on the tableland north of the
Volcán de Colima. From thence eastward, becoming progressively more mixed with Mexican
Pyramidal. Endosperm bright yellow. Ears and kernels small, somewhat compressed. Ears dis-
tinctly enlarged at the base and with irregular rowing there (pl. 5).
pe. Ears long with large, strong cob,
firmly held on a big shank. end of ear distinctly enlarged, upper part sub-cylindrical. In
Mexico known in a pure form only from the State of Chiapas.
(2) Tbe origin of denting.—The genetics of denting in maize has scarcely
advanced beyond the point where it was left by the pioneer investigations of
Hayes and East (715). Whatever the origin of denting, its genetical background
seems to be complex. The survey reported above demonstrates that in Mexico,
denting reaches its most extreme development neither in the Mexican Pyramidal
varieties nor in the Mexican Narrow Ear, but in kinds like maíz chino, which
seems to be a mixture of both races. This assertion is no more than suggested by
the figures of Table I, but in Mexico it is a matter of common observation that in
traveling from western Jalisco to Mexico City one passes out of a region of un-
dented or slightly dented Mexican Narrow Ear varieties into the variable but
strongly dented varieties of the Bajio and eventually reaches another region of less
denting, the Mexican Pyramidal center of pointed popcorns and pointed "sub-
dents." This would suggest that denting is in some way associated with race
mixture in maize. These results are in conformity with Jones’ experimental evi-
dence (724). He crossed a pointed popcorn with a flour corn and obtained
dented kernels in the second generation. It may be that in some manner not yet
clear the dent corns of Mexico are hybrid derivatives of the pointed popcorns and
the crescent-seeded flint and flour corns.
(3) The genetics of multiple factor characters in maize.—The survey re-
ported above demonstrates that certain complexes of characters tend to be found
[Vor. 33
172 ANNALS OF THE MISSOURI BOTANICAL GARDEN
together in Mexican maize. Since all of the dent corns of the United States are
derived from Mexican varieties, at least in part, we may expect to find some of
these complexes still playing an important role in the maize of the United States
Corn Belt. One who has studied character combinations in the cornfields and
corn cribs of Mexico cannot walk through a modern breeding field in the United
States without noticing the resemblances of certain widely grown inbred varieties
to Mexican corn. The two inbreds, K43 and Illinois HY, for example, present
combinations of characters that, while very different from the open-pollinated
varieties from which they are derived, are almost identical with certain Mexican
combinations. It has been shown from theoretical considerations (Anderson, '39),
as well as by experimental test, that the total effects of specific and racial co-
herence are incredibly strong. If this be true in maize, we might therefore expect
that our American dent corns would still be characterized by whole blocks of
genes which tend to stay together because they came in together in Mexican white
dents. If these complexes of genes still tend to be found together, on the average
in American corn, then to understand and to analyze the genetics of multiple-
factor characters in maize we shall have to pay attention to certain combinations
of characters. In other words, the genetics of multiple-factor characters in maize
has а good deal in common with Oenothera genetics. The complexes іп Zea аге
probably much less loosely knit than they are in Oenothera and will therefore be
more difficult to recognize and to locate in the germ-plasm. Оп the other hand,
Zea is more completely analyzed genetically than any other plant. With concerted
action it should eventually be possible to find which chromosomes carry the dis-
tinctive character combinations of Mexican Narrow Ear and Mexican Pyramidal,
and then to use that information in maize breeding.
SUMMARY
1. The great variability of Mexican maize is described. It varies from plant
to plant in the same field, from field to field in the same locality, and from region
to region. For the first two of these categories the variation is like that in the
United States but greater in magnitude; the regional variation is beyond anything
encountered in the United States.
2. The customary pigeon-holes of maize agronomists (dent corns, popcorns,
flint corns, etc.) are shown to be inefficient and misleading in Mexico.
3. It is argued that in classifying a variable, cross-pollinated crop like maize
the population is a more significant unit than the individual.
A method of measuring and describing maize populations (fields) is de-
~
veloped and applied to the collections made іп Mexico.
5. By means of this method the association and geographical distribution of
high row number with narrow, pointed kernels is analyzed.
Other characters are shown to be associated with these trends in row num-
ber and shape of kernels. High row number, pointed kernels, wide, hairy leaves,
condensed tassels, constitute a complex characterizing the Mexican Pyramidal race.
19461
ANDERSON—MAIZE IN MEXICO 173
It is centered in the state of Mexico. Low row number, long, irregularly tapering
ears, wide kernels, narrow leaves, long, wiry tassel branches, form a complex char-
acterizing the Mexican Narrow Ear race of western Mexico
The available evidence suggests that the ultimate reasons for these dif-
ferent races are largely historical but that their persistence in modern times is in
part due to their adaptation to different areas.
8. Evidence is presented for the existence of a third race, Mountain Yellow.
It resembles Guatemalan maize, is restricted to high altitudes, and from Jalisco
eastwards to Toluca becomes increasingly mixed with Mexican Pyramidal.
9. Several varieties grown for special purposes are described in detail:
A: Varieties with colored aleurone used as green corn. They apparently
spread from western Mexico in ancient times and have become more or less
mixed with the common varieties of each region.
B: A large-kernelled flour corn known as "Cacahuazintle" apparently de-
rived from the "Salpor" variety of Guatemala.
C: Maíz dulce, whose significance in the history of American sweet corns is
briefly indicated.
The ancient popcorn of the west coast, maíz reventador, and the very
different rice popcorns of the state of Mexico.
10. Three general problems are discussed in the light of the above survey:
the classification of the maize of Mexico; the origin of denting in maize; and the
genetics of multiple factor characters in North American maize.
LrrERATURE CITED
Anderson, Edgar (1939). Recombination in А ар crosses. ее 11 668-698.
-------, (1944a). Maíz reventador. Ann. Mo. Bot. Gard. 31:3
, (1944b). “Maiz del aut in California. ойт. Calif. ва ee . 52:2—6.
, and Hugh C. Cutler (1942). Races of Zea Mays: Y. Their tesognitión and classifica-
tion.
Beals, Ralph L. (in press). Cherán, a Sie «Mi 5 Village. Smithsonian Mid eee Social
Anthropology Publ. No. 2. (Includes " Repor on Maize of Cherán by Edgar Ander
e | F., and Edgar Anderson asa. A preliminary survey of maize in the ы
n United States. Ann. Мо. Bot. Gard. 32:297-322.
Dos H к. and E. Um an (1915). Further experiments on inheritance in maize. Conn. Agr.
ius = qun 188:
Jon (1924) о gin of flint and dent corn. Jour. Hered. 15:417
Kelly, ыы (1945). Тһе о of the Autlan-Tuxcacuesco area of Jalisco 1: The Autlán
98
ana e .
------, and Edgar Anderson (1943). yes: corn in Jalisco. Ann. Mo. Bot. Gard. 30: ео
ко E N. (1930). The maize of Guatemala, C Panama and Colombia.
Mex uba, p.
йу 1, in saan 5. М. The rte 2, p ts of Mexico, mund and Colombia. Bull,
in , Genet., & Plant Breed. занд 47:470-553 (English summary).
Lum © 2 (1902). о wn Mexico. Scribne
Sturtevant, E. Lewis (1889). Varieties of cork, UL 5. Dept. Agr. Off. Exp. Sta. Bull. 57:1—108.
[VoL. 33, 1946]
174 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATES 2 то 9
Most of the photographs in these plates are due to the energy and foresight of Prof. Carl
Sauer, of the Department of Geography of the University of Cali fornia, who not only assembled
igni о
a significant collection of Mexican maize saw to it tha s expertly photographed. The
negatives are e property of the Museum of Anthropology of t institution, an am
indebted to Mr. E. W. Gifford for permission to use them h The ears illustrated in pl. 5 and
the lowermost ear in pl. 4 are from my own
collections; 2” кеде were collected by Dr. Isabel
Kelly. All are reproduced at a little lon Fs natural s
Plate 2. Three ears of Mexican Narrow Ear varieties. Left to right:
pan.
Tuxpan
from Ameca, Tuxpan,
Four ^ i Intermediate or maíz chino varieties.
eca
Plate 3. From top to bottom: from
Sayula, Pm Sayula, A
Plate 4. Four ears in which the Mexican laugh i influence is predominant. The
nda ears are from Nochistlan, Zacatecas.
top
The botto а о Morelos. It is an
e example of the variety known as pipitillo whi ich i is aw red by corn merchants зде it
shells readily and produces a maximum volume for sale by the
Plate 5. Five ears from high altitudes, showing Mexican ч influence. Тһе
to the left аге from the village of Tlacotepec near Toluca maller ones show strong
evidence of mixture with Mountain Yellow. Right above, жй. black нуе zround, the co
orn of Toluc i
Mexic ity.
Mexican Pyramidal with a strong indication of Mexican Narrow in the ed rd husk
striations, lower row number, ad plant type (See Appendix No. 64).
Plate 6. Four oe of Mountain Yellow from Tapalpa, Jalisco, on the northern flank
of the Volcán de Colim
ads 7. a examples of "elote" varieties with colored aleurone. The the left
om Unión de Tula, Jalisco; the one to the right is from Ameca. Note 42 sale "white seed,
which probably fine pollination with dominant white (Ci)
Plate 8. Two ears of maíz dulce, a primitive source of sugar, from Unión de Tula, Jalisco.
Plate Two of maiz reventador from Tuxcacuesco, Jalisco (badly damaged by rw
and one des бі, "Jalisco. Lower right, two ears of maíz chapolote from an ejido near Culiac
Sinaloa, Mexi
>
~
PLATE
Vor. 33, 1946
ANN. Mo. Вот. GARD.,
об
ога
Ne
pido
uu
"(ТЇ
00100000006
НІШ
M
MEXICAN NARROW EAR
ANN.
Mo. Вот. Garb., Vor. 33, 1946
) = |
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i
|| ||
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| |
А
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ae p ;
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207 «eU Ш 709009000 T у
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PLATE
INTERMEDIATES
3
ANN. Mo. Вот. Сакр., Vor. 33, 1946 PLATE 4
MEXICAN PYRAMIDAL FROM LOWER ALTITUDES
PLATE 5
33, 1946
VoL.
ANN. Mo. Bor. GARD.,
— ЕР
бан = +“ NT а < БИР”
ес 47. 42%
азығы ый
TM eT Ф, „АМА МА MX
T ERU IUDA
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DES
MEXICAN PYRAMIDAL FROM HIGHER ALTITU
ANN. Мо. Вот. Сакр., Vor. 33, 1946
PLATE 6
»
4 j j
ў | : I3
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MOUNTAIN YELLOW
PLATE 7
GARD., Vor. 33, 1946
ANN. Mo. Bor.
VARIETIES
ELOTI
ANN. Mo. Вот. Garb., Vor. 33, 1946 PLATE
MAIZ DULCE
ANN. Мо. Вот. Garp., Vor. 33, 1946 PLATE 9
MAIZ REVENTADOR
19461
APPENDIX 175
Maize COLLECTIONS FROM MEXICO
FOREWORD
The following records and photographs are an attempt to sample and record
the common, every-day corn of Mexico. Much of it is from villages off the
highway and the railroad; of the 72 samples only two represent the finest corn
which could be raised in that particular vicinity. Varieties of spectacular pro-
ductivity can be, and are, grown in Mexico; in this survey they were rather
deliberately avoided. The old-fashioned varieties of the small growers give a
clearer picture of what indigenous Mexican maize was like before its variation
pattern had been blurred by modern commerce.
The grid used in each diagram has been explained in fig. 2 of the paper to
which this is an appendix. Row numbers from 8 to 22 are diagrammed on the
vertical axis at the left; kernel widths in mm. are diagrammed on the horizontal
axis across the base of the diagram. The first division of the scale represents 4-5
mm., the second 6-7, and so on up to 16-17 at the right. The same grid to the
same scale has been used throughout. For a few of the collections with very wide
kernels only the right-hand end of the scale is represented. The vertical scale
(though not drawn) is the same as in all the other diagrams and can be obtained
by measurement. The horizontal scale begins at 10-11 mm. instead of at 4-5
mm. and is so indicated in each diagram.
It might be well to point out that although these grids resemble the scatter
diagrams made in preparing a correlation table, they are a cruder device. They
are nothing but a set of pigeon-holes classifying all the ears of maize simultaneous-
ly according to row number and kernel width, and diagramming their denting
and kernel pointing in a graphic manner. The ears that fall in each square (each
separate pigeon-hole, that is) are diagrammed as close to the center as is practi-
cable; the arrangement within the square has no significance.
The photographs are all snap-shots made with a Leica camera. All but one
have been printed to the same scale. Most of the pictures include the calipers
used in making the measurements, and in each case the calipers were set to exactly
5 cm. In so far as possible the ears which are photographed represent a random
selection from the field or crib. Where local pride prevented the illustration of
a typical sample, this has been indicated in the notes.
Kernel color was scored separately for each ear that was measured, and the
summaries below each picture are an exact digest of accurate notes. Cob color
was not recorded in so systematic a fashion, and the lack of any mention of red
or purple cobs is no indication that they were not present in the sample.
For a variety of reasons the Appendix does not exactly coincide with the
collections reported in Table I but in general the collections are represented in
both places. In the Appendix, as in the table, the collections are arranged accord-
ing to Mexican states and roughly from west to east within each state. They
were mostly made on the plateaus of central Mexico and the lower flanks of the
mountains which rise above these plateaus. Aside from a few scattering collections
from other parts of Mexico they are from a region about 300 miles in extent, from
a little west of Guadalajara to the vicinity of Mexico City.
[Vor. 33
176 ANNALS OF THE MISSOURI BOTANICAL GARDEN
о өө ©
-
E о -
E аға” „ы о ө
E өө ee @
L 1 1 о 1 Q E шай
10-11 Kernel width
1. Аш ап, Jalisco. Maíz criollo
ўе» а small pu at the ee = town. m ailingly white though a majority = seh ears
had a yellow and a few ears had n One ear had a tan pericarp and o
жен oak wie ен three of whieh were ie and one Pr. Horizontal scale shifted. to the
left (as indicated) to accommodate wide kernels. Vertical scale as іп No. 4.
19461
ANDERSON—MAIZE IN MEXICO 177
ee ө
ә?”
E e ©
eo ©
— e e ә o -
o © ©
1 o о
1 | 1 1 @ | 1
10-41
2. El Grullo. Maíz humiado
The name ing cd 'smoked") refers to the tan | eaa which тај кји all but eight of
the ears ably most of these Pie exhibited this character but i o dilut state for recog-
nition vithout dissection of the pericarp. Over
endosper r bore a single kernel with colored aleurone (Pr.). Horizontal scale as
the Di Pur cis not shown, is the same in the first three collections as in those that fallow.
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
178
oo ө
10-11
El Chante, near Autlan, Jalisco.
Maíz Ритіайо
3 .
2.
Very similar to No.
A sample of thirteen ears obtained for me by Don José Maria Corona.
ии ve
S унин ина, "
генін - Т $)
ent i кеннен
Өш.
“бі,
-— iini TUM
19461
ANDERSON— MAIZE IN MEXICO 179
т E
с
=
Е
5 22
т k-
20
18 o
[о]
16 o
© O O
14
O O
12 O O
10
8
1 L 1 L 1 4 4
4-5 6-7 8-9 10-11 12-13 14-15 16-17
Kernel width
4. El Limón, Jalisco. Maíz reventador
sample of ten ae abtained by Don José Maria Corona. Prevailingly white. e
colored kernels were all pr. Ears similar о to those po in Pi. 9 and in the following Sn ene
Scale as in fig. 2. аа cal scale: numbers from 8 22; horizontal scale: kernel width
from 4-5 to 16-17 т
[Vor. 33
180 ANNALS OF THE MISSOURI BOTANICAL GARDEN
5. Chachahuatlán, Jalisco. Maíz reventador
Maíz reventador, the ancient popcorn of western Mexico, has been the subject of a special
monograph (see Bibliography, Anderson, '44
~
19461
ANDERSON—MAIZE IN MEXICO 181
ee Фе ө
odds 1 1 1 1 1 J
10-11
6. Chachahuatlán, Jalisco. Maíz colimote
rom an old hacienda a few miles down the river from Tuxcacuesco. The corn was prevail-
Na ie though most of the ears had at least a few = kernels. Four of the ears had
kernels with colored n (mostly Pr, one pr). Four of the ears had faint pericarp color
(P The name me "from Colima," which is not pi distant. Note change in position of
ike к
[Vor. 33
182 ANNALS OF THE MISSOURI BOTANICAL GARDEN
-
[^] ә ©
e?
oe
ву =.
© ә 9 427 949 E
© © ә ө
а L © 1 1 Г
10-11
7. San Gabriel, Jalisco. Maíz blanco
wo of the ears showed a tan pericarp; one pow ЕР pericarp (P?!9), The corn was prevail-
Te white, а few ears showed yellow kernels. 5 shifted to left because of very wide kernels.
1946]
ANDERSON—MAIZE IN MEXICO 183
e
ee o 9
Ф
ө ө
&
ә жө 9.9 ы ы
See өө өө
1 L. @ | )
8. Tuxcacuesco, Jalisco. Maíz blanco
The kernels of this variety were all heavily and
Prevailingly white, a few ears with occasional yellow
Pry.
conspicuously capped with soft white starch.
kernels and a very few with colored aleurone
2%
ij
B
E
ж
=
=
$
de e
5;
[Vor. 33
184 ANNALS OF THE MISSOURI BOTANICAL GARDEN
9. Palmar, Jalisco
From clearings in the mountains а few miles west of Tuxcacuesco. Prevailingly white with
many yellow kernels and some yellow ears.
1946)
ANDERSON—MAIZE IN MEXICO 185
ө 9
өө Ф
©
© ө Ә е O9 o
ә 9
оо оф ee
=
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| | | 1 1 1
10. Ameca, Jalisco. Maíz amarillo
ought in the sack by the family who let me measure it. Probably came from some of the
mountains near by. Shows a strong infusion of Mountain Yellow. Prevailingly yellow. Eight of
the s had tan pericarp, one a pale pink pericarp. v of the ears were segregating for bright
versus она yellow
[Vor. 33
186 ANNALS OF THE MISSOURI BOTANICAL GARDEN
оо x y ө
оо
| © о
оо оо OO
оо -
о о2о оо
11. Ameca, Jalisco. Maíz liso
ooth, glistening ia as d name M: Said to be very common in the region pier
Ген per the coast. wc ound in a casual survey o n being stored or
in the central part of the do ty. ET to т used sometimes as a popcorn. "Prevailingly white with
a few yellow i on nearly every
1946]
ANDERSON—MAIZE IN MEXICO 187
o
ее оф # в
“ e
Өс яо сө
* e d e
e
e
e e
LE
10-11
12. Ameca, Jalisco. Maíz areneno
This variety 4 so wide- ups that the scale has had to be shifted as noted in the diagram
Yellow and white, heavily capped with soft starch and uniformly dented with a smooth, even dent.
A few kernels wih Aerei pore (Br).
[Vor. 33
188 ANNALS OF THE MISSOURI BOTANICAL GARDEN
© ө
| e w^
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ә ө о 6€ e
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13. Tlajomulco, Jalisco. Maíz dulce (See plate 8)
үшү has been the subject of a special monograph by Kelly and ои It had been
his v
grown in а den with a red aleurone "elote" variety, apparently for more than one season since
ear was segregating ll the contaminated kernels were pr er vidi Pr. Maíz d E. is
weet
e
red like Mexican
contaminated kernels. The pr 1s En were usually
All the ears had yellow endosperm with varying
).
опе е
2. for sweet endosperm, but а
and these ears all had high
Pyramidal. The denting was scored o
rather than evenly flushed ки 4.
ейіп he pericarp (various alleles of Рс”
few of the ears showed partial d of the
ve) and were y
spotted
amounts of dull r
1946]
ANDERSON—MAIZE IN MEXICO 189
~ ©
ө о
ө
ө
ео e
Nue
е 9 ө
0-11
14. Tlajomulco, Jalisco. Maíz criollo
ajo — is an ancient town of great importance in colonial times but now isolated from the
aed and t ighway. Samples of maize from it are less ES to show the influence of
modern varieties. es that he extremely wide kernels have made necessary to poesi thé
ina as indicated in the diagram. This ых was prevailingly seri most of the ears showing
a few yellow EL Two of the ears showed colored aleurone on one or two kernels (pr). One
ear d a brilliant red flush (probably some allele of R).
[Vor. 33
190 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ob e
10-11
15. San Pedro, Tlaquepaque, Jalisco. Maíz criollo
rom a little pottery-making town a fcw miles east
i r
F of Guadalajara. Yellow and white with a
few ears showing occasional kernels with colored aleurone is vari ied i
i > 1 t 5
Thi iety was studied in the field
the veins on the lower sheaths with slight to no interveinal color. e upper sheaths showed
ith
little color. ‘Tassel branches varied from 5 to 24, with an average of 13. The male glumes were
10 to 14 mm., with an average of 12.
1946|
ANDERSON—MAIZE ІМ MEXICO 191
10-11
16. 5.Р. Tlaquepaque, Jalisco. Maíz ancho
rown on a field which was better watered than the rest of the fields in town and therefore
had a longer growing season. ‘This and the next diagram are two selections from the same field
made at husking time this one I accepted all the ears handed me by the workmen to avoid
offending them. The Aer selection is strictly at random. Both кенінен аге кайры boran
with one EET ear and a few yellow kernels on most of the There was one ear with a
crowned red pericarp (Pr). The field was outstanding in uniting « a i hish id. of Radon levity
with great “variability i in type of ear. One extreme ear shown in fig.
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
10-11
17. S.P. Tlaquepaque, Jalisco. Maíz ancho
second sample from the same field; see the previous accoun
tive. This photograph is not to the same scale as
nt. Extremely variable and
the others in the Appendix.
produc-
19461
ANDERSON—MAIZE IN MEXICO 193
а)
18. S.P. Tlaquepaque, Jalisco. Maíz chino
Prevailingly white. Maíz colorado (pr) was planted at the end of the field. Cross pollinations
indicate this variety is mostly Pr, C'. Sample of 40 ears.
+>
ayy
у =.
эү
ud
№
=
Ж
^
^
y»
+
фи
ТЕТІ,
ғ»
жә
‘cps
[Vor. 33
194 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ж әс өө 9
ө
өө о ә ©
© ©
oe ө Ө
| 1 1 1 1
10-11
19. 5. Р. Tlaquepaque, Jalisco. Maíz colorado y maíz negro
A badly mixed field of “elote?” corn grown by a poor widow. Note that the scale has been
changed because of the wide kernels.
19461
ANDERSON—MAIZE IN MEXICO 195
-
-
© өө Ө
е5" о
өс
[^] y^ ui оф e
се сө
1 1 1 1 1 4
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20. S.P. Tlaquepaque, Jalisco. Maíz criollo
vailingly white. Hal
— ord aleurone (pr).
f of the ears showed
yellow kernels. Two of the ears had
a few
One ear had faint pericarp ерм (Pcr).
196 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Ф
ә
où осе
о
Г ©
e
eo об ө ©
e e »
ое өс ее
[ | | | 1 )
10-41
21. Tonolá, Jalisco. Maíz criollo
onolá is a still more. iba pottery town near Guadalajara.
Te Prevailingly white with few to
many yellow kernels. A f r kernels.
a>
pra
%%
2
ж
M
ж-
=
ж
ж
=
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bod
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ж
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[Vor. 33
19461
197
ANDERSON—MAIZE IN MEXICO
,
valenciana
ә
i
Ma
22. Tepatitlan, Jalisco.
Prevailingly yellow, many white kernels and a few colored
market.
On sale in the local
aleurone (Pr).
LS
+46,
ns р
> У» morro
ысы
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
198
фо оф ө ө
еф оф Өө
o e
bo eb +:
o o
Maíz amarillo
1
1
Tepatitlán, Jalisco.
23.
ears had kernels with
the ears with white kernels. Four
of
A
Pre
colored aleur one
minority
(Pr), two ears Pcr,
yellow.
vailingly
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19461
199
ANDERSON—MAIZE IN MEXICO
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9
24. Tepatitlán, Jalisco. Maíz blanco
he preceding variety.
From the same small
with yellow kernels.
Prevailingly white, a minority of the ears
farm as t
Grown in an adjoining field to the preceding.
eus Дари
км" "еметін
ay АИ
=> See AS
SES T mem
AIL
[Vor. 33
200 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1 1 4 1
~
5. Apango, Jalisco. Maíz amarillo
Apango is a tiny town on the north flank of
ny the Volcan de Colima. The ears were selected
from a field and represent the extremes. Yellow; i
one ear strongly colored crown pericarp (РС”).
UY D
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1946)
ANDERSON—MAIZE IN MEXICO 201
обе обе
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229 өсө ө ©
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1 1 1 1 1 1 1
26. Tapalpa, Jalisco. Maíz amarillo
Tapalpa is a large and handsome town in the middle of a small, ne = plateau on the
north flank of the Volcan de Colim This variety was prevailingly yel with many white
kernels. Six of the ears had colored Secale of some shade or other ES pre of P and Рс").
obs.
Seven had red c
33
| Vor.
ANNALS OF THE MISSOURI BOTANICAL GARDEN
202
eo e e
27. Tapalpa, Jalisco. Maíz amarillo del cerro
The majority of the ears have colored pericarp
Prevailingly yellow with many white kernels.
(P and Рс”) of various intensities.
“Ж...
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19461
203
ANDERSON—MAIZE ІМ MEXICO
9 o
e 6
ә 9
28. Tapalpa, Jalisco. Maíz amarillo de tierra más templada
Light
As the name indicates, grown on fields well below the town, hence with a longer season.
yellow and white, mostly red-cobbed.
tU VR (di л
УЛ
111111
ГҮҮ"
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pie MER Ti A
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117 17
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ПАРУ:
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м week:
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[Vor. 33
204 ANNALS OF THE MISSOURI BOTANICAL GARDEN
-
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29. Tapalpa, Jalisco. Maíz criollo blanco
White, a few of the ears with а few bright yellow kernels. One ear РС”,
19461
ANDERSON—MAIZE ІМ MEXICO 205
=
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9.
i бе
1 1 1 1 1 1
30. Tapalpa, Maíz espiga blanca
Unlike most of the p or varieties, t t have a brightly colored tassel. Pre-
ee к а few of the ears with white pe pos ба with colored pericarp (P and Рс”),
Cobs whit a very faint =. The ears of this eu were similar to those illustrated in pl. 6,
ја угы pod at the 52 and the top of the plat
[Vor. 33
206 ANNALS OF THE MISSOURI BOTANICAL GARDEN
оо
©
оо © “
[^]
© оф о ё
| 1 1 1 1 1 _——
31. Coalcoman area, Michoacan. From clearing A.
his and the two following samples were sent me by Sr. F. Vargas Tentory, a member of a
cultural mission to backward areas. These three 5. were made іп three о —
clearings in the Coalcomán area in western Michoac The 14- and 16-rowed specimens in each
collection are very similar to maíz 2. the remainder of the collections looks like crosses
between that and typical Mexican Narrow Ear varieties
1946]
ANDERSON—MAIZE IN MEXICO 207
оо о 9
О
оо oe
ә ә
№
- ә © о
1 L 1 1 Д. 1 = |
32. Coalcoman area, Michoacan. Clearing B.
See notes on the ране collection. 7 wee were variable as to color. w had
dark red pericarp (strong allele of P), a few had colored aleurone (Pr), and 2 ol a
crowned pericarp (P°"). The rest were i and yale, One of the Pr ears had a single white
kernel, indicating the presence of dominant white (СІ).
[Vor. 33
208 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1 1 1 1 1 1 1
33. Coalcoman area, Michoacan. Clearing С.
See notes on two previous collections. These three collections do not represent samples ifa E
single field, as do most of the examples in this Appendix, but are from all the varieties gro
each clearing.
1946]
ANDERSON—MAIZE IN MEXICO 209
Г *
әс өсе се
"5 ш e
қ o o
ео eo ФО
Ө оо ө
————— BÉ | 1 1 1 1 1
Prevailingly yellow
34. Apatzingán, Michoacan. Maíz amarillo
Two ears had white kernels. Eight of the ears had few to many kernels
On
with colored aleurone (Pr). ne had tan pericarp.
%
+:
&
$.
4:
ii
i
4:
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1
:
210 ANNALS OF THE MISSOURI BOTANICAL GARDEN
со ә
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өс ә © 6
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35. Apatzingán, Michoacan. Maíz amarillo
Prevailingly yellow. Eleven of the ears had white kernels; one had a tan pericarp.
pintada в 5%
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=
t
2
=
2
=
Е
имеет
ріпті тім
ОРА 222”
[Vor. 33
1946)
ANDERSON—MAIZE IN MEXICO
e
є>
©
Сс Ф
©
[ 1 > 1
4 —
36. Jiquilpan, Michoacán. Maíz pipitillo
Prevailingly white, seven of the ears showing yellow kernels. Us ears had a
P па и but they were all very faint іп color. Apparently the white of
almost dominant. ле name p/pitillo usually denotes a digo "eared, man
more or less pointed kernels, dented and wrinkled below the poi
few kernels with
this variety is
h
y-rowed variety with long,
[Vor. 33
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ое ее фе
© ®
| Li ә
ec e) о
oe е.
Е “> ә
ip e р
1 1 1 1 | | 1
37. Jiquilpan, Michoacán. Maíz amarillo
at La 2. ia the hills above Jiquilpan. Prev = yellow. A few ears had white
jene four had a kernels with colored aleurone (Pr Гуо ears had faint pericarp color
(a low allele of P Su
19461
ANDERSON—MAIZE ІМ MEXICO 213
1 1 1 1 1 1 4
38. Jiquilpan, Michoacán. Maíz rosquera
This variety was identical with the maíz reventador of near- E Jalisco and not at all like the
pointed popcorn sometimes called by the same name near Mexico City. All the ears bore both
yellow and white kernels. Six had a few kernels with colored Boone (both Pr and pr).
[Vor. 33
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN
| Е
«o 005 5
$6 во
e Ô
е
1 1 1 1 д
39. Jiquilpan, Michoacán. Maíz grueso
Prevailingly white. А majority of the ears had a few to many yellow kernels. Five had
kernels with colored aleurone (Pr). One ear was flushed with brilliant color (allele of R).
19461
Nine of th
ANDERSON—MAIZE IN MEXICO
1 Е | 1 1 ES ^
215
show
40. Jiquilpan, Michoacán. Maíz negro
iae mi bis blue Aron ko па Pr). Several of the ears were a dicas. for pr
e ker
pollination ud dominant white s rather than segregat
fewer-rowed
under “elote”
ed whit s but in such small cete rs that it must hav
e been due to
N dae ida seeded a
this variety is, on the average, than the EGER Miis from Jiquilpan.
varieties.
nd
See discussion
[ Мог. 33
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN
©
=) е e ә
в
e o 59 со e •
% ео e
e © ә Ф
ее
Ны релі 1 | 1 1 Р ЕНЕ )
41. Uruapan, Michoacan. Maíz criollo
Prevailingly light yellow. Two ears with a few white kernels; two with a few kernels with
colored aleurone (one Pr and one pr).
1946]
ANDERSON—MAIZE IN MEXICO 217
©
ә ө e e =
ж
-
о о О © ~
e. -
о ө e v
- ә о
е
pul П ! ! | 1 ==)
42. Uruapan, Michoacán. Maíz breve.
Prevailingly reddish aleurone (br), though a major of the ears show few to many dark
kernels (Pr) and one was segregating. es had тек; вена y and Рс”),
[Vor. 33
218 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
o 6
è
ео 5%
e u
à e 6 5
° б ө
|.
“ eo 0
0
1 = 1 1 1 1
43. San Lorenzo, Michoacan. Maíz del cerro
Prevailingly white and yellow. A few ears with dark aleurone (Pr) on a few kernels. Two
ears flushed pink (allele of R?) Crop stunted by new volcanic ash.
1946)
213
ANDERSON—MAIZE IN MEXICO
44. Cherán, Michoacán. Maíz tulekenio
Seven of the ears had bright pink or red in the peri-
Six ears had a few kernels with dark aleurone (Pr).
White and yellow in various mixtures.
carp (various alleles of R?).
: S
D РАЉАМА НА"
л» эзсе, „ы LN
"о onroro a |
Ваља
ааа! ааа Кэ,
5%%%4% PUT 0902
мы MODA
(P, ie "
[Мог. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
220
pinto
iz
45. Cherán, Michoacán. Ma
Prevail-
2).
This is the black corn discussed in detail in the Appendix to Beal’s report on Cherán.
ingly dark aleurone (Pr), many with light to dark red purple in the pericarp (alleles of
2%
Saat SS
* МА МАЊУ |.
En
4%
tut
^I.
; У à
: E | ж
3 # Peete aan tae
МУ Те
ыи: du tu 3.
паче ETT
19461
ANDERSON—MAIZE IN MEXICO 22%
1 1 1 1 1 1 1
46. Cherán, Michoacan. Maíz blanco del cerro
Prevailingly white with a few yellow kernels. Two ears had a few kernels with dark aleurone
(Pr). Two showed color in the pericarp.
села ое
ы. Wee E ры
Ü
4
%
*
zi
+ |
Digi
% à
iii
* я
til
* 2
фе
id
„$
Yos
CM
[Vor. 33
222 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ее
ее әс
o ө
[ ә
e ео ФО
+ “
eco од
ее
e 9
1 1 1 1 L 1 і
47. Pátzcuaro, Michoacan. Maíz pinto
Grown on a small farm, "Las Paredis.” Prevailingly white. Nearly all the ears with some
kernels showing colored aleurone (usually Pr, sometimes pr).
1946)
ANDERSON—MAIZE IN MEXICO 223
с
©
48. Patzcuaro, Michoacán. Maíz blanco
From a field one mile above the town. Prevailingly white and T У за yellow. Three ears
with a few kernels with dark aleurone (Pr); two ears with colored peri
x.
pros 3
geet шақа к.
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
224
be
Maiz blanco
49. Lake Patzcuaro, Michoacan.
Four ears with crowned peri-
Most of the ears with some yellow kernels.
Prevailingly white.
Several ears with colored aleurone (Pr and pr).
carp (Pcr).
“а.
< УУ
Cj №
a%
AT EH ts
ЈИ ТИ Пети
мын aga ы
e
f.
RIT es
by free iJ adea
wa I P
НН
245r. y
19461
ANDERSON—MAIZE IN MEXICO 223
© © әс
49
ә
Г eee
ее өө
ә фе
о e “%
1 1 1 1 1 1 Í
50. Lake Pátzcuaro, Michoacán. Maíz colorado
2: т е (Pr), though many of the ears are i divini for reddish cde
(pr) a ar pr. Over half the ears with a few t ny white kernels, some
them io to x. d dominant white (20 • опе ear, for Ik had Pr xe pr kernels e
a single kernel of pure white with no trace of co
226
ee ө
e ©
6.9% e o
| 2
ә
L
се
e
L 1 1 L 1 1 )
ANNALS OF THE MISSOURI BOTANICAL GARDEN
neers by E. J. Wellhausen. Prevailingly white and pale yellow.
faint reddish pericarp. Many of the ears were 2. іп spite of being quite short.
slight error іп this figu re. Of the three ears indicated as having eight rows only
was eight-ro wo others the same square
wed.
should жей fis
51. Zitácuaro, Michoacán
are really ten-rowed,
t in
slightly higher in the diagran
[Vor. 33
Four of the sli rd a
is
the 1. опе
and these two dots
19461
ANDERSON—MAIZE ІМ MEXICO ` 227
>
e ©
Ё УСУ 0
6,6 фе
90,9
| boto
9 фо ёо
e
52. Quiroga, Michoacán. Maíz de tierra fria
Collection by E. J. Wellhausen. Prevailingly white and pale yellow with a few ears with faint
pink pericarp.
[Vor. 33
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN
-
L
m
о ж ә
© О о O
1 1 1 1 1 1 4
53. San Francisco, Villa Allende
A small collection made by Dr. Wellhausen. АП dark aleurone (Pr). On half the ears the
color is intensified by red pericarp (P and Рс”) which makes the dark blue into a shiny black.
19461
ANDERSON—MAIZE IN MEXICO 229
-
о
Ё ©
© оо (e) pe ©
< оо
Ё @
© ә9о
ә ә © Ж
ОО
1 1 1 1 1 1 |
54. Mexico-Toluca Highway, Km. 60. Cacabuazintle
Nearly all pure white. One ear with five yellow kernels, all brilliant yellow and showing no
capping of ek uA zu ears with a few kernels of dark aleurone as all of pe very dark.
Two ears sh , lower right. Two ears of this variety and from the same town are
illustrated in т
[Vor. 33
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN
| 1 \ 1 | |
55. Mexico-Toluca, Km. 57
pM асуы white 4. modifi ed by the pinky tan and smoky colors of the pericarp.
of the ears are st ғ fasciated and look almost like ‘ oes Hull-less’” popcorn. One
ear is шеші іп pl. 5, ue ove. Note extension + са scale
19461
ANDERSON—MAIZE ІМ MEXICO 241
ee 0 ө
©
| ф % о
s S obeso
| ео
99 о
Q
©
1 |: 1 1 1 | - 2 |
56. Mexico-Toluca, Km. 17
Collected by E. J. Wellhausen. Prevailingly white and light yellow. Four ears have a pinkish
tan pericarp. One ear is strongly fasciated.
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
232
о
Mexico- Toluca, Km. 17
57.
Prevailingly bright yellow, with some light yellow and white.
Collected by E. J. Wellhausen.
Varying greatly from ear to ear in the
ew kernels with dark aleurone (Pr).
amount of capping with soft starch.
Two ears with a f
ж.
ЕН КҚ К +
me Beet
S etat
quita E
Хоу;
=
Pete et ЫЫ e tii
er
ный M
петиции, у 9
*
аш minsay
%
+
ae ои,
жек каиыр а?
dapi
esty
»k
dat
Baa *
lla m mA ч
aos ыы их РА
Ри _ ғ”
ig petet
HOMI
A ata едм А,
ге (4-4
"ipsae. A
1946)
ANDERSON—MAIZE IN MEXICO 235
=
-
©
© ô ô 9
49
e
об бе ©
+
6,6 ео
0% 6
Ф ә
Г | | 1 | 1
58. Mexico-Toluca, Km. 57 (Bag Хо. Ш)
Collected by J. Wellhau Prevailingly white and yellow, the latter mostly very pale.
Many kernels қ ім dark ood (астана Pr and а semi-dominant white).
[Vor. 33
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN
59. Chalco, Mexico
Collected by E. J. Wellhausen. АП dark aleurone, one ear segregating for white. Most of the
ears with occasional white or nearly white kernels. Six of the ears with red pericarp (P and Рс”
19461
ANDERSON—MAIZE IN MEXICO
1 1 1
Collected by E. J. Wellhausen.
60. Metepec
All white and light ivory yellow.
235
33
[ Vor.
GARDEN
ANNALS OF THE MISSOURI BOTANICAL
236
61. Tepotzotlán, Mexico
Two ears with pericarp color.
Prevailingly white and light ivory yellow.
“eons
en 5%
aie
ашқан wee
Pig а 4 жен"
вать
=~ T a t <= ~
t as уценили rp)
orm wm
Жейн» mrt fugaces
d = s
ANS mamam >
Pn en Ee
~ А Р
M E Y ~
Be eS асаа оос КҮ,
%. ” ^ T У -
"е. —"
"vest Қым 222224”
- ланы ы” e
sar
іе
19461
222
ANDERSON—MAIZE IN MEXICO
62. Tepotzotlán, Mexico
*
Small samples of two early varieties used for pinole and elotes.
xg Б>
“oe
* vat ымға
абы etat" m.
* "an авз"
[Vor. 33
MISSOURI BOTANICAL GARDEN
ANNALS OF THE
238
е 9 <
о
63. Huehuetoca, Mexico. Maíz blanco
The ears photo-
Four ears showed a few kernels with dark aleurone (Pr).
graphed are the superior ones and not a random sample
Nearly all white.
A.
““%;
M
by all
4% ымы
сабай
ten hee
He.
gene теа
+.
жа ede еә,
^. PPS И ;
ом
ы
9909019605
Ld
LEE PALI rn
5 яз»
аҚ;
Мр
бед
be TL.
19461
ANDERSON—MAIZE IN MEXICO 239
e 090 о
о
ә ©
обе о ©
[4
ео
L о
ее ое
Ô ©
0 e
1 1 1 | 1 1 саба
64. Acupula, Mexico. Maíz morado араз айо
Predominantly reddish aleurone (pr), though every ear showed а few Pr and two ears were
largely so. Most of the ears with a few white kernels (Ct). Two ears with red pericarp. Said
to be early seasoned. One ear illustrated in pl. 5, right below.
[Vor. 33
240 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ео Ө ©
Ө
| e
де со ө
• “.
| ©
ое 9
[ 1 1 1 Кк... ыы
65. Huehuetoca, Mexico. Maíz negro
Prevailingly Pr. Many
B1) І
presence of dominant white (
of the ears with a few kernels of clear white, indicating the probable
"i
19461
ANDERSON—MAIZE ІМ MEXICO 241
+ %
eo Фе
&
Г 9
ё o .2 “
e 6
ое € E
ray ее ә
=
1 1 L 1 1 1 РР
66. Leon, Gto. Maíz vaquereno, tipo I
An improved variety produced by Sr. Eduardo Limón. White with heavy capping of soft
starch.
ANNALS ОЕ THE
[Vor. 33
MISSOURI BOTANICAL GARDEN
1 L 1 1 1
67. León, Gto. Maíz vaquereno
collection made from several small farmers in the 4. It тө the original
"Вало,"
Мехісап m belt
ears with yellow ker ls
> his selections and is ther representativ the maize of the
A few
© Prevailingly white ud 22. capped with MA starch.
19461
ANDERSON—MAIZE IN MEXICO 243
о бө се өе
e ©
m
1 1 1 1 1 1 1
68. Salamanca, Gto. Maíz blanco
collection made by E. J. Wellhausen. Lus ep with a strong api ency to fasciated ears.
There is a slight difference in scale in this figure. the vertical scale the row numbers run
from 10 to 26 instead of 8 to 24 as the m entire "om should be pushed up-
rhe
wards one unit to make the grid Маи to all 5. pc Pe,
Horizontal scale unchanged.
[Vor. 33
244 ANNALS OF THE MISSOURI BOTANICAL GARDEN
| б
$ > Ф
8 e
0 ô ёо 6
4
об 60
% m
ô be 8
-
* 5
ea _ 1 1 1 1 1 1 }
69. Tlaltizapan, Morelos. Maíz pipitillo
White and light yellow. One of the extreme ears is illustrated in plate 4, below. None of the
extreme type with long, pointed kernels showed any yellow stronger than a pale ivory. The less
pointed kernels (of the type known as maíz perla) nearly all showed light yellow kernels here and
there. It seems probable that the extreme "pipitillo" has a strong yellow-inhibitor.
19461
ANDERSON—MAIZE IN MEXICO
ее e
ә
9 %
ә ө "^ ә 9
ee обе ә ә
3 ж
e Ф []
1 1 1 [ 1 Ou ETIN
70. Tlaltizapán, Morelos. Maíz colorado
Prevailingly pr with several ears showing Pr and a few with white kernels.
All were red-cobbed.
246
ANNALS ОҒ THE
MISSOURI BOTANICAL GARDEN
71. Otlatlan, Puebla. Cacahuazintle
Obtained through К. Н. Barlow from а remote mountain village. — Prevailingly
many
whit
[Vor. 33
white with
ge
kernels of yellow and of dark aleurone (Pr). Obviously extensively mixed with the local
e corn, samples of which were also obtained.
1946]
ANDERSON—MAIZE IN MEXICO
L
© ө
о ~
1 1 4. 4 1 1 4
72. Chihuahua
Obtained through E. J. Wellhausen. White and yellow. Very similar to the maize of the
eblo Indians in Arizona and New Mexico and probably derived therefrom. This conclusion is
supported by unpublished archaeological evidence from several sources.
tree
Tea) rut
САНЯ
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teens
$4444
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ҚАЛУДЫ ТА
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Annals
of the
Missouri Botanical Garden
Vol. 33 SEPTEMBER, 1946 No. 3
A CYTOLOGICAL STUDY OF YEAST (SACCHAROMYCES CEREVISIAE)!
LILLIAN NAGEL
Associate Professor of Biology, Harris Teachers College, St. Louis
Formerly Instructor in the Henry Shaw School of Botany of Washington University
INTRODUCTION
A century ago Nageli (1844) first described the yeast cell as having “‘a little
nucleus of whitish mucus, lying on the membrane.” From that time to this the
cytology of yeast has been a subject of controversy. Extensive bibliographies and
discussions of the earlier literature may be found in Wager (798), Wager and
Peniston (10), Guilliermond (720), Moore (733), and Badian (737). Brandt
(741) has reviewed the more recent literature, as well as given the results and in-
terpretations of his own extensive investigations. Recently Lindegren (745) and
Subramaniam and Ranganathan (745, '46a, '46b) have published differing in-
terpretations of nuclear organization in yeast.
In spite of the great amount of work which has been done on the cytology of
yeasts, there is still no general agreement (Table VI) among students of the sub-
ject even on fundamental points. There are several reasons for this lack of agree-
ment: (1) The most important is the small size of the yeast cell which makes
accurate observation of details extremely difficult and causes interpretation to be
more or less speculative. (2) The use of only one or two stain techniques by
many investigators has increased the confusion because interpretations are based
on incomplete information. (3) Until the work of Winge and his associates
(35, 737) investigators were handicapped by lack of understanding of the life
cycle of yeasts. (4) Not until the recent work of Lindegren and Lindegren (44)
was it possible for a cytologist to secure adequate sporulating material.
The present work was undertaken with the hope that additional information
could be gained concerning the structure of the yeast cell through the thorough
and methodical use of a variety of techniques on the excellent material available
from the Lindegren laboratory.
investigation carried out in the graduate laboratory of the sae Shaw School of Botany
An
of eae gton University, assisted by a grant from Anheuser-Busch Inc., and submitted as a thesis
in partial a5 mé of the requirements for the degree of Doctor of Philoso phy.
(249)
[Vor. 33
250 ANNALS OF THE MISSOURI BOTANICAL GARDEN
MATERIAL
Saccharomyces cerevisiae was used in all of the work reported in this paper
except as noted below. The cultures were secured from the Lindegrens, and all
culture designations refer to their material. Most work was done with the dip-
loids LK, with the hybrids EFF, 30 x 20 сс., and with BC20—, and several
similar haploids. EFF is a hybrid of three commercial baking yeasts; LK is
isolated from a Canadian pressed yeast. The haploid BC20— is a single ascospore
isolate from a strain studied by the Lindegrens as were the other haploids used.
While working on the identification of wild yeasts found on native fruits ob-
tained from the Missouri Botanical Garden Arboretum, an especially large one
which showed peculiar multiple budding was isolated. It is probably an unde-
scribed species of Syringospora or a related genus. Limited cytological observa-
tions were made on it for comparison with Saccharomyces cerevisiae. A few
observations were also made on a species of Cryptococcus (?), another wild yeast.
METHODS
In so far as possible all techniques and observations were carried out with all
material used; that is, a general systematic survey was attempted. Parallel ob-
servations of living material were also made.
Culture Metbods.—Budding cells of various diploid cultures were sometimes
obtained from yield tests in the Lindegren Laboratory (Lindegren and Lindegren,
43). Both haploids and diploids were also cultured in liquid nutrient medium
TABLE I
PROCEDURE FOR SECURING SPORULATION
(LINDEGREN AND LINDEGREN, ’44)
MEDIUM
TIME Ма ы d TECHNIQUE
Kind Formula Preparation
Beet (leaves) extract ....10 cc. Streak on slant; in-
Beet (roots) extract......20 cc. cubate at 25°
pe Iu oar vs 35 cc. (room temp.). Aft-
EIU etm 1615 cc. Steam mixture 10 | er 2-3 days pour
48—72 | Presporulating Yeast -=n UNUM E 2gm.| minutes. Tube and | 1 cc. sterile water
hrs. Glyc 215 gm.| autoclave. Slant. over yeast. Stir to
TN 5 gm. make thick suspen-
1 gm. sion of cells. Take
Vater [EM 100 cc. up in sterile pipette.
final v
Pour over upper part
of gypsum slant.
Mix, pour, and slant г dieser
16-48 | Gypsum slant Water 100 сс.| rapidly. Dry 24 enough acetic acid
hrs yP Plaster of Paris .......... 100 gm.| hours at 50° С. ve и
Autoclave. Oe еи
4.0 into lower half
of slant. Incubate
19461
NAGEL—CYTOLOGICAL STUDY OF YEAST 251
and on nutrient agar slants. Little work was done with copulating yeast cells or
with old or starved cultures.
Material for the study of stages of sporulation was cultured in accordance
with the method developed by the Lindegrens on medium supplied by them
(Table I). Тһе diploid strains LK and EFF were usually used because they
sporulate well.
Smear Technique.—All permanent preparations were made as smears on cover
glasses and stained in cover glass Coplin jars (cover glass wells of Thomas & Co.).
The yeast cells were stuck to the cover glass with a thin coating of Mayer’s
albumen fixative or applied directly to a thoroughly cleaned cover glass. The
cover glasses with the layer of albumen were generally heated gently over an
alcohol burner “until a gray smoke is given off” as suggested by La Cour (741)
before applying the cells. Many more cells remained on the cover glass if fixative
was applied, but with some techniques the albumen tended to stain and if drying
occurred artifacts frequently appeared. Many cells were lost in succeeding
operations if applied directly to the cover glass.
For applying the yeast to the cover slip a micropipette was found useful. An
ordinary glass dropper was heated just above the narrow end, drawn out thin, cut
off to a length of seven or eight inches, and the rubber bulb replaced. A very
small drop of water was placed on the cover glass with the micropipette and cells
were added from a slant with a sterile loop; in other cases a droplet of cells was
pipetted onto the cover slip from a suspension. The cells were spread evenly by
passing the thin flexible end of the micropipette across the surface of the cover
glass. This was done rapidly to avoid drying, which causes severe distortion of
the cells. The cover glasses were immediately floated cell-side down on the surface
of the fixing solution in a Petri dish. If long fixation was required, they were
transferred after about fifteen minutes to a cover glass Coplin jar for convenient
storage.
Fixation and Staining Procedures.—Many different fixations were tried, and
the types which apparently caused least shrinkage, distortion of the vacuole, and
granulation of the cytoplasm were generally used (Table II). Drying, even after
fixation, tended to cause shrinkage and distortion. Good fixation was apparently
related in part to the condition of the cells; fresh, actively growing cells showed *
better fixation than old cells from giant colonies, and actively budding cells better
than the highly granular sporulating cells. None of the methods used was com-
pletely satisfactory.
The general staining procedures are outlined in Table III. A further discussion
of outstanding points is given in the following section of the paper.
252
[007733
ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE II
FIXATION*
Fixative Formula and/or Modifications Time Evaluation
Mercuric Saturated aqueous НЕСІ + 0. 15-24 Most satisfactory general
chloride -1.0% glacial acetic Med hrs. fixation
Iodine- 1% о таи be 1% KI..20.0 сс. 12-24 Good with Giemsa technique
ormalin- Formalin Ба 4.0 с hrs. (2, Table III)
acetic acid Glacial ir acid ...... 0.5 сс.
Distilled water ........ 5-10.0 сс.
Navashin Plant tissue formula (Lee, 737) 1-12 Fair. Distortion of vacuole
"Craf" Modification (Johan- hrs. common
sen, '40)
Osmic acid Vapor from 1% osmic acid 3-5 Fair. Cytoplasm granular,
vapor solution min vacuole often distorted or
invisible
Picric acid Sat же, E ees water or 4-12 Fair with methylene blue
70% a hrs. eosin ie i oig TA "Table Ш)
*Other fixations tried but not used because of perc rper. and/or granulation of
др were pua no lemming's niu
y,
»
Carnoy-Lebrun, Perenyi,
cro-formol-acetic acid mixtures, glacial сей acid vapor, " chrome- Bowen-urea
weak solution,
(Sem 42
fluid, у “кейді bath alone and in various combination
TABLE III
STAINING METHODS EMPLOYED IN THE STUDY OF THE YEAST CELL*
the
Uranium fixative
Stain Usual fixation Staining procedure Remarks
1. Feulgen Mercuric chloride 1. Hydrolyze 7-8 min, in не positive body
(Johansen, '40; |Navashin НСІ at 60° C. (Hillary, 739) ре, in all cells.
де Tomasi, 36; (Unnecessary ta wash |2. Stain E in n Feulgen. С коң сІеаг апа
Coleman, 73 out fixing solution |3. in each of three} colorless.
before hydrolysis—
Bensley & Bensley,
38)
ash 1
HCI- KS. rad hs.
2. Robinow's bac-
terial Giemsa
(Robinow, '42;
Dubos, '45)
Mercuric chloride
odine
Osmic vapor
~
. 7-10 min.
. Stain іп Giemsa (1
tures as fo
20:1
. Hydrolyze in 1 М НСІ at 60°
drop stain
per ml. dilute buffer— рН 6.9-
7.0) 15-45 min
Pass through acetone- xylol mix-
llows:
li COTES 4—30 sec
B Amon 10-30 sec
Ка 10—15 ѕес
е ayla <i. ЕК, 10-15 min
Very good; apparently
differentiation.
19461
NAGEL—CYTOLOGICAL STUDY OF YEAST 253
TABLE III (continued)
1. Stain in әй aqueous adii
violet 10 min
Results only fair;
nothing added to
2: er through: died from other
0.5% picric oan in 70% techniques.
Methyl violet “Te ohol 1
(Johansen, '40) Navashin ee неге E in 95%
hol 1
о: rir ты ТШ 4 drops
H per 100 cc.) 15 sec
4. 100% alcohol qd
e. Clove oil 30—45 s
. Mordant y hrs. "x 196 osmic|Results poor;
acid in 2% chrom nothing to himétory-
. Stain in standard інді solu- lin; could probably
tion be imp by
4. Flemming's 3. Stain in 1% aqueous crystal ificatio
i Navashin violet 10-15 min
triple stain
(Johansen, '40)
4. Dip twice in 95% alcohol, 3 or
4 times in absolute.
5. Drop sat. sol. orange G in clove
il on с 10 sec.
6. Wash in clove oil.
ДЕ Мајнц іп КАР ferric am-
Shorter mordanting,
je pa ns - destain
E Huh Ресей ii а. zu hrs. ng ga different
f jnenge Mercuric chloride езе eb. T he picture TL long.
пика 40 Navashin Ди P E 596 hematoxylin. Cytopla asm tends to
; : : hrs. or overnight.
cL М i3 hrs
3. Destain in sat. aqueous рісгіс
acid (Tuan, '30).
1. Mordant 6 hrs. in 1% ferric|Results similar to
ammonium sulfate іп 70% Heidenhain's hema-
alcohol. toxylin.
6. Brazilin pit M ссн 2. Stain 6 hrs. іп 0.5% Brazilin in
ematoxylin 70% alcohol.
3. Destain in sat. picric in 70%
alco
7. Methylene Picric acid . Overstain in 1% aqueous methy-|Results uncertain.
blue-eosin Navashin lene blue 24—48 hrs Tends to lose stain
(Badian, '37) Mercuric chloride 2. Differentiate very ШОЛУ with | in alcohol
dilute eosin (0.5% or less).
8. Aceto-, lacto-, and | Alcohol-acetic acid. |All combinations of the acids and Ep unsatisfac-
Propionic carmine, | Also various ants| stai i ious ons Aceto-orc ein
orcein, Bismar
brown
success.
n: gr after ae No per-
t slides prepar
id.
a. some better
than other combina-
tions
ынны were ru
n up thr ough
P
пећина stated. Usual washing or rinsing steps not
alcohol series into xylol and mounted in balsam unless
t listed.
[Vor. 33
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN
OBSERVATIONS
INTRODUCTION AND TERMINOLOGY
As mentioned in the Introduction there is as yet no general agreement among
yeast cytologists even as to fundamentals. Тһе body most frequently referred to
as the nucleus (Guilliermond, '10, 20; Badian, '37; Beams, Zell, and Sulkin, '40)
is called the centriole by another school (Lindegren, '45) and the nucleolus by a
third (Wager and Peniston, '10). In the following observations new terms which
are purely descriptive and have no previous connotation have accordingly been
chosen in order to present a picture of results, independent of interpretations of
nuclear organization in yeast. The term "parvicorp" (small body) will be used
(text-fig. 1) to indicate that part of the yeast cell frequently designated as the
perv icorp
companion body
entral granule (in motion)
agnicorp
particle іп magnicorp
refractive cytoplasmic
ranule
cyto hn m
Diagram of the yeast cell with parts labeled according to the term-
Text-fig. 1.
inology used in this paper.
nucleus. The term “‘magnicorp” (large body) will be used to designate the part
usually called the vacuole, but also named the nucleus (Janssens and Leblanc, '98)
and the nuclear vacuole (Wager and Peniston, 710; Lindegren, '45). А diagram-
matic representation of these parts is shown in text-fig. 1.
Descriptions of budding material precede those of sporulating material The
results for each technique are given separately and in the order listed in Table III.
The drawings on the plates follow the observations of the stained slides as closely
as possible. All observations were made with a binocular microscope equipped
with a 90 x apochromatic objective and 15 x compensating oculars.
1946]
NAGEL—CYTOLOGICAL STUDY OF YEAST 255
SACCHAROMYCES CEREVISIAE
STUDIES or BUDDING CULTURES
Feulgen Technique.—In all cells except small buds the parvicorp is at least
faintly Feulgen positive (pl. 10). However, not all of the parvicorps exhibit the
same color depth; a few are small and stain much more strongly than the larger
ones. In some cells the parvicorp is evenly colored throughout, in others there is -
apparent variation in color intensity (pl. 10, figs. 4, 5, 11, 13). Тһе buds often
lack the parvicorp, but in that case it is frequently dividing in the mother
cell or has finished its division or may not have started to divide, in which
event it is commonly larger than usual (pl. 10, figs. 1-15). The division
may be completed in the mother cell and the parvicorp then pass into the bud,
but in many strains it more often appears to divide directly into the bud. In
actively growing cultures the parvicorp is usually located proximal to the small bud
and divides directly into it (Lindegren, '45). Where the budding is not quite so
rapid the Feulgen positive body may be located between two parts of the magni-
corp or even opposite the bud. After the buds acquire parvicorps the latter usually
assume a distal position in mother and daughter cells (pl. 10, fig. 16). Тһе parvi-
corp may vary in shape from round to oval, may be crescent-shaped, irregular in
outline, or divided into two bodies, as noted earlier by Margolena (732), Winge
(35), Badian (737), and others. These bodies have often been interpreted as
chromosomes (Table VI). Conventional mitotic figures were not seen in the
dividing Feulgen-stained bodies of Saccharomyces cerevisiae. The Feulgen stain
is faint and the parvicorp small, so that details are somewhat speculative.
The magnicorp, which was unstained but generally quite clear in outline, showed
no Feulgen positive material with the procedure used. A discussion of the
magnicorp will be deferred to the sections on methylene blue-eosin and Heiden-
hain’s hematoxylin in which its structure is more readily observed. However, it
might be mentioned here that the parvicorp almost universally lies іп contact
with the magnicorp.
Robinow's Giemsa Tecbnique.—The use of this technique was suggested by
La Cour (personal communication) after a number of others were found inade-
quate. The Robinow Giemsa bacterial stain (742; Dubos, '45, with appendix by
Robinow) was used with the following minor modifications. Osmic acid was
slightly less satisfactory than mercuric chloride or iodine-formol-acetic acid as a
fixation. Samples of stain were obtained from three sources and were found to
vary somewhat in their staining ability. Destaining was not always even. Insuf-
ficiently destained and over-destained cells sometimes occurred on the same slide
probably due to inadequate pH control. Difficulty with fading was overcome by
controling the pH of the various solutions, especially the balsam which was
neutralized with sodium bicarbonate following the directions in Lee (9th edition,
'28) and by storing the finished slides in darkness. The better slides produced by
this method were the most satisfactory for observation of the parvicorp. How-
ever, good results were obtained only with the correct balance of all variables.
[Vor. 33
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Giemsa stains the parvicorp in much the same manner as the Feulgen reaction
but gives a much deeper color that shows more internal differentiation and greater
contrast with the cytoplasm. Because of this sharpness of contrast the irregularity
of the outline of many of the parvicorps is much more apparent than with
Feulgen. Although many of the cells show parvicorps with relatively regular
outlines more of those with irregular outlines are illustrated in pl. 11 in order to
indicate the kind and degree of variation observed. The difference in color in-
tensity within the stained bodies was often quite marked, resembling somewhat
the chromocenters of higher plants. One or two small extrusions or appendages
were often found. In older agar slant cultures, apparently aberrant divisions of
the parvicorps without consequent division of the cell seemed to occur, giving such
forms as shown in pl. 13, figs. 92, 93, 95-97. Haploid cells and their parvicorps
were smaller than diploid, but measurements of the parvicorps were not attempted
because their size variation in different stages of cell division made exact com-
parisons of doubtful value. Division of the parvicorp appeared much the same
as with the Feulgen technique. A few of the parvicorps exhibited the bipartite
structure noted in the preceding technique but the phenomenon seemed less in
evidence, especially in the haploids.
р The magnicorp was generally not visible with this technique, but when it was
its relation with the parvicorp was usually clear and unquestionable. Іп these
cells it lay in direct contact with the parvicorp as described above (pl. 11, figs. 30,
40, 42, 46). In a few cases various other relationships between these two bodies
were observed, and several of these are illustrated (pl. 11, figs. 45, 47; pl. 13, figs.
96, 97). In these exceptional cells, however, the definition of the two bodies was
somewhat obscure.
Methyl Violet and Flemming’s Triple Stain.—]ohansen's methyl violet stain-
ing method was used without appreciable change. Methods developed by Smith
(34), Newton (Johansen, 40), and Hancock (742) did not give good results
but would probably prove satisfactory if time were taken to modify them.
The parvicorp generally stained in much the same manner as with Feulgen
(pl. 12, figs. 48-60), but sometimes only part of it retained the stain, the slides
thus resembling certain Heidenhain’s hematoxylin slides (pl. 12, figs. 51, 53, 56).
The magnicorp showed no particles or stained structures. Similar but less
certain results were obtained with Flemming’s triple stain (pl. 12, figs. 61, 62).
As these staining procedures provided little additional information to that ac-
quired from other techniques and as they were more troublesome to carry out,
they were not used extensively.
Heidenhain’s Hematoxylin and Brazilin.—After long mordanting and staining
with Heidenhain's hematoxylin (5a, Table III) the entire parvicorp remained black
(рі. 13, figs. 72-81). With the short staining procedure (5b, Table III) usually
a black area designated here as the companion body (text-fig. 1) stained at one
place and the remainder of the parvicorp was lighter than the surrounding cyto-
1946]
NAGEL—CYTOLOGICAL STUDY OF YEAST 257
plasm (pl. 13, figs. 82-89). This differentiation resembled that shown in Wager
and Peniston’s figures of the "chromatin patch” except that the “peripheral layer
of chromatin” was not generally apparent, nor did the companion body exhibit as
great a diversity of shape as the authors illustrate (Wager and Peniston, 710, pl.
16, figs. 53-80). This same phenomenon shows to some extent with Brazilin
and methyl violet techniques but is not apparent with any of the others that were
used, although identical fixations were frequently employed.
In either budding or resting cells this companion body was often double and
occurred in any one of several positions, probably due to both the orientation
under the microscope and the condition and stage of development of the cell.
Division of the companion body at mitosis is difficult to follow even in slides of
actively budding cultures. It seems to elongate greatly and divide by thinning
іп the center (pl. 13, figs. 83, 89). In the non-budding cell the parvicorp ap-
pears to have a very regular outline; in the budding cell it is not distinct as the
contrast between cytoplasm and parvicorp is not great. With the longer staining
procedure results are similar to those of the Feulgen and Giemsa techniques, but
no differentiation within the parvicorp is visible, the boundary is usually regular,
and no extrusions or appendages are apparent. The parvicorp may destain in
various unusual patterns, especially in old or poorly nourished cells, and give rise
to figures such as pl. 13, fig. 76, which, although they may resemble conventional
mitotic stages, probably do not represent a division stage at all. Because the
cytoplasm sometimes destains irregularly, results are difficult to interpret. There-
fore, in spite of the recent work of Subramaniam and Ranganathan (745), the
author does not consider this technique one of the most satisfactory for the study
of yeast cytology, certainly not the technique to use as the sole basis of interpre-
tation.
The magnicorp appears clear and optically empty in budding material. How-
ever, if it is shrunken or distorted in fixation, as sometimes happens, the folds
tend to retain the stain and give the appearance of strands passing over the sur-
face (pl. 13, fig. 73). In well-expanded and preserved magnicorps this phenom-
enon was not apparent. Wager and Peniston reported a “chromatin network”
over the surface of the magnicorp as a regular cell component, possibly because
they regularly dried their preparations after fixation. There are times when denser
strands of cytoplasm seem to radiate from the parvicorp and may or may
not extend out over the magnicorp depending upon the relative positions of the
two cell bodies (pl. 13, figs. 77, 78). The early entrance of the magnicorp into
the bud is especially apparent with this stain when it follows mercuric chloride
fixation. Subramaniam and Ranganathan have used a Carnoy fixing solution
which usually leaves the magnicorp invisible and the relative positions of cell
components uncertain.
Brazilin gives results similar to Heidenhain’s hematoxylin, staining either the
entire parvicorp or only the companion body depending upon the length of stain-
ing and destaining (pl. 12, figs. 63-71). With the shorter procedure the parvi-
— -——
[Vor. 33
258 : ANNALS OF THE MISSOURI BOTANICAL GARDEN
corp sometimes shows shadowy material extending from the companion body
(pl. 12, fig. 65).
Methylene Blue-Eosin Tecbnique.—Yhe Giemsa-eosin technique of Badian
(37) was modified by substituting methylene blue for Giemsa. It can be said
of it also: "L'application de cette méthode de différenciation aux Levures n'est pas
toujours ҒасПе--” [Badian, '37, p. 64]. Staining and differentiation of the cells
in bulk in a centrifuge tube were more easily controlled than on the cover glasses
although both methods were used.
Certain particles or granules in the yeast cell stained a brilliant blue-red when
the differentiation was carried just far enough; further differentiation in eosin left
the entire cell a faint, even pink. The position, number, and size of these particles
apparently depended in part on physiological conditions, in part on fixation. In
a 48-hour unshaken broth culture the stained bodies, if present, were found within
the magnicorp (pl. 14, figs. 98-107); in a similar culture that was shaken and
crowded colored particles appeared to lie at the periphery of the magnicorp or
occasionally in the cytoplasm (pl. 14, figs. 109—112). In the unshaken culture,
the parvicorp stained faint pink in the blue cytoplasm and was clearly visible in
98 per cent of the cells, but little or no differentiation was noted. The outline
was quite regular and clear. In this same culture approximately 85 per cent of
the yeast cells contained no particles at all in the magnicorp; the rest contained
one or more of various sizes and shapes (Table IV). In the shaken culture the
TABLE IV
NUMBER OF PARTICLES PER CELL IN THE MAGNICORP OF YEAST CELLS STAINED
WITH METHYLENE BLUE-EOSIN TECHNIQUE (100 CELLS SELECTED AT RANDOM).
Number of particles | | | | | | | |
сей 0 | 1 2 IN E. 8 9 10 or
| E rn more
| | | | | | | |
Number of cells | 85 | 4 | 2 | 2 | 2 | -- | 1 | -- 1 2
particles were exceedingly numerous and were present in most cells where differ-
entiation was not carried too far. The parvicorp was not visible in these cells.
Granule number per cell was counted in 100 granule-containing cells (15
per cent of the total cell number). The results are diagrammed in text-fig. 2.
Most of the cells in this culture contained few particles which at times appeared
irregular in outline as if they were possibly aggregates, but the total amount of
material in the magnicorps was not uniform.
In a few instances the particles appeared paired (Table V). Тһе pairs of
granules were similar in staining reaction, size, and shape. At times they occurred
as separate particles; at times the two were joined in dumbbell (“diplokokken
ähnlich”, Henneberg, 716) or V-shaped masses with the two ends alike in shape
and size. Rarely all of the particles of a cell appeared paired (pl. 14, fig. 107);
1946) 5
NAGEL—CYTOLOGICAL STUDY OF YEAST 259
more often there were only one or two such pairs, the other particles not showing
this condition. The number of pairs of granules was not constant. Due to the
position and arrangement of granules in some cells it was not always possible to
be certain whether or not they were paired. These are listed in Table V as
“Possibly paired.”
RO
МУ
40:
50:
Number of cells
1-2 3-4 5-6 78 9-0 11:12
Number of granules
Text-fig. 2. Number of granules per cell in the magnicorps of 100 granule-containing cells.
TABLE V
PER CENT OF YEAST CELLS SHOWING PAIRING OF THE En IN THE
MAGNICORP WHEN STAINED WITH METHYLENE BLUE
O6 of gran p of total —
containing cells f cells іп cultur
No apparent pairing 82.+ 12.4
One or more pairs present 9.— d.
Possibly paired 9. 1.4-
[Vor. 33
260 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Actively budding yeast from a yield test (4 hour 20 minute shaken culture)
showed very faintly differentiated parvicorps and a few cells which contained as
many as thirty to fifty stained particles. The latter were in or at the periphery
of the magnicorp (pl. 14, fig. 108) and in several instances were passing into the
bud. Due to the large number of somewhat similar granules in the cells pairing
could not be determined.
Aceto-Carmine and Related Smear Tecbniques.—As mentioned in Table Ш,
various concentrations and mixtures of acetic acid, lactic acid, and propionic acid
were tried with the following dyes used singly and in combination: carmine,
orcein, and Bismark brown. Results were generally unsatisfactory. Granular
cytoplasmic material stained so deeply as to obscure the parvicorp in most cases
and the magnicorp was generally shrunken and distorted. Various fixations be-
side acetic-alcohol and a number of other treatments and mordants suggested in
the literature (Darlington and La Cour, '42) were tried with little success. Aceto-
orcein and propionic carmine were somewhat more satisfactory than other solu-
tions. Gentle heating helped bring out contrast, and the slides improved a little
after standing for two to five days at room temperature. The cells which stained
most clearly were those from the edge of giant colonies two to four weeks old,
grown on nutrient agar in a moist chamber (pl. 14, figs. 113-119). Probably this
is due to the relatively small amount of granular material in the cytoplasm of
these cells (author's research, unpublished). The parvicorp may be rounded, or,
more frequently, somewhat star-shaped, or may occasionally appear bipartite,
especially in dividing cells. Тһе magnicorp (if visible at all) is usually shrunken,
distorted, and empty. Although much time was spent in trying to adapt this
technique to yeast, it was finally abandoned as unsatisfactory.
STUDIES OF SPORULATING CULTURES
Feulgen and Giemsa Techniques.—Because of the similarity of results with
these techniques, they are considered together. In any sporulating culture only
a few cells were in meiotic division at any one time, and the extremely small num-
ber found in certain stages would indicate that at least parts of the process are
very rapid. Preceding spore formation, the parvicorp becomes approximately
twice its normal diameter, appears filamentous, and resembles the prophase of the
conventional meiotic division as closely as could be expected in anything as small
and unique as a yeast cell (pl. 14, figs. 123-127; pl. 15, figs. 128-130, 141-144).
This phase appears to be of long duration as it is quite common in slides of sporu-
lating cultures. The next steps in the division are not too certain and probably
proceed with great rapidity as few instances were found that could be definitely
assigned to this phase in the many slides that were examined. Probably such
stages as pl. 15, figs. 131, 145, 146, 149, 150 belong in this category. Very
1946]
NAGEL—CYTOLOGICAL STUDY OF YEAST 261
rarely two parvicorps lying beside or above each other were found toward the
center of the cell (pl. 15, fig. 151; pl. 16, figs. 173, 174). While this position
might be the logical expectation, its extreme rarity leads one to believe that the
second division normally follows the first so rapidly that the stage with two
parvicorps may not actually become organized in most cells. The second division
appeared to start near the center of the cell. Two elongate, slightly dumbbell-
shaped masses were formed, the ends of which passed toward the periphery of the
cell, usually toward opposite poles. А gradual thickening of the extremities of
the masses occurred at the expense of the center (pl. 15, figs. 132—136; pl. 16,
figs. 152-158). Such division figures were relatively common. Cells with elon-
gate parvicorps which appeared bipartite or possibly four-partite were also rela-
tively common in all sporulating material (pl. 15, figs. 147, 148). These may
represent a stage which either follows the more diffuse early stage or might be the
beginning of the second division. Instances of a single parvicorp at each pole of
the cell were not positively ascertained; careful study of apparent examples usually
showed that at least at one pole one body could be resolved above the other, and
that the second division had actually taken place. When division is first com-
pleted, the parvicorps usually lie at the periphery of the cell close to the wall and
the spore plasm lying between them is frequently more dense than the epiplasm!
(pl. 16, fig. 160). The organization of the ascospores seems to occur in a manner
similar to that of other Ascomycetes. Apparently the spore is delimited by being
cut out by rays extending from the parvicorp. At times one or two of the asco-
spores develop more rapidly than the others. Ray-like strands of cytoplasm
frequently extend out from the parvicorp in the fully formed ascospore. If only
two or three ascospores develop, the other parvicorps can be observed lying free
in the ascus (pl. 15, figs. 139, 140). Single spores with two or more parvicorps
were not seen. Old agar slants of diploid cells, some sporulating, showed aberrant
divisions in which the parvicorp apparently divided without subsequent cell
division. In some of these cells the parvicorp was divided into separate particles
(pl. 13, fig. 97).
Meiotic division of the magnicorp could not be followed in either technique.
Although this body is generally visible in budding material stained with Feulgen,
it was imperfectly visible, if at all, in sporulating cells (except in the fully
developed ascospores where it again appeared empty).
Notes On Other Tecbniques.—Aceto-orcein was unsuccessful as a stain for
the division figures of spore formation because of the deeply staining, granular
cytoplasm. However, the formation of the ascospores was similar to that observed
with Giemsa and in the fully developed ascospores an empty magnicorp was gen-
erally visible (pl. 17, figs. 193, 194).
lIn Ascomycetes, the cytoplasmic contents of the ascus not used in spore formation.
[Vor. 33
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Methylene blue-eosin was not especially useful as a stain for sporulating
material, nor did the conventional volutin reaction give an enlightening picture
of the magnicorp and its particles in meiosis. With both techniques, if magni-
corps were visible, stained particles in them were few (exceptionally there were
many particles or none). Мо regular organization of magnicorp and its contents
was ascertained.
Slides stained by the longer procedure for Heidenhain's hematoxylin (5a,
Table III) or by Brazilin gave much the same results as Giemsa and Feulgen, but
less distinct due to the retention of stain by the granular cytoplasm (pl. 16, figs.
165—175). If stained by the shorter method (5b, Table III), the results were so
variable as to be uninterpretable and the companion body could not be followed.
The magnicorp, though evident in budding and resting cells with these tech-
niques, could not be followed in sporulating material.
STUDIES OF LIVING AND SUPRA-VITALLY STAINED CULTURES
Comparative studies on living cells of cultures were made whenever slides
were prepared. In budding cells the parvicorp was only rarely visible in living
cells, although its position could often be determined by a slight invagination of
the magnicorp. The latter body could almost always be seen except in certain
small, very dense cells where it was probably obscured by the highly refractive
surrounding cytoplasm. It occurred usually as one body, occasionally as two,
more rarely as several to many, depending on the age, nutrition, aeration, and
other cultural conditions of the cells. At times one to several particles could be
seen moving in the magnicorp, and occasionally these appeared to be paired.
The cytoplasm of many living cells from presporulating cultures was so highly
granular that all other structures were obscured. It was from these that spores
developed. Although a number of methods of observing the formation of spores
in living cells was tried, only the one described below was found satisfactory.
Nothing externally visible occurred in the living cell the first twelve to sixteen
hours on the gypsum slant, the variation in time depending upon the culture,
temperature, presporulating medium, etc. After the culture had been on the
gypsum slant for twelve hours samples were examined at half-hour intervals. The
movements of yeast cells are sufficiently rapid that for accurate continued ob-
servation it was found imperative to restrain them as much as possible. The fol-
lowing technique prevented excessive motion, yet kept the yeast in viable condition
under the microscope for five to seven hours. A tiny frayed-out fragment of lens
paper was placed on a No. 1 cover glass and both were dipped in alcohol and
flamed. With a little practice the lens paper fibers spread flat and evenly and are
not charred. A drop of sterile water acidified to about pH 4 with acetic acid
was placed on the lens paper and a few cells from the gypsum slant were added
with a needle. The cover glass was placed cell-side down over a concave (drop)
slide to provide an air chamber and the edges were sealed with wax. The
19461
NAGEL—CYTOLOGICAL STUDY OF YEAST 263
yeast cells gathered along the lens paper fibers, and Brownian movement
was thus greatly reduced. Where the cells were not crowded, an observation
every fifteen to twenty minutes kept a given cell or group of cells in view. Thus
individual cells could be watched from the first sign of spore delimitation to com-
plete spore formation, a process that took three to five hours at room temperature
under these conditions. In acidified water with no air space, in agar, or in oil,
sporulation also occurred but only sporadically and after one to five days.
In living cells the spores appeared to be delimited in the same manner as
described for prepared slides. Several instances were observed in which the four
spores were formed from the protoplasm at one end of a large cell. The other end
was occupied by a body which resembled the magnicorps of other cells on the
slide although no proof of its actual identity therewith was ascertained. A cell
of this kind from а Giemsa-stained slide is illustrated in pl. 17, fig. 195. Gen-
erally the magnicorp was not visible in the living cells during the process of
sporulation.
When iodine was applied to budding cells, it had the effect of temporarily
bringing the parvicorp into "relief" and of emphasizing the particles іп the magni-
corp, often revealing them in rapid Brownian motion.
Toluidine blue was also applied to budding cultures. This dye at times caused
„197919590
К
Bud dead in
Q (2)
SERIES 2 а о А d e
Text-fig. 3. Series of sketches of two cells under the influence of toluidine blue solution.
[Vor. 33
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN
very striking reactions in the magnicorp, but the parvicorp was never visible.
Changes in the magnicorp were especiall apparent with continuous observation
of a single cell. Text-fig. 3, Series 1, shows sketches of the changes in the magni-
corp of a budding cell observed continuously for forty-five minutes. In “h” and
"n" the magnicorp was seen to burst rapidly out into the bud. Series 2 shows a
similar non-budding cell. Granules of various sizes often stain in the magnicorp
with this procedure; occasionally also parts of the cytoplasm stain, and the cells
finally die and become deeply colored throughout.
WILD YEASTS
Studies of Budding Cultures.—Several yeasts were collected in the wild and a
few cytological observations were made on two of them. А brief account is
given here of these fragmentary observations since they are rather suggestive when
compared with the results with Saccharomyces cerevisiae. Had these yeasts been
obtained earlier in the progress of the investigation more detailed observations on
them would have been made.
The yeast represented in pl. 17 is one of a number which was isolated from
wild fruit gathered late in the fall of 1945 at the Missouri Botanical Garden
Arboretum near Gray Summit, Missouri. While it has not been positively identi-
fied, since it has not yet been induced to sporulate, it can be placed tentatively in
the Fungi Imperfecti, possibly in Syringospora or Blastodendrion or a closely ге-
lated genus and probably represents an undescribed species. It is an especially
large-celled yeast that produces from two to twelve easily detached buds in bi-
polar position when growing actively on nutrient agar. А few Giemsa and
Feulgen slides were made in order to study its cell organization when producing
the higher number of buds.
From one to twelve parvicorps (Feulgen and Giemsa positive) were found
per cell, the number depending on their size and the amount of budding. Com-
monly there were more parvicorps than buds. Ав was noted in the microscopic
study of the living cells, the buds were very delicately attached and some of them
may have been broken off in handling. Probably also the division precedes the bud
formation in this species. While no attempt was made to check this fact, the very
large number of parvicorps in certain cells (pl. 17, figs. 176, 177, 186) would
suggest such a possibility. However, small, detached, immature buds without
parvicorps were frequently observed. When numerous parvicorps were present
they were often of two distinct sizes, the smaller ones staining more intensely.
In contrast to Saccharomyces cerevisiae the parvicorp is often separated into
particulate units at division (pl. 17, figs. 178, 180, 183, 187, 189). When several
parvicorps were present all seemed to divide synchronously, as cells were noted
with two, four, or eight in division at the same time. Dividing and non-
dividing parvicorps were not observed in the same cell. Wherever the magnicorp
1946]
NAGEL—CYTOLOGICAL STUDY OF YEAST 265
was visible it was empty, but when many parvicorps were present they sometimes
seemed to fill the whole cell.
Another of the wild yeasts from the same collection (a Cryptococcus?) should
be mentioned because under certain cultural conditions (usually when several days
to a week old) moving particles in the magnicorp were the rule rather than the
exception and many of the particles seemed paired in dumbbell fashion. In some
sections of one particular slide almost half the magnicorps had two equal particles,
closely paired, each pair moving as a unit. Under other conditions, especially in
young or older cultures, this phenomenon was not apparent.
DISCUSSION
The very nature of vegetative reproduction in budding yeasts would lead one
to expect their mitoses and nuclear organization to exhibit certain exceptional
phenomena. As there is nothing quite comparable to this budding process in the
plant or animal kingdom, interpretation of nuclear organization should take into
account as inclusive a picture of all cell components as possible. Both parvicorp
and magnicorp seem to be associated with vegetative cell division and a complete
interpretation must take cognizance of both. From genetics there is now exact
indirect evidence on the organization of the yeast cell. The excellent work of
Lindegren, Spiegelman, and Lindegren (744) and of Winge (739, '44) has demon-
strated the regular Mendelian segregation of characters at meiosis. Diploid strains,
reproducing only vegetatively, are stable and retain their cultural characteristics
as would be expected; haploid cultures exhibit a far greater mutation rate. This
is easily observed in giant colony structures where haploids show frequent sector
mutations which are generally lacking in the more stable diploids. Therefore in
spite of its small size and peculiar vegetative reproduction yeast cannot be greatly
different in its fundamental organization from other organisms although in various
superficial details of mitosis and meiosis one may confidently expect considerable
modification.
Similar superficial modifications of mitosis and meiosis have been reported for
a variety of tissues and organisms. In the pollen tube of the spermatophytes,
when mitosis of the generative cell must occur within narrow confines, it is often
somewhat atypical; the metaphase plate may be greatly elongated and there may
be no visible achromatic figure (Trankowski, '30). In the Protista, Belar ('26,
'28) has shown that fundamentally regular but superficially atypical nuclear or-
ganization is not uncommon. As mentioned before, several conflicting interpreta-
tions of nuclear organization of yeast have been presented in the past and there is
to date no general agreement. Table VI presents in tabular form the varying in-
terpretations by previous authors of the disputed entities of the yeast cell.
In the light of the work of Магла and Jaeger (°39), Caspersson and Schultz
(38), Caspersson ('39a, '39b, '40), Mirsky (743), Mirsky and Pollister ('43a,
'43b), Pollister and Mirsky (743, '44), Greenstein (’44), Davidson and Way-
[Vor. 33
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mouth ('44), and many others, the Feulgen positive staining of the parvicorp
would seem indicative of its nuclear nature. Тһе constancy of the parvicorp as
a cell component and its non-homogeneous character when stained with Giemsa
supports this interpretation. Opponents to this view point out that the division
of the parvicorp appears to be amitotic. Since division is intra-nuclear in many
fungi, this fact plus the very small size of the parvicorp make resolution of indi-
vidual chromosomes difficult with the ordinary microscope. Іп the wild yeast
described above particulate units were observed at certain division stages in both
Feulgen and Giemsa slides. Іп the early stages of meiotic division in S. cerevisiae
the parvicorp becomes enlarged and appears filamentous, resembling the usual
early meiotic prophase nucleus, but its small size makes accurate resolution diffi-
cult. As is true of many nuclei, the parvicorp stains more intensely with Feulgen
at some stages than it does at others. It is visible in all stages of division in all
cells stained with Feulgen and Giemsa and with Heidenhain's hematoxylin and
Brazilin if destaining is not carried too far, but is rarely visible in the living cells.
In ultra-violet photographs (Caspersson and Brandt, '41; Brandt, '41) the two
types of nucleic acid are indistinguishable. "These authors state that this technique
does not generally differentiate the nucleus in yeast. In the very actively budding
cells, however, the parvicorps appear in the photographs as diffuse, somewhat
lighter areas in the deeply absorbing cytoplasm, sometimes seeming not much more
absorbent than the magnicorp which they state contains very little nucleic acid.
The magnicorp enters the bud before the parvicorp and is probably of im-
portance in bud initiation as suggested by Lindegren ('45). If particles are visible
within the magnicorp they also enter the bud, and at times they seem to be paired
or assume form or position difficult to explain as "reserve stuff." "These phenomena
plus the apparent amitosis of the parvicorp have led to the interpretation (Jans-
sens and Leblanc, '98; Wager and Peniston, '10; Lindegren, '45) of the magnicorp
and its contents as the nuclear vacuole and chromatin respectively. Although the
magnicorp and its contents are Feulgen negative at all stages of growth and
division, Lindegren (745) has pointed out that the desoxyribosenucleoprotein
nature (Feulgen positive) of nuclei of higher organisms does not necessarily indi-
cate its universality in the chromatin of simpler organisms. However, if the
magnicorp is interpreted as the nuclear vacuole and the parvicorp as the centriole,
the latter is not usually Feulgen positive. If visible, the particles in the magnicorp
vary greatly as to size, shape, and number: (a) with physiological conditions—
age, nutrition, aeration, etc.; (b) with the strain; and (c) with fixation. Their
chemical composition is not known with certainty, but common nuclear fixing
solutions are not generally effective, and special fixation, as, for example, with
formaldehyde, must usually be employed. As pointed out earlier, the magnicorp
is usually a single body, but often seems to occur as two bodies, one at either end
of the parvicorp. In old or starved cultures it may appear to be divided into a
number of parts (Brandt, '41), in which case the particles may occur in any or
19461
NAGEL—CYTOLOGICAL STUDY OF YEAST 267
all the parts of the magnicorp. Budding cells very commonly show no stained
particles with the techniques employed. The number of particles counted in
cells of different cultures stained with methylene blue-eosin varied from none in
most cells to as high as fifty. Van Herwerden (718) found the number of
particles related to phosphorus content of the nutrient medium. With dark field
illumination the rate of Brownian motion of particles in the magnicorp and in the
surrounding medium is similar, indicating a very low viscosity. On the contrary,
the lack of Brownian motion in the vicinity of the parvicorp and its tendency to
bulge into the magnicorp would indicate a higher viscosity for the former. The
pressure changes of the magnicorp with toluidine blue are likewise indicative of
low viscosity, a character more usually associated with vacuolar behavior than
with nuclear "sap." Difficulty in following any organization of the magnicorp
through meiotic division in this investigation (also by Janssens and Leblanc, 798;
and Wager and Peniston, 710) points strongly away from a nuclear interpretation
but improved techniques may alter this. However, the presence of the magnicorp
in the fully developed spore indicates its importance as a cell entity.
Various chromosome numbers in various parts of the cell have been suggested:
two—both haploid and diploid—by Badian (738); approximately eight—diploid
—by Kater ('27) in the parvicorp; and twelve by Lindegren (745) in the magni-
corp. Although a number of investigators have noted the division of the parvi-
corp into two bodies at times and have called these chromosomes, present genetic
evidence (Lindegren, personal communication, unpublished) does not point to
this chromosome number, and cytological evidence is as yet uncertain. Undoubt-
edly the parvicorp is divided into two bodies at times, but if these are chromosomes
they are large enough that they should be seen occasionally in metaphase or
anaphase configuration. If the parvicorp is nuclear, further refinement of tech-
nique is necessary to be certain of chromosome number. If the magnicorp is
nuclear, the variability of the number of particulate units is suspiciously great.
However, because of the small size of the yeast cell and its lability under various
cultural conditions, this objection has less force than it would have in other
material. As mentioned previously, the parvicorp of the wild yeast described
above appeared to separate into particulate units during mitosis but these were
too crowded together for the number to be certain.
At least four parts of the yeast cell have been interpreted as the nucleolus
(Table VI). Wager and Peniston (710) considered the parvicorp as the nucleolus
with the chromatin network extending from it over the magnicorp. This places
the “nucleolus” entirely outside of the “nucleus” as they interpreted it, certainly
an unusual position for it. Brandt (741) and Caspersson and Brandt (741) con-
sider the ribonucleic acid-containing granules scattered throughout the cytoplasm
(designated as "volutin" by them) the equivalent of the heterochromatin and
nucleolus of higher organisms. Janssens and Leblanc (798) sometimes called
the central granule of the magnicorp the nucleolus, and Guilliermond (’20) and
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
268
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NAGEL—CYTOLOGICAL STUDY OF YEAST
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[Vor. 33
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Kater (27) thus designated the deeper-staining body or bodies in the parvicorp.
At times one or more such bodies are visible in the Giemsa-stained slides, but
whether these are sufficiently consistent to be considered the nucleoli is questionable.
Whether or not a membrane surrounds the parvicorp has been a question of
debate. Its presence is indicated by the regularity of outline of the parvicorp
after certain treatments, for example, with methylene blue-eosin and with Heiden-
hain's hematoxylin (short procedure; pl. 13, figs. 82-88; pl. 14, figs. 98-108).
It is true that the parvicorps in Giemsa and Feulgen slides often show an irregular
outline. Acid hydrolysis has been carried out with both of these, and although
meee б 52) n жаши that this treatment does not alter the structure of
the containing material, there is no reason to believe
that a нЕ, if picot, would stain. Studies of the magnicorp indicate that
there is a membrane separating it from the cytoplasm. At times it appears to be
a double membrane for occasionally after fixation the magnicorp seems to shrink
away from the cytoplasm in places as if both possessed membranes.
If the nucleo/cytoplasmic ratio is considered, the parvicorp alone gives a ratio
which is possibly slightly low, the combined parvicorp and magnicorp an excep-
tionally high one.
It is thus apparent that after one hundred years of cytological work, the or-
ganization of the yeast nucleus is still a matter for debate among authorities, even
as to the most elementary points. Direct observation, uncorrelated with other
techniques, must probably continue to be relatively unproductive with objects as
microscopically refractory as the yeast cell. However, direct cytological observa-
tion in combination with other techniques presents a more encouraging picture.
Among the most promising of these developments are the following:
(1) Additional genetic studies such as those of Lindegren and Winge.
Eventually they should demonstrate the chromosome numbers, their comparative
size, and even chiasma frequencies and positions.
(2) Further biochemical studies along the lines of those of Mirsky and
Pollister on higher organisms, combined with studies of the effects of enzymatic
action on the several cell entities. A more complete knowledge of the chemical
nature of the particles in the magnicorp and of the refractive granules in the
cytoplasm should aid in an understanding of their role in cell organization and
metabolism.
(3) Comparative cytological studies of wild yeasts. Species with larger
cells or clearer cytoplasm should reveal more details of cell structure. Wild yeasts
which reproduce by budding but which also develop true or pseudo-mycelia are
frequently encountered. A cytological study of these species would relate the
recognized entities of the yeast cell to the more usual mycelial type of growth.
Additional investigations with ultra-violet and "phase-difference" micro-
scopes and the study of thin sections with the electron microscope.
(5) Further refinements of cytological techniques. A method of removing,
softening, or breaking the firm cell wall (possibly by enzymatic digestion, chem-
19461 |
NAGEL—CYTOLOGICAL STUDY OF YEAST 271
ical maceration, or pressure) which would permit spreading of the cell contents
without excessive distortion should make further details of the parvi- and magni-
corp resolvable. A study of the effects of cold treatment on different stages of
spore formation may make it possible to obtain many more cells at one time in
certain phases of meiosis. New techniques, as well as improvements in present
fixation and staining procedures, should provide additional information for cor-
relation with other methods.
SUMMARY
1. A review of the literature on the cytology of yeasts showed great con-
fusion in:
(1) use of terms,
(2) interpretation of nuclear structure.
(This is demonstrated in tabular form in Table VI.) An attempt was made to
apply systematically a whole battery of techniques to both sporulating and bud-
ding material of yeast (S. cerevisiae).
2. All of the techniques tried out are listed and described. Those used ex-
tensively are discussed in detail.
3. А descriptive terminology for the major cell entities (parvicorp, magni-
corp, companion body, text-fig. 1, p. 254) is adopted in order to permit
presentation of observations unbiased by earlier interpretations.
Fragmentary observations on two wild yeasts are included for comparison
(рр. 264-265).
5. Тһе “рагуісогр” is a Feulgen positive, non-homogeneous, constant cell
entity. Vegetative division does not appear to be typically mitotic in S.
cerevisiae, but at least the prophase of meiosis resembles conventional configura-
tions. А large-celled wild yeast showed particulate units of the parvicorp at
mitosis.
6. The "magnicorp" (vacuole or nuclear vacuole of most authors) is Feulgen
negative. It is almost universally present in budding material and enters the bud
before the parvicorp, but is generally invisible in presporulating and sporulating
material. Granules of the magnicorp are lacking in most cells with the techniques
БЕРІШ, but, when present, occasionally appear paired.
Relationships of these observations to the several interpretations of nuclear
organization are discussed briefly (pp. 265-271), and suggestions are made as to
possible future advances in this field.
[Vor. 33
272 ANNALS OF THE MISSOURI BOTANICAL GARDEN
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211 39). Saccharomycodes Ludwigii Hansen, a balanced зача 0 Ibid.
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[Vor. 33, 1946)
274 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 10
Cells from budding culture of Saccharomyces cerevisiae (Feulgen technique).
Figs. 1-19. Shaken broth culture, diploid (LK) strain, mercuric chloride fixation.
ANN. Мо. Bor. GARD., Vor. 33, 1946 PLATE 10
Budding material— Feulgen
NAGEL—SACCHAROMYCES CEREVISIAE
[Vor. 33, 1946]
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 11
Cells from budding cultures е E cerevisiae (Robinow's Giemsa technique) —all
12—24-hr. broth cultures except fig.
Figs. 20-41, 43. Haploid (Вс20--) strain.
Figs. 42, 44, 45, 47. Diploid strains.
тч cell from sporulating culture 24 hrs. on gypsum slant, diploid
intra- "specific hybri
Figs. 20-34, 38-42, 44, 45, 47. Todine-glacial acetic acid-formalin fixation.
Figs. 35-37, 43, 46. Mercuric chloride fixation.
ANN. Mo. Вот. Garp., Vor. 33, 1946 PLATE 11
Budding material — Giemsa
NAGEL—SACCHAROMYCES CEREVISIAE
[Vor. 33, 1946]
278 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 12
Cells from кн cultures of S. cerevisiae (methyl violet, Flemming’s triple, ог
веша technique).
Figs. 48-60. Johansen’s methyl 08 I broth culture, diploid (LK) strain,
Navashin fixation (figures not drawn to s
Figs. 61, 62. Flemming's triple stain, otherwise as above.
Fig ‚ 71. Brazilin (with short еуро на non-sporulating cells from
gypsum Лы, diploid (LK) strain, mercuric chloride fixatio
—67, 69, 70. Brazilin (with long ошо), broth culture from yield test,
"TE intra-specific hybrid, mercuric chloride fixatio
ANN. Мо. Вот. Garp., Vor. 33, 1946 PLATE 12
Flemmings triple
Brazilin
Budding material.
NAGEL—SACCHAROMYCES CEREVISIAE
[Vor. 33, 1946]
280 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ExPLANATION OF PLATE
PLATE 13
Cells from budding culture of S. cerevisiae (Heidenhain's hematoxylin) and from
agar slant (Robinow's Giemsa technique).
Figs. Heidenhain's hematoxylin stain Pe m II), 24-hr., unshaken
broth culture, diploid (LK) strain, mercuric chloride fixatio
Figs. 82-89. Heidenhain’s hematoxylin stain (5b, Table III), otherwise as above.
Figs. 90-97. Robinow’s Giemsa stain, 7-week-old nutrient agar slant, diploid (LK)
strain, osmic acid vapor fixation
ANN. Мо. Вот. Garb., Vor. 33, 1946 PLATE 13
24285
Budding, material Hematoxylin
~ ви
94
К 95
Giemsa
Op “м
NAGEL—SACCHAROMYCES CEREVISIAE
ANN. Мо. Вот. Garp., VoL. 33, 1946 . . PLATE 13
NAGEL—SACCHAROMYCES CEREVISIAE
[Vor. 33, 1946]
282 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 14
Cells from budding cultures of S. cerevisiae (methylene blue-eosin or aceto-orcein
technique) and from sporulating cultures (Feulgen technique).
. 98-107. Methylene blue-eosin stain, 48-hr., unshaken broth culture, diploid
intra- саќе hybrid, fixation іп sat. picric acid іп 70 рег cent alcoho
Fig. 108. Methylene деді -eosin stain; 4-hr. shaken yield test, broth culture, dip-
loid, osmic acid vapor fixatio
Figs. 109-112. Same as figs. 98-107 except culture was shaken.
Figs. 113-117. Aceto-orcein stain, edge cells from giant colonies, various haploid
strains.
F Aceto-orcein stain, budding cells from sporulating culture 24 hrs.
on nina s diploid (EFF) strain
120, 122. Robinow’s Giemsa stain, week-old nutrient agar slant, diploid
(EFF) бів, osmic acid vapor fixation
. 121, 124-126. Vii stain, 48 hrs. on presporulating medium, diploid (LK)
strain, mercuric acid fixatio
Figs. 123, 127. Feulgen ұлан, кайи: culture 17 hrs. on gypsum slant, diploid
(LK) strain, mercuric acid fixatio
ANN. Мо. Bor. GARD., Vor. 33, 1946 PLATE 14
2 7109
Budding material
113
Aceto-orcein
124 129. 126 .
Sporulatmg material Feulgen
NAGEL—SACCHAROMYCES CEREVISIAE
ANN. Mo. Bor. Garp., Vor. 33, 1946 PLATE 15
%
х д
&
Giemsa
Sporulating ‘material
NAGEL—SACCHAROMYCES CEREVISIAE
| | [Vor. 33, 1946]
286 ANNALS OF THE MISSOURI BOTANICAL GARDEN :
EXPLANATION OF PLATE
PLATE 16
s from sporulating cultures of S. cerevisiae коо $ Giemsa, Heidenhain's
hematoxylin, or Brazilin techniq
ue).
А 152-1 164. Robinow's гае stain, sporulating culture 24 ке оп gypsum
x: d mercuric chloride fixation.
Heidenhain's — stain (5a, Table ied sporting c re
16 ы оп Sun diae diploid (LK) strain, fixation in 48 p
chloride, 2 per cent glacial acetic acid, 50 = "cent вя асі
Figs. 170-175. Brazilin stain, otherwise as above.
[Vor. 33, 1946]
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ExPLANATION OF PLATE
PLATE 15
Cells from sporulating cultures of S. cerevisiae (Feulgen or Robinow's Giemsa tech-
nique).
igs. 128-140. Feulgen stain, ini. culture 17 hrs. on gypsum slant, diploid
(LK) strain, mercuric chloride fixat
Figs. 141-151. Robinow's Giemsa stain, sporulating culture 24 hrs. on gypsum
slant, diploid, mercuric chloride fixation. Figs. 141, 142 are same cell shown at two
levels (өлік scale than other figures
ANN. Mo. Bor. GARD., Vor. 33, 1946 PLATE 16
173
Sporulating material
NAGEL—SACCHAROMYCES CEREVISIAE
[Vor. 33, 1946]
288 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 17
Budding cells of wild yeast (Robinow’s Giemsa or Feulgen technique) and sporu-
lating cells of 8. cerevisiae (aceto-orcein or Robinow’s Giemsa technique).
Figs. 176-190. Robinow’s Giemsa stain, 24-hr. agar slant, wild yeast, osmic acid
vapor fixation. In figs. 176, 177, 186 parvicorps are shown with solid outlines, magni-
corps with dotted outlines.
Figs. 191, 192. Feulgen stain, otherwise as above.
Figs. 193, 194.
E (EFF) strain.
Robinow's Giemsa stain, sporulating culture 24 hrs. on gypsum slant,
Aceto-orcein stain, sporulating culture 24 hrs. on gypsum slant
195.
mH mercuric chloride fixation
ANN. Мо. Вот. Garp., Vor. 33, 1946 PLATE 17
NAGEL—SACCHAROMYCES CEREVISIAE
CONTRIBUTIONS ТО OUR KNOWLEDGE OF AMERICAN
CARBONIFEROUS FLORAS
IX. SOME PETRIFIED SEEDS FROM IOWA
ELLEN M. KERN
AND HENRY N. ANDREWS
This is a third contribution that has accrued from our study of the coal-ball
collections of Mr. Frederick O. Thompson of Des Moines, Iowa, the two previous
accounts (Parts VII and VIII of "American Carboniferous Floras") giving a
detailed description of the origin of the specimens. All of the fossils described
herewith were found in coal balls from the Urbandale mine located immediately
west of Des Moines.
During the decades that have elapsed since the time of Brongniart’s (781)
foundational memoir on the silicified seeds from Saint-Etienne a rather vast as-
semblage of fossil seeds have been described from Paleozoic deposits. Although
relatively few have been found attached to the plants that bore them they have
contributed very appreciably to our knowledge of the three great groups of
Carboniferous seed-bearing plants—the Pteridospermeae, Cordaitales, and Lyco-
podiales. Comprehensive summaries of the many genera and species are included
in the works of Seward (717) and Arnold (738).
With the exception of Lepidocarpon, few structurally preserved seeds have
been recorded from the Carboniferous of North America, most of the known
species having been found in European deposits. There is now some evidence to
indicate that certain areas of the Pennsylvanian forests of Illinois were predomi-
nantly pteridophytic, and since much of the American coal-ball work to date has
been confined to collections from that state the apparent paucity of seeds is
accounted for at least in part. It should be added, however, that only a few of
the coal mines of even Illinois have been subjected to intense study, and it will
certainly be some years before we arrive at a clear picture of the composition of
the ancient forests that are represented in the coal balls. Іп a previous discussion
a brief contrast was drawn between the coal-ball flora of Illinois and that of
Iowa, seeds being present in Iowa in much greater numbers. Although there are
at least seven species of seeds, exclusive of Lepidocarpon, in our present collection
we have selected for description only the better-preserved ones.
Conostoma Williamson, 1877.
Conostoma oblongum Williamson.
Only one specimen of this species has turned up in our Urbandale collection
thus far, and since it has been described with considerable precision by Oliver and
Salisbury (711) from the Lower Coal Measures of Lancashire a detailed account is
not warranted here. More recently it has been reported by Krick (732) from the
(291)
[Vor. 33
292 ANNALS OF THE MISSOURI BOTANICAL GARDEN
upper part of the Carbondale group at Harrisburg, Illinois.
The Iowa record given here is based on a specimen that was exposed in longi-
tudinal section, although the cut had been made slightly beyond the median plane.
The seed (pl. 18, fig. 2) measures nearly 4.5 mm. long by 2 mm. in diameter and
shows no pronounced tapering toward either end. The apical portion of the
nucellus (fig. 3) is quite well preserved, showing the distinctive generic features
of the pollen chamber in which two pollen grains may be noted. Тһе integument
is lobed at the micropylar end, somewhat more distinctly so than in the previously
described specimens.
Conostoma oblongum is described by Oliver and Salisbury as being platy-
spermic although in their own words it is "only trifling in amount," and their
figures adequately bear this out. In view of the general wide range in cross-
sectional shape of the petrified Carboniferous seeds there can be little doubt that
more fundamental structural features, such as the organization of the pollen
chamber and the nature of the integument, are of greater significance. In other
words, Seward's classification of the Paleozoic seeds into three groups, the Lage-
nostomales, Trigonocarpales, and Cardiocarpales, is certainly preferable to the
older system based on cross-sectional shape.
Aside from Krick's citation of Conostoma oblongum in an Illinois coal ball,
the only previously known American species are those described by Graham (734).
His well-executed figures of C. platyspermum leave no doubt as to the generic
identity of this fossil, and although it is very close to C. oblongum its segregation
as a distinct species seems justified.
Rhabdospermum Seward, 1917.
Rhabdospermum spinatum Andrews, sp. nov.
The fossil seeds, preserved as both petrifactions and impressions, that are
of Cordaitean affinities already compose a striking assemblage displaying con-
siderable variation in size, form, and anatomy. The fact that the vast majority of
these seeds (included in the Cardiocarpales as classified by Seward, '17, pp. 332-
356) have been described from European Carboniferous deposits by no means
implies their restriction to that region. Cordaitean stem, root, and foliage remains
are of frequent occurrence in the Illinois and Iowa coal balls. Although the
present description is based on a single, incomplete seed, it seems worth recording
inasmuch as it presents characters that distinguish it from any previously described
species.
The seed is heart-shaped in longitudinal section and relatively large, measuring
15 mm. broad by 12 mm. long. The integument, which is composed of four
clearly defined tissues, is especially well preserved. Following the nomenclature
used by most previous workers, the outer two tissues will be referred to as the
sarcotesta and the inner, more sclerotic two, as the sclerotesta (text-figs. 1, 2).
The outer sarcotesta, which is appreciably thicker than the three inner zones
1946]
KERN & ANDREWS— AMERICAN CARBONIFEROUS FLORAS. IX 293
combined, is composed of large, relatively thin-walled cells averaging about 20044
in diameter (text-fig. 2, So). In life this must have been of a fleshy texture,
probably comparable with that of the outer coat of a Cycas circinalis seed. This
outer sarcotesta is bounded by an epidermis of much smaller, vertically elongated
cells averaging about 115и x 70g. Тһе cells of the inner sarcotesta (Si) аге
nearly isodiametric, like those of the outer tissue, although much smaller, aver-
aging 50и.
Text-fig. 1. Rbabdospermum spinatum. Diagram of the seed shown in median longi-
tudinal section, n um chamber restored: SO, outer sarcotesta; SI, inner sarcotesta;
SCO, outer sclerotest
The sclerotesta appears as a conspicuous dark brown band, the color being due
partly to the cellular contents. Like the sarcotesta, it is composed of two clearly
defined tissues (text-fig. 2, Sco, Sci), although relatively much thinner and not as
readily distinguishable at lower magnifications (pl. 18, fig. 1). Тһе outer com-
ponent (Sco) consists of cells similar in shape to those of the inner sarcotesta
All cell measurements of these i Wd qe tissues have been taken from the plane of the
ДЕ; median longitudinal section shown іп рі. 18, fig. 1.
[Vóor..38
294 ANNALS OF THE MISSOURI BOTANICAL GARDEN
c
ње
gne IER атуды
MES
d
" > er
е ж (^ 4 wen
о * ч 5 |.
‚4 7 7X4 de.
‘о ./ Apr У ray f
^ PNM $
b
Text-fig. 2
= Rhabdospermum
spinatum
Cellular detail taken
from the equatorial re-
gion: E, epidermis; So,
outer sarcotesta; Si, in-
ner sarcotesta; Sco,
outer sclerotesta; Sci,
inner sclerotesta; nm,
remnants of nucellus
and megaspore mem-
brane.
19461
KERN & ANDREWS—AMERICAN CARBONIFEROUS FLORAS. IX 295
although thicker-walled. The most conspicuous feature of this tissue is the ir-
regularly shaped spines projecting into the inner sarcotesta, and it is in recognition
of this character that the specific name has been chosen. The inner sclerotesta
consists of cells that are very poorly preserved but which apparently were elon-
gated in the plane of the longitudinal axis.
The remnants of the nucellus are clearly attached to a raised cushion at the
base of the seed although it otherwise appears to be quite free from the integu-
ment. It should be noted that, since the innermost tissues of the sclerotesta and
of the adjoining nucellar tissue are not perfectly preserved, it is not certain
whether the two were originally in organic connection. Yet the space between
them is so uniform as to imply that it is natural rather than due to shrinkage or
decay. .
Very little remains of the nucellar tissue. At its apex, however, it appears to
have developed in the form of a broad conical pollen chamber (pl. 18, fig. 1,
text-fig. 1). Although little more than the cuticularized remains of the epidermal
cells are left, the form of the integument in this region also suggests such a
terminal structure of the nucellus. Thus the internal conical cavity presents a
distinct contrast to the flattened one іп Rhabdospermum cyclocaryon (Ad.
Brongn.) Seward (Brongniart, 781, pl. XII, fig. 1).
Affinities: —The affinities of this seed appear to lie with the closely related
genera Cardiocarpus, Rhabdospermum and Mitrospermum of the Cardiocarpales.
The last two of these genera are distinguished from Cardiocarpus chiefly in the
way that the vascular strands originate from the main bundle. In Cardiocarpus
the outer (lower) bundles depart from the main vascular axis before the latter
reaches the sclerotesta, while in Rbabdospermum. and Mitrospermum they аге
given off from the sclerotesta region and recurve back into the outer portion of
the integument (cf. Seward, '17, figs. 500B and 501E).
As stated above, the only available specimen of this seed was exposed on a
previously cut slab from Mr. Thompson’s collection. The fact that the initial
cut was apparently made directly through the central vascular strand leading up
through the base of the seed makes it impossible to determine whether the integu-
ment branches departed from the main bundle before or after reaching the
sclerotesta. The one observable integumentary strand passes out between the outer
and inner sarcotesta, more closely comparable with the position of the bundles in
Rhabdospermum (Seward, 717, fig. 501E) than in Cardiocarpus (Seward, 717, fig.
500B). Moreover, the strikingly distinct integumentary tissues of Rhabdo-
spermum spinatum compare very closely with Rhabdospermum cyclocaryon (see
Brongniart, '81, pl. XII, fig. 3). Тһе spinose nature of the sclerotesta of the
new seed sets it apart from R. cyclocaryon or the apparently closely related
Mitrospermum compressum (Will.) A. Arber.
[Vor. 33
296 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Diagnosis:—R babdospermum spinatum: platyspermic seed 15 тт. broad
x 12 mm. long; integument composed of four tissues: an outer conspicuously
thick sarcotesta, inner sarcotesta, outer spinose sclerotesta, and inner sclerotesta of
longitudinally elongate cells; pollen chamber shaped like an inverted shallow
teacup.
Locality: Urbandale Coal Mine, Des Moines, Iowa.
Horizon: Des Moines Series, Pennsylvanian.
Type specimen: No. WCB493, Henry Shaw School of Botany paleobotanical
collections.
Kamaraspermum Leeanum Kern, gen. et sp. nov.
One of the Urbandale coal-balls received from Mr. Thompson contained a
dozen or more specimens of a seed which, because of its highly distinctive struc-
ture, is designated herewith as a new species, Kamaraspermum Leeanum'.
pim
Text-fig. 3. Kamaraspermum Leeanum.
Diagrammatic median longitudinal section along
the minor axis (from еей А, peel 475-Т 21).
Horizontal broken lines icate corresponding
positions of transverse sections prit seeds C and
| 74-0 D (see text-fig. 5). The seed tissues are indicated
| T12-C as follows:
ETIO Epidermis—outer black line.
Outer sclerotic integument—striped area.
Outer parenchymatous integument—white area.
Inner sclerotic integument—
Inner parenchymatous integument — white area
delimited by inner sclerotic integument and
T 26-C thin black line
Nucellus and megaspore membra
= 7530,2 Probable course of vascular Жі ee line
T 732-C through basal chamber.
Parenchymatous cells of basal chamber—black
| T25-D
rT 24-D
| T23-D
722-2
ШЕСІ So,
2The prefix Kamara is from the Greek, meaning а chamber or room 22 ап arched covering.
The species is named for Mr. Arthur Е. т а Engineer of the Binkley Coal Company’s
ramid Mine, Pinckneyville, Illinois. Mr. most cordial AE ды the past six
years has been an indispensable aid in our Ed ball collecting.
19461
KERN % ANDREWS—AMERICAN CARBONIFEROUS FLORAS. IX 297
In spite of some pyritization the seeds are quite well preserved, and the orienta-
tion of two of them has allowed the preparation of satisfactory series of transverse
and longitudinal peel preparations. The remaining specimens, although less com-
pletely preserved, have been useful in checking structures throughout.
The seed possesses an integument with a distinctive succession of tissues, and
a large basal chamber, presumably a buoyancy mechanism which aided in water
transport. The only previously described seeds with which it may be compared
in a general way are those assigned to Brongniart's Codonospermum. | However,
the pronounced platyspermy of the Iowa seeds, as well as the nature of the integu-
ment, seems to render necessary a new generic name.
GENERAL ORGANIZATION—
The seed has the approximate shape of a double convex lens (pl. 19, figs. 4, 5),
slightly elongated in the micropyle-peduncle axis, with an extended micropyle in
the form of a flattened funnel. It measures about 12 mm. in length, and in a
median transverse section the major and minor axes measure 11 and 3 mm. re-
spectively. Thus, quite different aspects are presented, depending upon whether
the longitudinal section is taken through the major (fig. 5) or minor (fig. 4)
axis. In order to portray clearly the various aspects of the seed two sets of
diagrammatic drawings have been prepared: one, from a series of transverse
sections (text-fig. 4); and the other from a series of longitudinal sections (text-
fig. 5)? taken through the minor axis.
It is significant to note that the seeds, with the exception of the terminal
portion of the micropyle, show no evidence of any appreciable distortion as a
result of fossilization. Aside from the outer layer of the nucellus and whatever
may have been within the megaspore membrane at the time the seeds were de-
posited, the tissues are well preserved and the gross shape as outlined in the text
figures presents the true life form of the seed.
A longitudinal section (pl. 19, fig. 4, and text-fig. 3) shows that the seed is
composed of two clearly defined regions: a basal chamber and nucellar chamber,
with their attendant tissues. Simply as a matter of convenience these will be
described separately.
NUCELLAR REGION—
Integument:—The integument is bounded externally by a single layer of epi-
dermal cells which appear brick-shaped in both longitudinal and transverse sec-
tions. Within this epidermis is a succession of four clearly defined tissues. First
Ея the seeds on which this description is based were contained іп опе к. зд coal-ball ed smt
s not М to рге 25 al preparations of peels. Consequently in the lon
t wa
еді series E in t-fig. 5 ор н are slightly Ы. каар the micropyle E
basal chamber more Louis at one в. than the other (cf. Nos. 2 and 3 pid Nos. 15 and 16).
[Vor. 33
298 ANNALS OF THE MISSOURI BOTANICAL GARDEN
NN АА WY N
NNN
xt-fig. 4. Kamaraspermum Leeanum. А series of diagrammatic drawings of transverse
sections extending from below the basal chamber to the top of the nucellar chamber.
1. Sclerotic tissue below basal chamber. Seed D, peel 475-Т21.
rmost part of basal chamber containing prominent central vascular strand. Seed
D, peel 475-122.
3. Lower part of basal chamber. Seed D, peel 475- T2535.
4. Upper part of basal chamber. Seed D, peel 475-T24.
5. Тор of basal chamber. Seed D, peel 475-T25.
part of nucellar chamber, near nucellar attachment. This and the remaining
figures in this series are from Seed C. Peel 475-Т31.
7. Lower part of nucellar chamber. Peel 475-Т30.
8. Lower third of nucellar chamber. Peel 475-T28.
9. Slightly below center of nucellar chamber. Peel 475-T26.
19461
KERN & ANDREWS—AMERICAN CARBONIFEROUS FLORAS. IX 299
WH N
Text-fig. 4 (Continued).
Peel 475-T23.
Slightly above center of chamber.
Peels 475-T21, T19, T18, T16, T15, T12
10.
From upper part of nucellar chamber.
respectively.
17, 18.
Top of nucellar chamber. Peels 475-T8 and T4.
300
(Мог.
ANNALS OF THE MISSOURI BOTANICAL GARDEN
xt-fig. 5. Kamaraspermum Leeanum. A series of diagrammatic drawings of
longitudinal sections taken parallel to the minor axis of the seed. From seed A.
19461
ІХ 301
KERN & ANDREWS——AMERICAN CARBONIFEROUS FLORAS.
NN
СУ
Text-fig. 5. (Continued).
1; peel 475:133; 2, T32; 3, 1351; А 128082 12516, 124; 7, 125; 8, 121: 9,
'119; 10, Т18; 11, T16; 12, За ОЛТУ; 16; 12,
[Vor. 33
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN
is a dark-colored layer, three or four cells thick, of rather large, irregularly shaped,
thick-walled cells. Within this outer sclerotic tissue is a conspicuous layer of
light-colored parenchymatous cells, small and hexagonal when viewed in transverse
or longitudinal section. This tissue is massive in the lower micropyle region of
the integument (pl. 19, fig. 4), tapering down rather abruptly in the mid-nucellar
region, and tends to become slightly thicker in the transition zone between the
nucellar and basal chambers. Тһе third layer, like the outermost one, is relatively
narrow and consists of dark sclerotic cells. This expands to form a very con-
spicuous tissue between the nucellar and basal chambers. The fourth layer is
narrow, and is composed of longitudinally elongate, brick-shaped parenchyma cells.
Micropyle:—The micropyle of Kamaraspermum Leeanum is а striking struc-
ture and deserving of special comment. Ав may be noted in the median (minor
axis) longitudinal section (pl. 19, fig. 4; text-fig. 3), it consists of two quite
distinct regions: a massive continuation of the integument above the nucellar
chamber, and a much more slender tapering apex. These will be referred to as
the proximal and distal portions respectively. |
The structure of the proximal micropyle differs from that of the integument
below it only in the great development of the parenchymatous (second) layer,
and an absence of the innermost parenchymatous tissue. The distal portion of
the micropyle is approximately 1.5 mm. long and shaped like a much-flattened
inverted funnel, being twice as broad in the plane of the major longitudinal axis
as in the minor axis. Like the proximal region, this portion consists of epidermis,
outer sclerotic and outer parenchymatous layers, the inner sclerotic layer having
terminated in the upper portion of the proximal region.
Nucellus:—The nucellus consists of the remnants of a thin layer of small
brick-like parenchymatous cells surrounding the well-preserved megaspore mem-
brane, and is attached to the rest of the seed only at the base. No pollen chamber
was found in any of the seeds, although judging from what is known of petrified
Pennsylvanian seeds in general, it seems likely that one did exist. If such were
the case it was probably composed of delicate cells which were destroyed prior to
fossilization.
Megaspore membrane:—This appears as an orange-colored band immediately
within the nucellus. It becomes somewhat thicker at the base where the nucellus
is attached to the integument, and sometimes a few scattered cells may be seen
within the membrane.
BASAL CHAMBER—
The basal chamber consists of epidermis, outer sclerotic and outer parenchy-
matous tissue, each identical to and continuous with the respective layers of the
integument surrounding the nucellar chamber. An inner sclerotic layer surrounds
the chamber and appears to be a continuation of the inner sclerotic integument
1946 |
KERN & ANDREWS—AMERICAN CARBONIFEROUS FLORAS. IX 303
around the nucellar chamber. This layer consists of irregular cells, which are
smaller than those of the outer sclerotic layer but resemble them in shape. The
inner edge of this sclerotic layer is somewhat irregular, with a few small scattered
parenchymatous cells occasionally adjoining it. Although not present іп the
specimens at hand, a vascular strand (indicated by the dotted line in text-fig. 3)
probably passed from the peduncle through the center of the chamber. Evidence
for it is an area of vascular tissue visible in cross-sections at the base and at the
top of the chamber. Furthermore, in one of the median longitudinal sections the
chamber is strongly heart-shaped due to the extension of some sclerotic cells down-
ward from the top of the chamber. Тһе fact that some scattered thin-walled
cells are found within the chamber suggests that it might have been occupied with
a loose aerenchymatous tissue.
COMPARISON WITH OTHER SEEDS—
Kamaraspermum presents a number of structural features that render it of
very great interest, yet at the same time preclude it being assigned definitely to
the recognized orders of Paleozoic seeds (Seward, '17, рр. 300—365). Тһе strong
bilateral symmetry and lack of any semblance of trigonocarpous organization in
transverse section seem to rule out the Trigonocarpales. In at least three important
respects it diverges from characteristic Lagenostomalean seeds: Kamaraspermum
is strongly platyspermic; the nucellus is free from the integument; and the integu-
ment as a whole is comparatively thick. Its affinities lie closer to the Cardio-
carpales than either of the previous two orders, and it seems most expedient to
consider it tentatively as a member of this group. Тһе chief conflicting feature
here, however, is the structure of the integument. А typical Cardiocarp seed, as
the present authors understand it, has an integument with a conspicuous and
bulky outer fleshy sarcotesta, while Kamaraspermum presents in the sequence of
its integumentary tissues: first (outermost), a sclerotic layer, then a relatively
fleshy layer followed by another sclerotic one.
It is also appropriate to comment briefly on the prominent basal chamber.
Usually where there is a lack of tissue in a petrifaction the possibility of loss
through decay exists. Yet, since the Iowa seeds are generally well preserved and
a similar basal chamber occurs in the French Codonospermum species (Brongniart,
'74, 781; Renault, 796), it appears likely that the basal chamber existed as such
in life. However, since the chamber region contains some remnants of delicate
tissue it may be that it was occupied by a very loosely organized aerenchyma. In
either event there can be little doubt that it functioned as a float mechanism.
The possibility of Lepidocarpalean affinities has also been considered, There
are certain points of similarity between Kamaraspermum when viewed in median
longitudinal section and a tangential section of a Lepidocarpon taken through the
“heel” or distal end of the sporophyll. The symmetry of Kamaraspermum, its
complex integument and micropyle, nature of the remnants of the outer nucellar
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN
tissue are, upon more critical examination, found to be in no way related to those
of any described species of Lepidocar pon.
пе is almost tempted to apply the nebulous term "missing link" to this
curious fossil, with its non-conformity to established groups, yet the evidence
seems to point in the direction of the Cardiocarpales, to which order it is tenta-
tively assigned.
Diagnosis:—Kamaraspermum Leeanum: platyspermic seed 12 mm. long, 11
x 3 mm. in median transverse section; conspicuous chamber beneath nucellar
region; integument composed of thin outer sclerotic layer, conspicuous fleshy
layer, inner sclerotic layer, and thin inner parenchymatous tissue; micropyle of
two clearly defined regions: a massive continuation of integument above nucellar
chamber, and delicate distal portion shaped like a flattened funnel.
Locality: Urbandale Coal Mine, Des Moines, Iowa.
Horizon: Des Moines Series, Pennsylvanian.
Type specimen: No. WCB475, Henry Shaw School of Botany paleobotanical
collections.
Literature Cited:—
Arber, A. Los 10). ied hong structure of the Palaeozoic seed Mitrospermum compressum (Will.).
Ann. Bot. 24:491-
Arnold, C. A. насы Paleozoic — Bot. Rev. 4:205-
— A. (1874). = es sur les pm с эзчу à l'état silicifié dans le terrain
Houiller A саде Etienn Гом Sci. Мас . V. 20:234-265 (see p. 257).
18
— есһег xol sur les graines I silicifiées. Paris.
Graham, R. mde Pennsylvanian flora of Illinois as revealed in coal balls. I. Bot. Gaz.
95:453—47
Krick, Н. V. (1932). Structure of seedlike fructifications found in coal balls from Harrisburg,
Illinois. yee т :151-172.
Oliver, F. W. and E. J. Salisbury. (1911). E the structure and affinities of the Palaeozoic seeds
of the оета group. Апп. Bot. 25:1-50
E = (1896). Etude des T oen minéraux Ұн la France. Bassin Houiller ес Permien d’Autun
pinac. Flore fossile.
Seward, г "c. (19 кей Fossil Cin III. Cambridge
Williamson, W. C. (1877). On the organization "d the fossil plants of the Coal Measures. pt. viii.
Phil. Trans. Roy. Soc. London 167:213-271.
EXPLANATION OF PLATE
PLATE 18
Fig. 1. Rhabdospermum spinatum. A nearly median longitudinal к. See text-
fig. 2 M cellular structure of integument. From peel 493-T3. Magnified x
Fig Con a oblongum Williamson. 5 pem median ae section
Edict и“ а. ронй рае 1398 Ша
Fig. 3. The micropylar sr of the seed 2. in fig. 2, at a higher magnification.
From “lide 1398. Magnified x
Рі.АТЕ 18
1946
33,
Bor. GARD., Vor.
ANN. Mo.
X
ROUS ELDRAS.. |
IFI
CARBON
RICAN
ME
1
ANDREWS—A
%
| Vor. 33, 1946]
306 ANNALS OF THE MISSOURI BOTANICAL GARDEN
EXPLANATION OF PLATE
PLATE 19
Kamaraspermum Leeanum Kern
Fig. 4. Median longitudinal section along minor axis. Seed A, peel 475-T21. x
м
3.
Fig. 5. Longitudinal section along major axis. Peel 475-T14. x 12.
Fig. 6. Transverse section through lower part of nucellar chamber, just above point
of жар attachment. Seed С, peel 475- 2.
Transverse section through central portion of nucellar chamber. Seed С,
a 47%. Т23. х 12
ANN. Мо. Вот. GARD., Vor. 33, 1946 PLATE 19
KERN « ANDREWS—AMERICAN CARBONIFEROUS FLORAS. ІХ
THE GALLATIN FOSSIL FOREST
HENRY М. ANDREWS лмр LEE W. LENZ
The petrified forests of Yellowstone National Park have been a source of won-
der ever since the early explorations of Jim Bridger, and that wonder grew into a
more realistic admiration of geologic time and the forces that preserve plants
through the ages following the studies of Dr. F. H. Knowlton during the latter
years of the last century. In spite of the fact that some of the plant names in
Knowlton's monograph would probably meet with revision in the hands of a
present-day paleobotanist of Tertiary floras, it is likely that it will always stand
as a classic in the annals of the science. It brought to light a flora, or more cor-
rectly a series of floras, strikingly different from the one that exists there to-day,
and it made generally known the most spectacular of all petrified forests.
Fossil forests representing diverse ages in the earth's history, various modes of
preservation, and a wide range of plant groups have been acclaimed by numerous
authors. Perhaps the best known of all are the petrified trees of Arizona, althoug
there the wood is, for the most part, too highly replaced to be of botanical value,
and the great trees were transported some distance from their original habitat prior
to petrification. Certainly among the most unique fossil floras, from the stand-
point of the plants themselves, is the Devonian Eospermatopteris deposit near
Gilboa, New York, and the Jurassic Cycad forest of the Black Hills. Unfor-
tunately there is but little to be seen of these in the field. Through the work of
the New York State Museum the "Dawn-seed-ferns" of Gilboa have been made to
live again in an admirably executed restoration, and one may catch a glimpse of
one of the earliest forests that existed on the earth. And we may be consoled in
the knowledge that a large and representative series of the Dakota Cycads rests in
security through the vigorous collecting activities of Professor Wieland. The
same author has also given us a picture of the great Patagonian forest, especially
remarkable for the prodigious abundance of petrified Araucarian cones that it has
yielded.
In the coal balls and shales of the Carboniferous there is ample evidence of the
Pteridophytic and early seed-plant forests that once covered so much of the globe,
while occasionally, as with the Lycopod stumps preserved in Victoria Park, Glas-
gow, we see fragments of the forests in place. Another remarkable forest,
preserved in a more precise sense of the word, is the one at Florissant, Colorado.
Here a profusion of foliar remains, along with occasional representatives of the
animal population, are preserved in volcanic ash beds immediately surrounding the
stumps which were petrified in their original position in life. This is a rarely
enough encountered combination of the trees and foliage that they bore. The
impressions have been treated by a number of authors and are deserving of a
comprehensive revision.
(309)
[Vor. 33
310 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN |
The discovery, in some abundance, of the silicified trunks of the Cretaceous
Tempskya tree ferns in various of the northwestern states, and especially Idaho,
has revealed a widespread and unique forest tree. But, like the Arizona petri-
factions and numerous other western fossil wood deposits, these are not found in
their original place of growth and hardly deserving of the term "fossil forest."
All of these forest remains of the past are important and distinctive in their
own way and the age that they represent. Yet none of them can vie with the im-
mense grandeur in both space and time of the Yellowstone forests. Nowhere else
does there exist the succession of one forest directly above the grave of its prede-
cessor—a succession that emphasizes perhaps more forcefully than any other
plant fossil deposit the immensity of geologic time. Individually these forests
attained ages well exceeding 1,000 years, and there is a minimum of no less than
16 of them extending one above the other.
The petrified forests in the Specimen Ridge region south of the Lamar River,
being not far distant from the main stream of travel through the Park, are rather
well known. It is the purpose of this article to focus some attention on the fine
display of fossil forests in the extreme northwest corner of the Park. Although
this is a region that is by no means unknown, it is certainly worth more attention
than it has received from botanists and geologists, whether amateur or professional.
There is a fine camp ground about 300 yards up Specimen Creek from the
Gallatin Canyon highway (Route U. S. 191), and shortly to the north there is
noted a “Fossil Forest" оп the U. S. Geological Survey's map of the Park. From
our own observations the finest exposure of the forests is found about two miles
northeast of this point considerably closer to the summit of Big Horn Peak. While
it is quite possible to make the climb, see a good deal of the forests, and return to
the camp ground on the same day, a two-day trip allows a more leisurely and
profitable study.
An excellent trail leaves the highway and follows along the north side of
Specimen Creek. Two unnamed tributaries may be noted on the topographic map
flowing in from the north, the second of which departs from Specimen Creek
about one and three-quarters miles from the camp ground. About a quarter of
a mile north of the trail this divides into two branches which, for the sake of
clarity, may be referred to as the West Fork and East Fork, although no names
are designated on the map. Оп our trip of last summer we packed in provisions
for overnight and set up camp about 200 yards north of the point where the
two forks join. |
The finest succession of fossil forests that were encountered occur оп the
southwest slope of the spur on either side of which the two forks flow. In ascend-
ing this spur petrified stumps were found at about the 8,000-foot contour, and
splendid displays of at least ten successive forests were counted extending up the
rocky exposed southwest face of the spur.
It is well to emphasize that figures given here are only approximate inasmuch
as surveying instruments were not employed, and the number of forests given is
a very conservative minimum. A "forest" was recorded only where a series of at
1946]
ANDREWS & LENZ—GALLATIN FOSSIL FOREST 311
Upper: A petrified stump of Sequoia magnifica Knowlton, 14 feet in diameter.
Lower: A portion of the Gallatin fossil forest showing silicified stumps at three suc-
cessive forest levels.
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN
құқ ЖА.
ыы E :
~ ET Фе
a e
i
Sv
Upper: An exceptionally tall specimen in the Gallatin fossil forest.
trees weather to within a few feet of the surrounding breccias.
Lower: Two stumps showing roots intact.
Most of the
1946)
ANDREWS & LENZ—GALLATIN FOSSIL FOREST 313
least four or five stumps could be traced along the same horizon, or where rooted
specimens were observed. A more precise counting of the forests was hampered
by two factors—the cover of modern vegetation on the lower slopes of the spur
as well as the upper reaches on and immediately below the ridge, and the nearly
precipitous nature of the terrain above the uppermost reaches of the West Fork,
as is clearly indicated on the topographic sheet of the Park.
Modern vegetation covers the volcanics. at about the 8,500-foot contour of
the spur, and it is necessary to follow along in a westerly direction almost to the
Left Fork. The breccias are well exposed in the stream bed and its immediate
vicinity and above its uppermost limit for another 500 feet or more. Here six
more forests were defined at less regular intervals. It is almost certain that the
actual number preserved here must be at least twice that recorded, but due to the
steepness of the slope most of the stumps do not remain long in position once they
start to weather out.
The average vertical distance between the ten successive forests that were
counted on the spur slope was about 25 feet, with a variation of about 15 to 35
feet. To determine the distance more exactly between forests would require
leveling instruments and considerable excavation inasmuch as only occasionally
are the stumps exposed to the roots. Since the area is most unique and a
National Park as well, the latter treatment would hardly be justified. Thus, while
evidences of sixteen successive eras of forest growth were found on the spur and
the upper reaches of the Left Fork ravine it seems safe to suggest that half again
that number would be revealed by more detailed study. Such evidence is hardly
required to emphasize the spectacular nature of the forests.
On the second day we ascended the spur that lies between the two previously
mentioned tributaries of Specimen Creek. The fossil forests are first met at a
somewhat higher level here partly because the living vegetation cover extends up
higher, and partly because the beds dip toward the southeast. Following this spur
to about the 8,300-foot contour one may then traverse about 100 yards to the
west into an exceedingly rugged ravine where numerous stumps are exposed
through a vertical distance of some few hundreds of feet. The successive forests
cannot be traced as clearly in this sector although it is of interest for the large size
of some of the stumps, a Sequoia 14 feet in diameter being the largest that we en-
countered. Although a central core some 5 feet in diameter had been destroyed
in this tree prior to fossilization a study of wood specimens from the remaining
part of the trunk showed an average of 19 rings to the inch, indicating an age of
about 1,600 years for the tree.
Unfortunately there were few evidences of well-preserved foliar remains in
the territory that was covered. The forests do extend for some distance to the
northwest, however, and it is possible that leaf impressions might be found at
other points. If such were located the possibility of an ecological study is evi-
dent and should produce most interesting results. Aside from this the region is
well worth a day’s time for any naturalist with a paleontological bent.
ЕГОКА ОЕ РАМАМА
ВҮ
ROBERT E. WOODSON, Jr.
AND
ROBERT W. SCHERY
AND COLLABORATORS
PART ПІ
Fascicle 3
ORCHIDACEAE (Williams)
(second part)
ANNALS
OF THE
MissourI BOTANICAL GARDEN
Vol. XXXIII NOVEMBER, 1946
FLORA OF PANAMA
Part III. Fascicle 3
ORCHIDACEAE
Bv LOUIS O. WILLIAMS
27. EPIDENDRUM L.
Ершемокум L. Sp. Pl. ed. 2. 1347. 1763; Ames, Hubbard & Schweinfurth,
Genus Epidendrum іп U. S. & Middle America, pp. 1-233. 1936. Non L. Sp.
Pl. ed 1. 952. 1753.—Сопзегуед name, the generic type being Epidendrum
nocturnum. Jacq.
Amphiglottis Salisb. in Trans. Hort. Soc. Lond.1:294. 1812, nomen nudum.
Auliza Salisb. loc. cit., nomen nudum
Dinema Lindl. Collect. Bot., App. 1825, nomen subnudum.
Encyclia Hook. in Bot. Mag. 55:1. 2831.
Nanodes Lindl. in Bot. а yi і. 1541. 1832.
Physinga Lindl. in Bot. 4: Misc. p. 32. 1838.
Barkeria Knowl. & Westc. "EL. "Cab. 2:7, Е. 40. 1838.
Hormidium Lindl. ex Heynhold, фы Bot. Hort. Nachtr. 880. 1841.
Spathiger Small, Fl. Miami, p. 55.
Dimerandra Schltr. in Fedde Rep. Sp. Nov. Beih. 17:43. 1922.
E
Small to large, epiphytic or semi-epiphytic herbs or subshrubs with or without
the stems swollen into pseudobulbs. Stems (secondary) swollen into pseudobulbs
and the leaves borne at or near the apex of the pseudobulbs ($ Емсүсілом); or
the stems slender and not swollen and with the leaves scattered along the stem
($ Есершемркум); or slender or fusiform stems with the leaves scattered along
them but with the base of the stems swollen (6 ВАККЕКЛА). Leaves variable in
shape and size, usually coriaceous but occasionally fleshy. Inflorescence commonly
terminal but occasionally lateral or both terminal and lateral, rarely borne from
lateral, leafless shoots which later develop into pseudobulbs or stems; from simple
and often 1-flowered to compound and many-flowered. Flowers from small to
relatively large. Sepals equal or subequal, spreading, reflexed or subconnivent.
Petals similar to the sepals but usually smaller and narrower or even filiform, oc-
casionally broader. Lip unguiculate; the claw free, partially free, or adnate to
the column; lamina simple or 3-lobed, the lobes often lobulate; the disc ecallous
or usually bicallous and with or without lamellae or additional calluses. Column
stout or slender, alate or exalate, clinandrium variously developed. Anther term-
inal, operculate, incumbent, 2-celled and with each cell more or less divided by a
septum; pollinia 4, uniseriate, laterally compressed, with appendages, rarely with-
out, waxy. Capsule various, winged or wingless.
Issued December 7, 1946.
(315)
(247)
[Vor. 33
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Epidendrum is the largest genus of neotropical orchids. It contains about one
thousand species. Certain species are exceedingly abundant in their range and in
places might almost be called weeds.
The most recent study of this vast genus is Ames, Hubbard, and Schwein-
furth’s admirable monograph, cited above. We are much indebted to the authors
of the fine key in that work, which has been followed as closely as possible in the
following key to the Panamanian species. We are particularly indebted to Mr.
Charles Schweinfurth for suggestions and criticism during its preparation, and to
Prof. Oakes Ames for his generous permission to use many of the illustrations.
a. Stems with true pseudobulbs surrounded by scarious non-leaf-bearing
—€— or —M leaf-bearing sheaths; leaves 1 or more borne at
or the the pseudobulbs; lip free iis the column or
vario bd ay І. ENcvcLIUM (p. 322)
aa. Stems la vem pen I although sometimes thickened; рейн
distributed along the or at least leaf-sheaths surrounding the
о stem.
ib. Lip free from the column or adnate only at the base; the col-
umn usually winged; the stems swollen at the base or pp
cylindric ect. П. BARKERIA (р. 338)
bb. Lip trien adnate to the entire length of the column oe
less) ; the column not winged laterally; stem never swollen....Sect. Ш. EuEPIDENDRUM (p. 338)
Sect. І. ENcYCLIUM
a. Lip entire or at most obscurely lobulate.
b. Pseudobulbs 1-leaved.
с. u^ ligulate or pandurate, slightly er — the apex.
vary 15 Һап
muricate; sepals more than 3. E. Воотнп
Or vary smooth; sepals less 2 10 mm. = 13. E. OTTONIS
се. Lip ovate to чы rbicular-cordat
d. Leaves wly linear 20. E. STANGEANUM
dd. Leaves aee to oblong-elliptic.
e. Lip hastate-rotund, acute; sepals oblong-lanceolate 18. E. SPONDIADUM
ee. ты ovate to ovate-subrotund, acuminate; sepals linear-
lanceolate 10. Е. FRAGRANS
bb. Pseudobulbs 2- or more leaved.
c. Inflorescence — ied peduncle short or 1. E. ABBREVIATUM
cc. PM rescenc t abbreviated; peduncle more or tes yim
d. ina of ss lip к longer than
е. Apex of the lip r а d, obtuse, truncate ог retuse [Species
reported in fir, Ser 0. E. LIVIDUM
ee. Apex of the lip acute or acuminate 4. E. BRASSAVOLAE
dd. Lamina of the lip as broad as long or broader than lon
e. Bracts e inflorescence conduplicate and clasping the
edicel, conspicuous; rac exuose; pseudobulb stipitate......22. E. VOLUTUM
ee. Bracts of the inflorescence | concave to flat; rachis straight or
i nconspicuously flex
Callus of the lip a a sail or bilobed cush
в. Disc of. the lip occupied by a central "bilobed cushion-
like callus; pope т petals usu Шу maculate 9. Е. CRASSILABIUM
gg. Disc of the Пр with a d мр ba И cushion-
like callus; жы апа peta tals maculate 11. E. IONOPHLEBIUM
ff. overs 29 the lip consisting ha or ыл basal keels ог
wellin 7. Е. COCHLEATUM
aa. Lip -lobed or 3 -lobul
b. Column with a ресми lateral auricle on each side about opposite
the TÉ
© Colu umn more or less sharply и in the middle; veins of the
mere lobe of the lip smooth or nearly 12. E. ONCIDIOIDES var.
RAMONENSE
(248)
19461
FLORA OF PANAMA (Orchidaceae) 317
cc. Column straight or nearly so; veins on the mid-lobe of lip often
d. Lateral lobes of the lip triangular-lanceolate to oblong.............. 12a. E. ONCIDIOIDES var.
GRAVIDUM
dd. Lateral lobes of the lip spatulate or oblong-spatulate.................... 12b. E. oNciDIOIDES var
bb. жола "em distinct lateral auricles on either side of th а.
. Mid-lobe of lip proportionally narrower at base, тай. ог
re ps: filifor
ateral liba of lip fimbriate-ciliate . E.c
4 pio 1 lobes of lip entire to somewhat депсаїе............................ T s турин Р
сс. еа lobe of lip и broader at base, not Тү: Uo pem
r linear-filiform
d. Outer margins o£ mid-lobe p lip dentate to fimbriate................ 19. E. STAMFORDIANUM
dd. Outer margins of mid-lobe of lip not dentate to fimi ria
e. Mid-lobe of 5 lip mid than either lateral lobe or 2 38
equal to either of them.
К 242 bracts conspicuous, . long or more,
p 3-lobate; mid-lobe бйлу: as broad as long.................. E. VOLUTUM
5 т - ply 3-partite; mid-lobe oblanceolate, longer E
broa
6a. E. CILIARE var.
ERSTEDII
ff. P bracts not very а екоо 10 mm. or less long.
. Mid-lobe М m lip triangular 16. E. PYGMAEUM
Pa 55 каф 12 not trian ngular
h. Petals is 2 ап one-half as broad as the lateral қы ай
iria esc commonly basal 7. E. ROUSSEAUAE
hh. dead at most only e: narrower than the iau
9. E. CRASSILABIUM
inflor
ee. yrs lobe of 5 lip ЕТ larger than each lateral lobe.
f. Pseudobulbs 1-leav
5 ps of the lip gradually narrowed below the ae
lobes; floral bracts large, chartaceous Е. CAMPYLOSTALIX
gg. Base of the lip ups Ју contracted below the был:
lobes; floral bracts small 12a. E. ONCIDIOIDES var.
GRAVIDUM
ff. Pseudobulbs 2- or more leaved.
g. ре contracted into a шт». с neck, 4-10 cm.
ong, iar ben leaves ied the swollen portion...... 21. E. VARICOSUM
£g. Pseudobulb not contracted into a _aulecen Mick, leaves
almost aia on pea swollen P
h. Mid-lobe of lip trulliform, acuminate ..15. E. PRISMATOCARPUM
hh. ra lobe of lip not trulliform nor acuminate.
iddle clinandrial tooth at summit of colum
y and em or ана.
rate, distinct from lateral t 8. Е. CONDYLOCHILUM
ii. Middle insueta tooth at summit of column
small, deltoid, incurved, obtuse or acute, separated
by as і
re
5 Е
2
Us
т
со
б
г
8%
us.
mid-lobe of lip about 4-10 times that
u
k. gin Ка mid-lobe of lip entire to eros
crenulat 2. E. ATROPURPUREUM
kk. Margin d mid-lobe of lip dentate-lacerate...... 2a. E. ATROPURPUREUM
var. LACINIATUM
. Area of mid-lobe of lip about 3 times that of
either жене lobe, or less
k. Column somewhat sharply recurved in the
idis veins of wb. lobe usually smooth........ 12a. E. qut
var. RAMONENSE
kk. Column ex: d Ы мн veins of mid-lobe of
lip often verru
1. Lateral lobes of lip spatulate to oblong.......... 12b. E. ONCIDIOIDES
var. MOOREANUM
ll. Lateral lobes of lip deltoid-lanceolate to
oblong
12a. E. oNcIDIOIDES
var. GRAVIDUM
(249)
[Vor. 33
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Sect. II. BARKERIA
23. E. STENOPETALUM
One species in Panama
a. Leaves equita
b. Flowers lo amd ааа and terminal
b. Flowers sessi «v or nearly so and axillary
Sect. III. EuEPIDENDRUM
43. E. EQUITANTIFOLIUM
50. E. LOCKHARTIOIDES
aa. ei
b. Lip dis ti er 5. oid or 3-lobulat
c. Leaves terete, or subterete and ры or flat and woes
d. Mid-lobe of lip vos plains тир apiculat 65. E. PUDICUM
dd. Mid-lobe of lip elliptic to narrowly oblong, и with a
inen
t lamellate callus 75. Е. TERETIFOLIUM
d usually more tha А
with a distinct dorsal horn at the apex 51. E. MICRODENDRON
dd. Column without a distinct dorsal horn at the apex.
e. Mid-lobe of lip narrowly triangular to linear, several times
longer t
than bro
f. Lateral lobes ч the lip laciniate-fimbriate
ff. Lateral lobes of the lip not pos fimbri
35. E. CRINIFERUM
nce subtended by one or more large spathes........ 54. E. OBESUM
d
g. Infloresc
gg. ae кука usually кеі subtended by spathes; if so
b all i
en by sma
inconspicuous one Е. NOCTURNUM
ee. Mid- lobe of the lip not інді pee to lin
Lip tubular-involute, usually fleshy, чы pnm
to ovate-tria ngular, longer dis "m a 72. E. SCULPTUM
н. eg not tubular-involute, yng flat.
of
h.
2 lateral lobe or even lar
Out
the mid- em of 7 "ne twice that of
ter surface of ан se је А уеггисоѕе 44. Е. EXASPERATUM
f
hh. ge! surface of the lateral sepals smooth.
of
mid-lobe of lip rounded or protuberant-
j. Floral bracts зыш, cucullate, only slightly short-
the flow
er than 5; ийи cence subcapitate with
1 flower appear time 41. Е. ELLIPSOPHYLLUM
. Floral dem sm mall үе if large then much short-
er than the flowers), not cucullate; inflorescence
not subcapitate се normally with several flowers
at a time.
k. Lateral lobes of the = dentate to lacerate; in-
florescence usually long-pedunculate 47. E. IMATOPHYLLUM
kk. Lateral lobes of = Бр пос — nor lac-
erate; inflorescence not long-pedun
1. Lip spatulate, base of the mid- be ы” апа
пос і . WARSCEWICZII
z
З
ТЕ
>
ті
9
ll. Lip not s one base of Т mid-lobe pet
a narrowed or unguicu ase . E. DENTILOBUM
ed o ulat
ii. Apex of the mi -lobe of - lip а, retuse or
e
bilobed, жеке apiculat
і. Lateral lobes of lip digita, lacerate or -
dentate; sehe e pink, maculate 7. E. CALIGARIUM
ji. Lateral lobes of the lip o entire to minutely crenu-
late or crenate- дерінің г bilobulate.
k. Leaves obtus E minutely bilobed at the apex,
usually ovate to elliptic or elliptic-lanceolate.
| Inflorescence paniculate; sepals ап4 petals
1
p
maculate SCHUMANNIANUM
Il эн racemose; sepals and petals not
aculat 2. E. ENDRESII
kk. Leaves acute minate, not bilobed at m
apex, usually Taper. lanceolate to lanceolat
l. Lobules of the mid-lobe of lip pin times
larger than the dentiform lateral lobes; in-
florescence usually subumbellate or subcorym-
bose
29. E. CENTRADENIA
(250)
19461
FLORA OF PANAMA (Orchidaceae) 319
Il. ре es of the mid-lobe
of the lip about
ualling in size rn oblong or falcate-lance-
e lateral lobes, or smaller; inflorescence
usually racemose E. CENTROPETALUM
gg. Area of mid-lobe of lip about equal to that of either lat-
era
1 lobe or 1
ess.
h. Ovary with a saccate vesicle at the summit 58. E. PHYSODES
hh. Ovary without a saccate vesicle at the
Colum
i
umm
n, when seen from the side, like an ox- Ee .39. E. DIFFORME
. Column, when seen from the side, not like an ox-
Ow, at most Ms. «Гулу: rched.
h pure 1 flowered or subumbellat
Sepa
k.
als owly lanceolate, long-acuminate;
stems usually straight c or nearly so; usually but
1 flower tim 40. E. EBURNEUM
kk. Sepals mas ме ре че to oblanceolate ог
obova
-oblong; stems flexuose.
1. Sepals more than 1 cm. broad; inflorescence,
f k
as far as known, 1-flowered 57. E. PENDENS
II. re less than 0.8 cm. broad; inflorescence
sually more than 1-flow
were
m. Mid-lob be of lip, when re nt, twice or
more broader Фа long, semi-elliptic to
orm 39. E. DIFFORME
mm. Mid- Е: - lip usually little p than
long; n ad, subquadrate to sub-
ne ES -OV
n. Apex of he mid- lobe truncate, retuse
Кый і bilobed, usually ын” 758 E es DIFFORME var.
MUM
nn. е of Bes mid-lobe of the lip tri-
gular, acu 39b. E. DIFFORME var.
SIMULACRUM
jj. пр racemose ог paniculate.
—
==
еа
. Mid- Pos m. the lip bilobed or re
Uv» v ње: је lip not ос пог retu
а dense spike-like raceme um
is more or Ele Б. та leaves linear-lanceo-
late 6
—
Е.Рохкілп
. Inflorescence not а dense spike-like raceme;
penes elliptic to lanceolate or oblanceolate.
m. Sum column n a denticulate,
sometimes 4-lobed w 40. E. EBURNEUM
mm. Д of y columan n.
Lip up to about . broad; stems
> 25. Е. АМСЕР5
2:4 12 mm. or more broad; stems
48. E. INCOMPTUM
l. Margins of ж ateral lobes of lips yc lac-
a or den
. Peduncle praed with close scarious
tubular sheaths and wit ers i
d es etimes paniculate;
s
mm. Peduncles not Wee gated o
the sheaths аут e т ог іт
pier 2 stems not bearing roots dimit
lea
the
п. Mid. lobe of the lip with two strongly
explanate lobules 56. E. PANICULATUM
nn. Mice of the lip with the pie
more or less porrect, never strongly
planate.
o. Lateral sepals 7-16 mm. long.............. 60. E. POLYANTHU
oo. Lateral sepals 4-6.5 mm. long............ 60a, E. POLYANTHUM
var. MYODES
(251)
[Vor. 33
320 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
Il. a of lateral =... оғ lip not digitate,
lacerate, nor dentate, s metim crenate, un-
ca
m. Spread between the tips of c divaricate
lobules of the mid-lobe of the lip about
tw = or more greater than the length of
the lobes 74. E. SUBNUTANS
mm. Spread between the tips of the
less porrect lobules of the mid- in іе lip
or brad of the mid-lobe less than twice
the length
n. Lip 2 cm. or more broad; raceme usual-
4 ани and more than 6 ст. іп
i 52. E. MOYOBAMBAE
nn. cA; 1.5 or less broad.
o. Pair of phe calluses of the lip ab-
nt; peduncle usually elongated 25. E. ANCEPS
se
oo. Pair of basal calluses of the lip present.
p. Peduncle conspicuously winged and
ancipi itous 24. E. ALLENII
pp. Peduncle not winged and ancipi-
tous.
а. Lateral sepals 6.5-16 mm. long.... 60. E. POLYANTHUM
qq. Lateral sepals 4—6.5 mm. long... 60a. E. POLYANTHUM
var. MYODES
bb. Lip simple, obscurely lobate or bilobed.
er portion of lateral margins of lip irregularly lacerate to
coarsely eod ate; lamina with a prominent central keel extending
almost to the apex 47. E. IMATOPHYLLUM
. Lower portion of lateral margins of lip neither lacerate nor
coarsely dentate.
d. Leaf- blade not articulated to the sheath, thus not deciduous.
e. Plants elongated б жай creeping; lateral sepals without a keel
on the outer surfac 2. E. CONFERTUM
ee. = not elongated and creeping; lateral sepals keeled on the
urfac
f. Lip con Ир acute; clinandrium entire or denticu-
ate 33. E. CONGESTUM
nifor piculate; clina праве lacerate......70. E. SCHLECHTERIANUM
dd. D T Made қак етер to the riven thus deciduo
. Ovary with a more or less semiglobose vesicle at np summit
r the lip.
nterior portion of the lamina of lip broadly rounded.
g. Leaves triangular-lanceolate or nins, scattered along
tem; li
the s lip not strongly conduplicat 58. E. PHYSODES
gg. e elliptic to ym MN congested on the upper
of the ; lip ngly conduplica 28. E. Canoni
ff. Anterior “portion a the gy pare of the "lip acute or
73. E. STROBILIFERUM
ee. Ovary d a semiglobose vesicle at the summit.
ack of the column terminating in a distinct, more or less >
a eading Bed 1. E. MICRODENDRON
ff. mcn of the column not terminating in a distinct spreading
hor
5. Lip with a tridentate (not қан) or V-shaped
callus at or near the base of the lam
h. Leaves usually sulcate, terete or cc if flattened
n bed т linear-lanceolate, often grass-like
and a 49. E. ISOMERUM
hh. Pow da at most conduplicate, ligulate or linear-
oblong to 52 or lanceolate to nearly elliptic, ob-
tuse, bilo
i. Lamina of the д suborbicular-cordate, commonly
broader than | 59, Е. PLATYSTIGMA
(252)
19461
FLORA OF PANAMA (Orchidaceae) 321
ii. Lamina of the lip triangular to ovate-triangular,
usually longer than broad.
j. Margins of x lip ых fleshy and thickened........ 69. E. SANCHOII
ji Margins of the lip not fleshy and thickened.
- 5 т. о
ua
eter below the inflorescence; leaf-sheaths and
bracts not maculate; inflorescence usually lax
and narrow 66. E. RAMOSUM
kk. vg p - usually 5-6 in Dr
ust below the PW фол scu sheaths and
айн ный Ё nely maculate; inflorescence
typically rather congested and broad.................. 66a. E. RAMOSUM
Var. ANGUSTIFOLIUM
85. he never with a e mee or V-shaped callus at or
ar the base of А lam
E Lamina of the ж triangular with a truncate base;
inflorescence 1-flow 78. E. TRIANGULABIUM
hh. Lamina of the lip never triangular with a truncate
base.
i. ку length of column adnate to base of lateral
61. Е. РОВРАХ
fone length dm column not adnate to the base of
the lateral sep
aring.
l. Clinandrium es dentate; iw ig m.
falcate and acumin plant dwarf................ . E. DENTIFERUM
ll. Clinandrium пог 34 erate- sd. E
sepals no
m. bein bracts very conspicuous, large and
31. Е. COCLEENSE
mm. For bracts not conspicuous, small and
о
п. Disc of lip bed earing more or less lobate
or retuse calluses, usually situated well
above the base 77. Е. TRACHYTHECE
пп. Disc of the lip not bearing more or less
lobate or retuse bees es.
o. Clinandrium 4 subr iig өң
pea lobes, bes central pair lon
est narrowest 76. E. TETRACEROS
е арі: lacking 4 lobes................ 26. E. BISULCATUM
kk. Lip eca A (mid-nerve sometimes thickened).
1. Inflorescence 1-flowered 67. E. REPENS
ll. pde. 2- or more flowered.
"I ess йу 1 cm. broad, linear 94
gs ar-o . E. SANCHON
mm. etn more ethan 1 cm. broad, oed
elliptic-o 36. E. CRYPTANTHUM
jj. Stems dim che
k. Floral bracts conspicuous due to their
compared to the flower, or to their гн
spreading chara ter.
Lami f the lip broader than long, reni-
form, bilobed or pie se 34. E. CORIIFOLIUM
ll. Lamina of the lip longer than broad or at
most ну slightly broader than long.
. Flowers clustered at the end of a long
kise pon ncle.
n. Peduncle doles NN winged and an-
cipitous 24. E. ALLENII
nn. Peduncle not conspicuously winged and
ancipitous 55. E. PANAMENSE
mm. Flowers not clustered at the end of a
long T peduncle.
n. Lamina of the lip cordate-orbicular,
Eos as long as broad 68. E. RIGIDUM
(253)
[Vor. 33
322 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
nn. Lamina of the lip oblong-ovate, longer
than broad
. E. RIGIDUM var.
ANGUSTISEGMENTUM
е
со
~
kk. Floral bracts not conspicuou
1. Inflorescence 1-flowered or subumbellate.
m. Lateral sepals narrow M lanceolate, long-
ually 1 flower open
lip white 40. E. EBURNEUM
3 5 br г ођ-
lanceolate to elliptic-oblong ovate-
hare, id leaf-sheaths dilated bud their
n. ‘Leaves semiterete and sulcate, linear-
65. E. PUDICUM
nn. i flat conduplicate, usually
18 ulat te to elli dr
ки af-sheaths strongly cuneate (viewed
у аи large and flaring, the folded
sheath 2 cm. broad at junction with
cm. br е жәй folded 45. E. HUNTERIANUM
oo. Leaf-sheat t br ced = y the
folded 2 ыы се . broad
at junction wit ығ ame more
or less flat, u se и not stro - con
duplicate, Bruns so, less t 1 cm.
broad when folded.
p. gin of = dpa rtm —
than lon
"uu Bette 39. E. DIFFORME
pp. Lamina of the lip about : as н broad
as long, жа "и ога 39Ь, Е, DIFFORME
var. SIMULACRUM
ll. Inflorescence racemose or paniculate, mostly
3- fl
о |
т. Lateral sepals 22 mm. ог more long
minate; lip subentire, white............... 0. E. EBURNEUM
nn. Lateral sepals narrowly spatul
acute or obtuse; lip denticulate or
nate-dent er. оннан 2 T DN 64. E. PSEUDEPIDENDRUM
mm. ioe sepals or less lon
п. 2 co duplicates “peduncle elongated;
aceme elongated and n
o. Apex of t E lip Eu. rounded and
retuse-apiculate; lateral sepals with а
orsal keel toward the арех................. 28. E. САКОШ
oo. Apex of the lip short-acuminate; lat-
eral sepals without a dorsal Кесі........ 63. E. PROBIFLORUM
nn. Lip not conduplicate, flat or at most
what convex 25. E. ANCEPS
Sect. I. ENCYCLIUM
1. EPIDENDRUM ABBREVIATUM Schltr. in Fedde Rep. Sp. Nov. 3:107. 1906.
Epidendrum prorepens Ames, Sched. Orch. 2:33. 1923.
Small, repent, epiphytic herbs up to 2.5 dm. tall. Pseudobulbs 3.5—10 cm. long
апа 0.3-1 cm. in diameter, fusiform, occasionally stipitate, normally bifoliate.
Leaves 4—14 cm. long and 0.3—1.5 cm. broad, obtuse, ligulate, subcoriaceous. In-
florescence a short, 1- to 5-flowered raceme up to about 4 cm. long. Dorsal sepals
10-15 mm. long and 2.5—3.5 mm. broad, linear-oblanceolate, obtuse or acute,
(254)
19461
FLORA OF PANAMA (Orchidaceae) 323
often carinate dorsally along the midrib. Lateral sepals 10-15 mm. long and 3-4
mm. broad, linear-lanceolate, obtuse to acute. Lip unguiculate, the claw adnate
to the column, free portion 7-10 mm. long and 5-8 mm. broad, ovate to oblong-
ovate; disc with a conspicuous callus-thickening at the base.
Guatemala, Honduras, Costa Rica and Panama.
CHIRIQUÍ: Boquete District, alt. 1300 m., Davidson 666; “Chiriquí”, alt.
Powell 152, 3560. cocrf: vicinity of La Mesa, hills north of El Valle de Ал, с
1000 m., Allen 2354. vERAGUAs: Santa Fé, alt. 500 m., Powell s. n.
2. EPIDENDRUM ATROPURPUREUM Willd. Sp. Pl. 4:115. 1805; Ames, Hub. &
Schweinf. Genus Epidendrum іп U. 5. & Mid. Am. 60. 1936.
Epidendrum auropurpureum Lindl. Gen. & Sp. Orch. Pl. 99. 1831.
Epidendrum macrocbilum "a іп Bot. Mag. 63: f£. 3534. 1836
Epidendrum macrochilum Hook. var. roseum Batem. rm Det & Guat. Ё. 17. 1839.
Encyclia atropurpurea atin Orchidéen, p. 208, fig. 49.
Encyclia atropurpurea var. leucantba Schltr. in Fedde ES МЫ Nov. Beih. 17:45. 1922.
Encyclia atropurpurea var. rhodoglossa Schltr. loc. с
Robust caespitose epiphytic herbs. Pseudobulbs 3-8 cm. long or more and
1.5-5 cm. in diameter, ovoid, 1- to 3-leaved, usually bifoliate. Leaves 8-35 cm.
long and 1.5-3.5 cm. broad, elliptic-ligulate or ligulate, obtuse or acute, coriaceous.
Inflorescence a simple few- to many-flowered raceme up to 6 dm. long. Dorsal
бергі 2.5—3.5 cm. long and 0.4-1.2 cm. broad, oblanceolate, obtuse. Lateral sepals
similar to the dorsal but usually arcuate. Petals 2.5—3 cm. long and 0.7-1 cm.
broad, oblanceolate to obovate, unguiculate, obtuse or acutish. Lip 3-5 cm. long
and 1.8-4 cm. broad, 3-lobed, free nearly to the base of the column; lateral lobes
oblong, obtuse or acute, relatively small, enfolding the column; mid-lobe large,
variable, oblong to obovate to suborbicular to flabellate, always more or less
emarginate.
Mexico, Guatemala, Salvador, Nicaragua, Costa Rica, Panama, the West Indies
and in South America.
NCE NOT KNOWN: “Panama”, Sinclair. снікюоі: between El Boquete and
Caldera, alt phage m., Pittier 3331. cocrÉ: Penonome, alt. 15-300 m., Williams 440.
en, north of Chepo road, Hunter and Allen 851; ва Juan, “all
pi 2 re Pacific side", foothills east of city, Powell 80, 148, 140, 3106, 3226.
The copy of the Plumier plate (ed. Burmann) depicting this, which I have
seen, is really not satisfactory for absolute identification. Lindley (Gen. & Sp.
Orch. Pl. 100. 1831) had seen a copy of Plumier's plate in Lambert's herbarium
and characterizes that of Burmann as a "caricature."
2a. EPIDENDRUM ATROPURPUREUM Willd. var. LACINIATUM Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:63. 1935.
Margins of the mid-lobe of the lip lacerate-dentate and the lateral lobes some-
what smaller than in the species (a malformation?).
Panama.
PANAMA: foothills east of Panama City, alt. sea-level, Powell 287.
3. ЕРРЕМРКОМ Воотни (Lindl.) L. Wms. in Ann. Missouri Bot. Gard. 26:282.
1939.
(255)
[Vor. 33
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Maxillaria pee Lindl. in Bot. Reg. 24: Misc. ‚Р К 1838.
nema paleaceum Lindl. loc. cit. 26. Misc. p. 1840
Epidendrum psu Lindl. loc. cit. 29: Misc. H 4. 43.
те paleaceum Reichb. a in Beitr. Orch. Centr.-Am. 80. 1866; Reichb. f. in
unders Ref. Bot. 2: f. 87.
E ema Bootbii Schltr. in Fedde E. E Nov. Beih. 17:43. 1922.
Repent or subcaespitose epiphytic herbs up to about 3 dm. tall Rhizome
creeping. Pseudobulbs up to about 5 cm. long, cylindric, fusiform or narrowly
ovoid, stipitate, unifoliate. Leaves (when mature) 5-22 cm. long and 0.4-1.2
cm. broad, linear-ligulate, obtuse or acute, coriaceous. Inflorescence a 1- to few-
flowered raceme; flowers small; bracts mostly 2-3 cm. long, lanceolate, cucullate,
paleaceous, relatively large. Dorsal вера! 12-17 mm. long and 3-4 mm. broad,
narrowly lanceolate to lanceolate, acute. Lateral sepals 11-15 mm. long and 2-3
mm. broad, linear-lanceolate to lanceolate, acute or acuminate, arcuate. Petals
7-12 mm. long and 2-4.5 mm. broad, lanceolate to elliptic-ovate, acute or acumi-
nate. Lip 8-11 mm. long and 2-3.5 mm. broad, oblong-oblanceolate, obtuse or
acute, slightly constricted above the middle, fleshy, bicarinate toward the base;
disc often verrucose toward the apex.
From Mexico to Panama, the West Indies and northern South America, mostly
at low elevations.
s DEL TORO: Rio Cricamola, alt. about 10-15 m., Woodson, Allen & Seibert 1892;
Isla "Colón, Wedel 123; vicinity of Chiriquí Lagoon, Wedel 1090.
A species of low altitudes, often taken to be а Maxillaria.
EPIDENDRUM BRASSAVOLAE Reichb. f. in Bot. Zeit. 10:729. 1852; Hook. A
in Bot. Mag. 93: ¢. 5664. 1867.
Repent or caespitose epiphytic herbs up to about 5 dm. tall. Pseudobulbs up
to 22 cm. long, from cylindric to ovoid, 2-, rarely 3-foliate. Leaves 12-26 cm.
long and 2-4.5 cm. broad, oblong-ligulate to narrowly ovate, obtuse, coriaceous.
Inflorescence a simple few-flowered raceme, exceeding the leaves; sheath incon-
spicuous, up to 12 cm. long. Sepals 3.5-5 cm. long and 0.3—0.5 cm. broad,
>
linear-lanceolate, acute or acuminate, spreading. Petals similar to the sepals but
slightly narrower. Lip 3-4.5 cm. long and 0.7-1.5 cm. broad, cuneate to a nar-
row base and adnate to the column for nearly the length of the column, simple;
lamina 2-3 cm. long and up to 1.5 cm. broad, lanceolate-ovate, acuminate; claw
with 2 short lamellate calluses.
Mexico, Guatemala, Honduras, Costa Rica and Panama.
CHIRIQUÍ: “Chiriqui Vulcan", Warscewicz.
This, like many another of Warscewicz’s collections, is said to have come from
Chiriqui, but has not been collected there since.
5. EPIDENDRUM CAMPYLOSTALIX Reichb. f. in Bot. Zeit. 10:730. 1852; in Saun-
ders Ref. Bot. 2: 7. 86. 1869; Summerhayes in Bot. Mag. 154: #. 0243. 1928.
Encyclia campylostalix Schltr. in Fedde Rep. Sp. Nov. Beih. 17:45. 1922.
(256)
19461 ;
FLORA OF PANAMA (Orchidaceae) 325
Caespitose epiphytic herbs up to about 4.5 dm. tall. Pseudobulbs 2-12 cm.
long and 2-4.5 cm. broad, ancipitous, narrowly oblong to ovoid, unifoliate. Leaves
4-30 cm. long and 1.3-8 cm. broad, elliptic, acute, subcoriaceous. Inflorescence
ап erect, 1- to many-flowered raceme subtended by a paleaceous sheath 1-7.5 cm.
long; floral bracts 0.8-4 cm. long, linear to linear-lanceolate, acute, cucullate,
paleaceous. Sepals similar, 16-20 mm. long and 3-4.5 mm. broad, linear-lanceo-
late, acute, the laterals slightly arcuate. Petals 13-17 mm. long and 1.5-2.5 mm.
broad, ensiform to narrowly linear-oblanceolate, acute. Lip 12.5-18 mm. long
and 6-9 mm. broad, unguiculate, only slightly adnate to the column at the base,
3-lobed; claw narrow, comprising about half the length of the lip; terminal por-
tion of lip subquadrate in outline; lateral lobes from small and rounded to oblong,
obtuse; mid-lobe rounded to obovate, disc with an inconspicuous callus thickening
or low raised lamellae. Ovary conspicuously 3-winged.
Costa Rica and Panama.
CHIRIQUÍ: forests around El Boquete, alt. 1000-1300 m., Pittier 3047; “Chiriqui”,
alt. 1600 m., Powell 74.
This species is variable in size, and many specimens appear depauperate.
6. EPIDENDRUM CILIARE L. Syst. Nat., ed. 10:1246. 1759; Lindl. in Bot. Reg.
10: 7. 784. 1824
Auliza ciliaris Salisb. in Trans. Hort. Soc. Lond. 1:294. 1812
Ros rum cuspidatum Lodd. Bot. Cab. 1: Ё. 10. 1816; Скат. in Bot. Reg. 10: #. 783.
ыыы: viscidum Lindl. їп Bot. Reg. 26: Misc. р. 81. 184
Phaedrosanthus ciliaris O. Ktze. in Post & Ktze. Lex. Gen. Es 429. 1904.
Repent or caespitose epiphytic herbs up to about 5 dm. tall. Pseudobulbs 8—22
cm. long and 0.6–2.2 cm. in diameter, cylindric, fusiform or tapering to the base,
1- to 2(—3)-leaved. Leaves 8—28 cm. long and 2-8 cm. broad, oblong-lanceolate,
elliptic-oblong to oval, obtuse, coriaceous. Inflorescence a few- to several-flowered
raceme; bracts 3.5—6.5 cm. long and 0.6—1.5 cm. broad, lanceolate to oblanceolate,
cucullate, pergameneous. Sepals 35-85 mm. long and 1.5-6 mm. broad, linear to
linear-lanceolate or linear-oblanceolate, acute or acuminate. Petals 35-70 mm.
long and 1.5-5 mm. broad, linear-lanceolate to linear-oblanceolate, acute or
acuminate. Lip 4-8 cm. long, 3-lobed, the base adnate to the column; lamina
3-6 mm. long, with 2 short lamellate calluses at the apex of the column; lateral
lobes about 1.5-2.5 mm. long, each essentially reniform in outline or obliquely
lanceolate-ovate, deeply dentate or usually lacerate; mid-lobe linear-filiform, up
to 6 cm. long. Column with the clinandrium dentate or sublacerate.
Widely distributed and common from Mexico to Panama, through the West
Indies and in South America.
CHIRIQUÍ: around El Boquete, alt. 1000-1300 m., Pittier 3048. "PROVINCE OF VERA-
GUAS AND CHIRIQUÍ”: alt. 910-1210 m., Powell тоо, 3406.
A variable species but a distinctive one. Often in cultivation. А putative
hybrid between E. ciliare and E. Parkinsonianum is recorded.
(257)
[Vor. 33
326 ANNALS OF THE MISSOURI BOTANICAL GARDEN
19461
FLORA OF PANAMA (Orchidaceae) 327
ба. EPIDENDRUM CILIARE L. var. Oerstedii (Reichb. f.) L. Wms. comb. nov.
Epidendrum Oerstedii Reichb. f. in Bot. Zeit. 10:937. 1852.
Epidendrum costaricense Reichb. f. loc. cit.
Epidendrum Umlaufii Zahlbr. in Wien. Illustr. Gartenzeit. 18:209, /. 2. 1893.
Similar to E. ciliare except the lateral lobes of the lip reniform, entire or at
most crenulate, and the mid-lobe from filiform to broadly oblanceolate.
Costa Rica and Panama.
cHIRIQUÍ: Boquete, alt. 1150 m., Davidson 738; Palo Alto, alt. 1360 m., Powell 239,
3325, 3339.
Although this is obviously only a variation of E. ciliare, in which the lateral
lobes of the lip are entire (or nearly so) and in which the mid-lobe is usually
broader, its relationship seems not to have been suspected in recent years. Perhaps
this is because of the very evident but superficial character of lacerate lateral lobes
of the lip in E. ciliare and the entire (and thus less conspicuous) ones in the
variety.
7. EPIDENDRUM COCHLEATUM L. Sp. Pl, ed. 2:1351. 1763; Ames, Hub. &
Schweinf. Genus Epidendrum in U. S. & Mid. Am. 79. 1936.
Extremely variable epiphytic herbs up to about 5 dm. tall. Pseudobulbs 3-25
cm. long and 0.5—4 cm. broad, cylindric to fusiform to ovoid, round or flattened
in cross-section, bifoliate or rarely otherwise. Leaves 10-35 cm. long and 2-5 cm.
broad, elliptic to lanceolate to oblanceolate, acute. Inflorescence from shorter to
longer than the leaves, a simple to branched raceme, subtended by a sheath which
may be up to 15 cm. long. Flowers rather pretty, lip purple-marked to almost
black, sepals and petals white to green, occasionally purplish. Sepals similar,
2.5-7 cm. long and 0.2-0.7 cm. broad, linear to linear-lanceolate, acute, usually
strongly reflexed. Petals 1.5-5 cm. long and 0.15-0.4 cm. broad, narrowly linear,
occasionally broadest near the apex, acute. Lip 1—3 cm. long and 1—2.5 cm. broad,
short-unguiculate, claw adnate to the column for about half its length; lamina
cordate-orbicular, usually broader than long, obtuse or apiculate, cochleate, with
а more or less prominent callus thickening under the end of the column. Ovary
3-winged, prominently so in fruit.
Mexico to Venezuela and Colombia; the West Indies.
DARIÉN: Cana and vicinity, Williams 974.
A common and variable species, curiously known by only one collection in
Panama.
8. EPIDENDRUM CONDYLOCHILUM Lehm. & Krinzlin in Engler’s Bot. Jahrb.
26:459. 1899.
ое tessellatum Batem. іп Bot. Кер. 24: Misc. р. 7. 1838, поп Roxb.; Hook. in
Bot. Mag. 65: t. 8.
a Deamii Schltr. in Beih. Bot. Centralbl. 36, Abt. 2:402. 1918; Ames, Hub.
& Schweinf. Genus Epidendrum іп U. S. & Mid. Am. 88. 1936.
Encyclia аярга Schltr. TA cit. 474, sphalm.
(259)
[Vor. 33
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Repent or caespitose epiphytic herbs up to about 4 dm. tall. Pseudobulbs 2-15
cm. long and 0.3-1.5 cm. in diameter, cylindric to usually fusiform or narrowly
ovoid, bi- to trifoliate. Leaves 6—25 cm. long and 0.5—2.5 cm. broad, linear to
elliptic, acute or obtuse, usually erect. Inflorescence usually shorter than the leaves,
simple or rarely branched, few- to many-flowered. Flowers small, dull-colored,
variable. Dorsal вера! 8-12 mm. long and 1.5-3 mm. broad, linear to linear-
lanceolate, acute. Lateral sepals 8—13 mm. long and 2—3 mm. broad, lanceolate
to lanceolate-ovate, acute. Petals 8-10 mm. long and 1.5-4 mm. broad, linear-
oblanceolate to oblanceolate, obtuse. Lip 8—12 mm. long and 4—6 mm. broad,
obovate, obovate-oblong to oval in general outline, distinctly 3-lobed to almost
simple, free from the column, provided with a long low puberulent callus extend-
ing from the base to about the middle of the lip, and mammillate calluses from
the middle toward the apex, these last often arranged in 3 rows; lateral lobes from
very small to quite prominent and explanate, obtuse; mid-lobe with the margins
undulate. Ovary winged.
Mexico, Guatemala, Honduras, Costa Rica, Panama, Venezuela and Colombia.
CHIRIQUÍ: “Chiriquí”, alt. 1050-1450 m., Powell 93, 05, 3345, 3350, 3351, 3350.
This species has gone under the name of Epidendrum Deamii Schltr. for many
years. It is perhaps not distinct from E. lividum Lindl., being separated by the
presence of more or less distinct lateral lobes on the lip. The degree of lip loba-
tion is of dubious value, for several specimens show almost entire lip, including
typical material of E. condylochilum.
9. EPIDENDRUM CRASSILABIUM Poepp. & Endl. Nov. Gen. & Sp. РІ. 2:1, £. 102.
1838.
Epidendrum variegatum Hook. in Bot. Mag. 59: #. 3151.
Epidendrum baculibulbum Schltr. in Fedde Rep. Sp. Nov. Већ. 19:116. 1923.
Erect repent or caespitose epiphytes up to about 70 cm. tall. Pseudobulbs
15-35 cm. long and 0.5-1.5 cm. in diameter, cylindric (often thickened at the
base) to fusiform, bearing 2-4 widely separated leaves. Leaves 14-40 cm. long
and 2.5-4 cm. broad, elliptic to oblong-lanceolate, erect or spreading. Inflores-
cence a simple raceme either shorter or longer than the subtending leaf. Sepals
similar, 10-13 mm. long and 0.3-0.6 mm. broad, oblong to ovate to obovate,
obtuse. Petals 7-13 mm. long and 2.5-6 mm. broad, oblanceolate to oblong-ob-
lanceolate, obtuse, oblique. Lip 7-8 mm. long and 4-6 mm. broad, short-unguic-
ulate, only slightly longer than the column; claw short, adnate to the column at
the base; lamina orbicular to ovate, acute or obtuse, provided with a longitudinal
callus at the base which is usually a single raised ridge and at the apex divided
into two Jamellae extending almost to the apex of the lip.
Costa Rica, Panama, the West Indies and to Brazil.
cocLÉ: El Valle de Antón, Allen 2205, Woodson & Schery 204.
10. EPIDENDRUM FRAGRANS Sw. Nov. Gen. & Sp. Prodr. 123. 1788.
Epidendrum fragrans Sw. var. pachypus Schltr. in Fedde Rep. Sp. Nov. Beih. 17:32. 1922.
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19461 B
FLORA OF PANAMA (Orchidaceae) 329
Repent or caespitose epiphytic herbs up to about 4 dm. tall. Pseudobulbs 4—11
cm. long and 0.5-3 cm. in diameter, variable, cylindric, cylindric-fusiform to
ovoid, occasionally ancipitous, unifoliate. Leaves 8-30 cm. long and 1.5-5 cm.
broad, ligulate to elliptic, obtuse, coriaceous. Inflorescence to 15 cm. long, shorter
than the leaves, few- to several-flowered; sheaths up to 6 cm. long, chartaceous.
Flowers greenish to yellowish, the lip usually lined with purple along the veins,
sepals and petals rarely maculate. Sepals similar, 1.5—3.5 cm. long and 0.3—0.5
cm. broad, lanceolate or linear-lanceolate, acute or acuminate. Petals 2-3 cm.
long and 0.5—1 cm. broad, broadly lanceolate to oblanceolate or obovate, acute or
usually acuminate, unguiculate at the base. Lip 1.5-3 cm. long and 1-1.5 cm.
broad, somewhat cochleate, base unguiculate and shortly adnate to the column;
lamina ovate to ovate-subcordate, acuminate, provided with an inconspicuous
callus at the base forming an inverted Y or with two small parallel calluses.
Mexico, Guatemala, Costa Rica, Panama, the West Indies and in South America
to Brazil and Peru.
ZONE: Maxon 4913; San Juan and Frijoles, Powell 30, 43, 3355, 3370, 3371,
39$. E 3428, 3448, 3474; Barro Colorado Island, Shattuck 551. cocré: El Valle
de Antón, Allen 1678.
11. EPIDENDRUM IONOPHLEBIUM Reichb. f. Beitr. Orch. Centr.-Am. 103. 1866.
Epidendrum pachycarpum Schltr. in Fedde Rep. Sp. Nov. 3:109. 1906.
Epidendrum Hoffmannii Schltr. loc. cit. 16:444. 1920.
Repent or caespitose epiphytic herbs up to about 4 dm. tall. Pseudobulbs 3-9
cm. long and 1.5—3 cm. broad, ovoid, rarely fusiform, bifoliate or rarely trifoliate.
Leaves 10—35 cm. long and 1.5—4 cm. broad, ligulate or elliptic, obtuse. Inflores-
cence up to 15 cm. long, shorter than the leaves, few-to several-flowered. Flowers
greenish to white with the lip more or less prominently lined with purple. Sepals
similar, 1.4—3.5 cm. long and 0.3—0.8 cm. broad, lanceolate-ligulate to lanceolate,
acute, the laterals somewhat oblique. Petals 1.2—2.4 cm. long and 0.4-0.9 cm.
broad, elliptic-oblanceolate to oblanceolate, acute or acuminate. Lip 1.2-2.2 cm.
long and 0.8—1.8 cm. broad, unguiculate; claw short, adnate to the column;
lamina ovate to orbicular, obtuse or apiculate, cochleate, provided with an oblong-
pandurate to quadrate, obscurely puberulent callus at the base under the apex of
the column. Ovary angled, when mature alate.
In all Central American countries from Mexico to Panama and in Venezuela.
CANAL ZONE: near Frijoles, Maxon gu Culebra, alt. 50— zs m., Piftier 3301; бап
Juan, Powell 187, 3182. CHRIQUI: uete District, alt. 00 m., Davidson 706;
“Chiriqui”, Powell 151, 187, 310, 3226, 724 3565. VERAGUAS: +“ ELI yo 300
m., Powell 384.
12. EPIDENDRUM ONcIDIOIDES Lindl. var. RAMONENSE (Reichb. f.) Ames, Hub.
& Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:103. 1935. ‘
Epidendrum ramonense Reichb. f. Beitr. Orch. Centr.-Am. 81
1866.
Encyclia ramonensis Schltr. in Beih. Bot. Centralbl. 36, Abt. 2: 473. 1918.
Encyclia Powellii Schltr. in Fedde Rep. Sp. Nov. Beih. 17:46. 1922.
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[Vor. 33
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Large caespitose epiphytic herbs up to about 1 m. tall. Rhizome short. Pseudo-
bulbs up to about 10 cm. long, cylindric to usually ovoid, 2- to 4-foliate. Leaves
10—60 cm. long and 1—2.5 cm. broad, ligulate or ensiform, acute or obtuse. In-
florescence paniculate, exceeding the leaves, few- to many-flowered, simple or
compound. Dorsal sepal 14-18 mm. long and 3.5-5 mm. broad, elliptic-lanceo-
late to oblong-lanceolate, acute or acuminate. Lateral sepals similar to the dorsal
except somewhat arcuate. Petals 14—17 mm. long and 3—5.5 mm. broad, nar-
rowly oblanceolate, acute or acuminate. Lip 12-15 mm. long; lateral lobes oblong,
triangular-oblong, oblong-ovate or subquadrate-oblong, obtuse to truncate, arcu-
ate; mid-lobe 7-10 mm. long and 6—8.5 mm. broad, ovate to suborbicular, obtuse
to subacuminate; disc with 2 longitudinal fleshy ridges which form a fovea on
the isthmus, extending onto the mid-lobe as thickened nerves. Column exauricu-
late, with terminal angles or with dentiform auricles.
Honduras, Costa Rica and Panama.
ша “Chiriquí”, alt. 1360-1510 m., Powell $3. cocrf: El Valle, alt. 1060 m.,
Bouché 10
For all practical purposes this and the two following varieties might well be
referred to Е. oncidioides Lindl.
12a. EPripENDRUM ONCIDIOIDEs Lindl. var. GRaviDUM (Lindl) Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:104. 1935
Epidendrum gravidum Lindl. in Jour. Hort. Soc. 4:114. 1849; Ames, Sched. Orch. 4:42,
А 1
Eneyelia grevide Schltr. in Beih. Bot. Centralbl. ЖА E 2:472. 1918.
Epidendrum alanjense Ames, Sched. Orch. 1:13.
Encyclia Hunteriana Schltr. in P fe. Sp. Nov. "Beh. 17:46. 1922.
Epidendrum amandum Ames, Sched. Orch. 4:36. 1923
Epidendrum peraltense Ames, loc. cit. 46
Similar to the preceding variety into which it intergrades, usually somewhat
smaller іп all parts. Mid-lobe of lip triangular-lanceolate to oblong. Veins of the
mid-lobe verruculose.
Mexico, Honduras, Nicaragua, Costa Rica and Panama.
CHIRIQUÍ: Alanje, Pittier s. п.; David, Powell 84, 3261, 3264, 3530, 3545. PANAMA:
San Juan Range, Powell 271, 3470, 3485.
12b. EPIDENDRUM ONCIDIOIDES Lindl. var. MoorEANUM (Rolfe) Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:106. 1935.
Epidendrum Mooreanum Rolfe in ad Bull. 1891:199. 189
Encyclia Tonduziana Schltr, in е Кер. Sp. Nov. Beih. s 132. 1923.
Encyclia Brenesii Schltr., loc. cit
Much like the Nc variety. Lateral lobes of the lip spatulate or oblong-
spatulate, veins verruculose.
Costa Rica and Panama.
CHIRIQUÍ: Volcán de Chiriquí, alt. 2120 m., Davidson 870; Lino Hill, alt. 1360 m.,
Powell 292.
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19461
FLORA OF PANAMA (Orchidaceae)
331
Fig. 121. Epidendrum oncidioides var. gravidum
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[Vor. 33
332 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
13. Еріремркі/м OTTONIs Reichb. f. in Hamb. Gartenzeit. 14:213. 1858.
Nidema ottonis Britt. & Millsp. Bahama Fl. 94.
Nidema € (Lindl.) Schltr. var. hint “беріс in Fedde Rep. Sp. Nov. Beih.
Small, repent or caespitose epiphytic herbs up to about 25 cm. tall. Rhizome
creeping. Pseudobulbs up to about 4 cm. long, fusiform, short-stipitate, uni-
foliate. Leaves 5-19 cm. long and 0.5-0.9 cm. broad, linear-ligulate, obtuse or
acute, coriaceous. Inflorescence a short, few-flowered raceme; flowers small,
white or yellowish; bracts mostly 1-2 cm. long, lanceolate, acuminate, paleaceous.
Dorsal sepal 7-9 mm. long and 2.5-3 mm. broad, elliptic-lanceolate, acute.
Lateral sepals 7-9 mm. long and 2-2.5 mm. broad, lanceolate, acuminate, more
or less arcuate. Petals 6-7 mm. long and 1.5-2 mm. broad, elliptic, elliptic-
lanceolate to oblanceolate, acute or acuminate, arcuate. Lip 6.5-7 mm. long and
about 1.5 mm. broad, narrowly oblong, acute, subpandurate, fleshy, canaliculate,
keeled toward the apex below. Column monandrous or triandrous, if triandrous
then the lateral stamens usually imperfect.
Panama, the West Indies, Venezuela, Colombia and Peru.
isa vicinity of Bejuco, Allen 081; east of Panamá City, Powell 131, 3440;
Tecümen, Standley 26611; between Las Sabanas and Matías Hernández,
Standley 31888, 31945.
Resembles Epidendrum Boothii (Lindl.) L. Wms. very much.
14. EPIDENDRUM PARKINSONIANUM Hook. in Bot. Mag. 67: f. 3778. 1840.
Epidendrum aloifolium Batem. Orch. Mex. & Guat. t. 25. 1840, поп Г.
ее falcatum Lindl. var. a Schltr. in Fedde Rep. Sp. Nov. Beih. 19:37.
Repent epiphytic herbs up to about 6 dm. long. Stems creeping, branched,
cylindric or somewhat swollen. Leaves 15-50 cm. long and 1-4.5 cm. broad,
ligulate to lanceolate-ligulate, acute, very thick, more or less flaccid, occasionally
pustulate, usually pendent. Inflorescence a very short, few-flowered raceme;
bracts short, ovate; pedunculate ovaries up to about 16 cm. long. Sepals 5.5-9
cm. long and 0.6-1.2 cm. broad, linear-lanceolate to linear, acute or acuminate,
spreading. Petals 5-8 cm. long, similar to the sepals. Lip 6—8.5 cm. long (from
the base of the column) and 3-4 cm. broad, 3-lobed, the base adnate to the col-
umn; lateral lobes 2.5-4 cm. long, broadly lunate to oblong-lunate; mid-lobe
3-6 mm. long, linear.
Mexico, Guatemala, Honduras, Costa Rica and Panama.
cHIRIQUÍ: Pueblo del Volcan, alt. 1800-1960 m., ем Upper Rio Chiriqui Viejo,
vicinity of Monte Lirio, alt. 1300-1900 m., Seibert 214, 217.
A curious species with large and handsome flowers. It is allied to E. ciliare
and E. pugioniforme but is easily distinguished from them by the flaccid, dif-
ferently shaped leaves.
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1946]
FLORA OF PANAMA (Orchidaceae) 331
15. EPIDENDRUM PRISMATOCARPUM Reichb. f. in Bot. Zeit. 10:729. 1852;
Reichb. f. Xenia Orch. 2:83, 2. 123. 1862; Hook. in Bot. Mag. 88: /. 5336.
1862.
Stout repent or caespitose epiphytic herbs up to about 5 dm. tall. Pseudobulbs
up to about 15 cm. long and 5 cm. in diameter, narrowly ovoid, often attenuated
above, bi-trifoliate, the leaves somewhat separated on the apex of the bulb.
Leaves 12-33 cm. long and 2-6 cm. broad, variable, from ligulate and oblanceolate-
oblong to narrowly ovate, obtuse, coriaceous. Inflorescence an erect, few- to
many-flowered raceme. Dorsal sepals 22-44 mm. long and 3-5 mm. broad, nar-
rowly lanceolate to narrowly oblanceolate, acute, arcuate. Petals 20-28 mm.
long and 3-5 mm. broad, elliptic-linear to lanceolate, acute, arcuate. Lip 18-25
mm. long and 7-9 mm. broad, 3-lobed, free from the column nearly to the base,
subsagittate, with a central callus from the base nearly to the apex; middle lobe
large, trulliform, acuminate.
Costa Rica and Panama.
CHIRIQUÍ: "Province of Chiriquí”, alt. 1200-1360 m., Powell 00, ТОТ, 3375, 3388,
3391, 3402; “Chiriquí”, Warscewicz; near El Volcán, White 203; Casita Alta to Cerro
Copete, alt. 2300-3300 m., Woodson & Schery 375.
Allied to E. brassavolae. One of the commonest of the Epidendrums in culti-
vation in greenhouses. It is a good bloomer and grows well, the flowers occasionally
being sold as cut flowers in the northern markets.
16. EPIDENDRUM PYGMAEUM Hook. in Bot. Mag. 60: /. 3233. 1833; Hook. Jour.
Bot. 1:49, Ё. 118. 1834; Ames, Hub. & Schweinf. Genus Epidendrum in
U. S. & Mid. Am. 160. 1936.
Hormidium pseudopygmaeum Finet in Bull. Herb. Boiss. 7:121, /. 3. 1899.
Small repent epiphytic herbs from a creeping rhizome. Pseudobulbs scattered
along the rhizome, 2-10 cm. long, cylindric to fusiform, bifoliate. Leaves 2-15
cm. long and 0.7-2 cm. broad, narrowly elliptic со oblong-oval, obtuse to acute.
Inflorescence sessile in the axils of the leaves, a short few- to several-flowered
raceme. Flowers small, inconspicuous. Dorsal sepal 5-10 mm. long and 1.5-3
mm. broad, lanceolate, acuminate. Lateral sepals 6-12 mm. long and 2-4 mm.
broad, lanceolate, acuminate, keeled dorsally toward the apex, shortly connate at
the base and lightly adnate to the base of the column. Petals 4-8 mm. long and
0.5-1 mm. broad, linear, acute. Lip 3-8 mm. long and 3-7 mm. broad, clawed
at the base and shortly adnate to the base of the column; lamina 3-lobed or 3-
lobulate; lateral lobes explanate or erect, ovate to orbicular; mid-lobe small,
apiculate. Ovary winged.
Florida and Mexico to Panama, the West Indies and to Brazil and Bolivia.
CHIRIQUÍ: vicinity of Monte Lirio, Seibert 230. сосіЁ: El Valle, Seibert 426; near
El Volcán, White 205.
A variable and widely distributed species. There seem to be two forms of
this,—one compact and small, the other rather loose and comparatively large. "Тһе
second form is Finet's Hormidium pseudopygmaeum.
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[Vor. 33
334 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 122. Epidendrum pygmacum
(266)
19461
FLORA OF PANAMA (Orchidaceae) 335
17. EPIDENDRUM RoussEAUAE Schltr. in Већ. Bot. Centralbl. 36, Abt. 2:407.
1918; Ames, Hub. & Schweinf. Genus Epidendrum in U. S. & Mid. Am.
170. 1936.
Epiphytic herbs similar to E. ramosum Jacq. Leaves ligulate, about 7 cm.
long and 1.2 cm. broad, obtuse. Inflorescence a lax several-flowered raceme up to
about 10 cm. long. Flowers fleshy, greenish. Sepals about 1.2 cm. long, lanceolate-
ligulate, acute, fleshy. Petals about 1.2 cm. long, linear or linear-oblanceolate.
Lip unguiculate; claw adnate to the column; lamina about 7 mm. long and as
broad, trilobate, provided with 2 parallel calluses extending from the base up to
the base of the mid-lobe, base deeply cordate; lateral lobes spreading, dolabriform,
obtuse; terminal lobe about 4 mm. long and 3.5 mm. broad, extended into a sub-
orbicular blade from a broadly ligulate claw, obtuse, apiculate.
Panama.
CANAL ZONE: Mrs. Rousseau, s. п.
The description is taken from the original publication. Specimens are missing
from the Ames Herbarium but Ames, Hubbard & Schweinfurth have cited the
following collections from Panama: Powell 112, 3303; Münch 27164; Shattuck
347.
Curiously enough, Schlechter contrasts this species with Epidendrum ramosum
Jacq., which belongs to the section EuEPIDENDRUM, while Ames, Hubbard апай
Schweinfurth have placed it in the section ENcYcLIUM.
18. EPIDENDRUM SPONDIADUM Reichb. f. in Bot. Zeit. 10:731. 1852; Hook. f.
in Bot. Mag. 119: #. 7273. 1893.
Epidendrum platycardium Schltr. in Fedde Rep. Sp. Nov. Beih. 17:36. 1922.
Repent or caespitose epiphytic herbs up to about 35 cm. tall. Pseudobulbs
3-12 cm. long and 0.6—1.3 cm. broad, cylindric to narrowly ovoid, unifoliate.
Leaves 10—28 cm. long and 2.5—5 cm. broad, ensiform to elliptic, obtuse. In-
florescence shorter than the leaves, few-flowered; sheath about 2.5 cm. long.
Flowers with sepals and petals greenish to white and the lip rose to purple. Sepals
13-20 mm. long and 3-6 mm. broad, lanceolate, acuminate or acute, the laterals
oblique. Petals 12-17 mm. long and 3.5—6.5 mm. broad, lanceolate to lanceolate-
ovate, acute or acuminate. Lip 12-15 mm. long and 10-12 mm. broad, short-
unguiculate; claw adnate to the column; lamina suborbicular-reniform to cordate,
apiculate, provided with a small subfoveate callus under the apex of the column
which usually has 3 thickened nerves extending outward.
Costa Rica, Panama and possibly Jamaica.
CHIRIQUÍ: Palo Alto Hill, alt. 1500 m., Powell 141.
Very closely allied to Epidendrum fragrans Sw. and perhaps only a variation
of that.
!
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[Vor. 33
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN
19. EPIDENDRUM STAMFORDIANUM Batem. Orch. Mex. & Guat. /. 11. 1838;
Hook. in Bot. Mag. 80: £. 4750. 1854.
Epidendrum Cycnostalix Reichb. f. in Bot. Zeit. 10:731. 1852.
Caespitose epiphytic herbs up to about 50 cm. tall. Stems pseudobulbose,
8-27 cm. long and about 0.5-2 cm. in diameter, cylindric-fusiform to fusiform,
bearing 1—4 leaves at or near the apex. Leaves 10—27 cm. long and 3—7 cm. broad,
variable, ligulate, elliptic, oblong-lanceolate, or oblanceolate to oval, obtuse,
coriaceous. Inflorescence up to about 60 cm. long, spreading or pendent, simple
or paniculate, usually many-flowered, lateral from a short fertile shoot, or rarely
terminal on the pseudobulb. Flowers rather pretty, fair-sized. Dorsal сера!
15-18 mm. long and 4—6 mm. broad, elliptic or lanceolate, acute or acuminate.
Lateral sepals 14-18 mm. long and 4-6 mm. broad, lanceolate or oblong-lanceolate,
strongly oblique, acute or acuminate. Petals 14-18 mm. long and 1.5—4.5 mm.
broad, linear, elliptic, linear-oblong to lanceolate, acute. Lip unguiculate; claw
adnate to the column; lamina 11-16 mm. long and 13-20 mm. broad, 3-lobed;
lateral lobes large, spreading, oblong, oblique, obtuse, entire or crenulate; mid-
lobe bilobulate, retuse, transversely oblong in outline, provided with a narrow,
often cuneate claw at the base, entire or usually crenulate or serrulate; disc pro-
vided with a pair of parallel, short, lamellate calluses, or a single deeply sulcate
callus at the base near the apex of the column, and with the median nerve thickened
or raised into a low lamella.
Mexico to Panama, Venezuela and Colombia.
CHIRIQUÍ: "Chiriqui", Warscewicz.
No recent Panamanian collections seen. Тһе species grows well in cultivation.
20. EPIDENDRUM STANGEANUM Reichb. f. in Gard. Chron. n.s. 15:462. 1881.
Epidendrum glandulosum Ames, Sched. Orch. 7:5, #. 20. 1924.
Small repent or caespitose epiphytic herbs from a slender creeping rhizome, up
to 2 dm. tall. Pseudobulbs 1.5—5 cm. long and 0.1—0.5 cm. in diameter, cylindric
to narrowly fusiform, covered with marcescent sheaths, becoming naked with
age, unifoliate. Leaves 4-12 cm. long and 1-3 mm. broad (or in diameter),
linear and flattened or appearing terete due to their inrolled margins. Inflorescence
a few- to several-flowered raceme, shorter than the subtending leaf; flowers small
and inconspicuous. Dorsal sepals 5-6 mm. long and 1.5-2 mm. broad, lanceolate,
acute, sparsely glandular dorsally. Lateral sepals 6-7 mm. long and 2-3 mm.
broad, oblique, lanceolate, acute or acuminate, sparsely glandular dorsally. Petals
4.5-5.5 mm. long and 0.5-0.7 mm. broad, linear, acute. Lip 5-6 mm. long and
3—3.5 mm. broad, unguiculate; claw short, adnate to the column; lamina deltoid
to ovate-lanceolate, apiculate, slightly cochleate, without calluses.
Costa Rica and Panama.
CANAL ZONE: on trail from Colón, Powell 338; (record without data), Shattuck 454.
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19461
FLORA OF PANAMA (Orchidaceae) 337
21. EPIDENDRUM VARICOSUM Batem. in Bot. Reg. 24: Misc. p. 30. 1838; Lindl.
Folia Orch. Epid. 23. 1853; Reichb. f. Xenia Orch. 1:163, Ё. 50. 1856.
mM chiriquense Reichb. f. in Bot. Zeit. 10:730. 1852, Xenia Orch. 1:164, Ё. 67.
bed chiriquensis Schltr. in Beih. Bot. Centralbl. 36, Abt. 2:472. 1918.
Small to large, repent or caespitose, epiphytic herbs up to about 1 m. tall.
Pseudobulbs up to 25 cm. long, usually ovoid at the base and prolonged into a
long slender neck above, 3- to 4-leaved. Leaves 5-35 cm. long and 1.5-5 cm.
broad, linear-lanceolate to elliptic, acute or acuminate. Inflorescence simple or
branched, shorter or longer than the leaves. Dorsal sepal 8-18 mm. long and 3-5
mm. broad, oblong to oblong-lanceolate, acute or obtuse. Lateral sepals 8—16 mm.
long and 3—5 mm. broad, oblong to oblong-ovate, oblique, obtuse or acute. Petals
8-14 mm. long and 2-5 mm. broad, oblanceolate to narrowly obovate, obtuse,
usually oblique. Lip 9-15 mm. long and 6-13 mm. broad, cuneate, narrowed to the
base and shortly connate with the base of the column, 3-lobed; lateral lobes arising
near the middle of the lip, usually small and explanate. Mid-lobe relatively large,
bifid, and the lobules more or less explanate and flabellate; disc with a large cen-
tral callus and with mammillate or varicose processes or lamellae from the middle
extending out to the lobules of the mid-lobe. Оуагу angled or narrowly winged
in fruit.
Mexico, Guatemala, Salvador, Honduras, Costa Rica and Panama.
CHIRIQUÍ: "Chiriqui", Warscewicz.
Reichenbach first gave the origin of Epidendrum chiriquense as "Costa Rica
und Chiriqui" and later as "Chiriqui: v. Warscewicz". The species has not been
seen from Costa Rica or Panama since and the basis of E. varicosum in Panama is
based on his record.
22. EPIDENDRUM VOLUTUM Lindl. & Paxt. in Paxton's Flow. Gard. 2:151, fig.
215. 1851-52.
Epidendrum Radlkoferianum Schltr. іп Fedde Rep. Sp. Хоу. 17:142. 1921.
Erect epiphytic herbs 18-25 cm. tall. Pseudobulbs when young 5-10 cm.
long and 0.5-0.6 cm. in diameter, cylindric. Leaves 6-8 cm. long and 0.8—1 cm.
broad, ligulate, acute, coriaceous. Inflorescence a lax raceme of 6-10 flowers,
about 13 cm. long; peduncle 2-4 cm. long. Sepals about 1.8 cm. long and 0.3
cm. broad, narrowly oblong-ligulate, acuminate, the laterals oblique. Petals about
as long as the sepals, linear, narrow and oblique, acuminate. Lip unguiculate;
claw adnate to the column; lamina about 9 mm. long and 11 mm. broad, ob-
scurely 3-lobed, subreniform-cordate, apiculate, apex triangular, apical part sub-
crenate and undulate; disc provided with 2 short, obtuse, parallel, lamellate
calluses at the base and 3 approximate linear calluses extending up to about the
middle.
Panama.
(269)
[Vor. 33
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CHIRIQUÍ: "im Veragua- und Chiriqui-Distrikt”, alt. 1000-1200 m., Powell $9;
"Provinz Chiriqui”, Wagner.
The description is taken from Schlechter's description of E. Radlkoferiamum.
The specimens indicated above are those cited by Schlechter.
Sect. II. BARKERIA
23. EPIDENDRUM STENOPETALUM Hook. іп Bot. Mag. 62: #. 3410. 1835; Ames,
Hub. & Schweinf. Genus Epidendrum in U. S. & Mid. Am. 179. 1936.
Dimerandra stenopetala Schltr. in Fedde Rep. Sp. Nov. Beih. 17:44. 1922.
Slender, caespitose, epiphytic herbs up to about 4 dm. tall. Stems terete,
cylindric, cane-like, often somewhat flexuose, leafy. Leaves 3.5—14 cm. long and
0.4—1 cm. broad, linear or ligulate to linear-oblong or linear-lanceolate, obtuse,
unequally bilobulate at the apex, scattered along the stem, one at each node. In-
florescence a short, few-flowered raceme. Dorsal вера! 10-13 mm. long and
3.5—4.5 mm. broad, lanceolate or elliptic-lanceolate, acute or acuminate. Lateral
sepals 10-13 mm. long and 3.5—4.5 mm. broad, lanceolate to lanceolate-ovate,
acute or acuminate, somewhat oblique. Petals 11-13 mm. long and 6—6.5 mm.
broad, obovate to subrhombic, acute or acuminate. Lip 11-13 mm. long and
8—9.5 mm. broad, obovate to obovate-flabellate, truncate; disc obscurely lined.
Column short, the clinandrium with 2 large lateral lobes.
Mexico, Guatemala, British Honduras, Honduras, Costa Rica, Panama, the
West Indies and South America at low elevations.
BOCAS DEL TORO: Water Valley, Wedel 765. CANAL томе: drowned forest near
Vigía - San Juan on Río Pequeni, Dodge, Steyermark t$ Allen 16586; hills near Panamá
City, Powell 17, 3451; Las Cruces trail, Standley 20101; between France Field and
Eu "Standley 30448. PANAMA: swamp east of Río Tecümen, Standley 26610.
This species is placed in section BARKERIA by Ames, Hubbard & Schweinfurth
(loc. cit.), but it has little relationship to the other species of the section; it
would perhaps be best placed in a section by itself.
III. Section EUEPIDENDRUM
24. EPIDENDRUM ALLENI Г. Wms. in Ann. Missouri Bot. Gard. 28:418, £. 22,
figs. 3-5. 1941.
Erect epiphytic herbs up to about 3 dm. tall. Stems ancipitous, covered with
the bases of the 3 to 6 leaves. Leaves 4-12 cm. long, 1.2-3 cm. broad, elliptic to
lanceolate, acute or short-acuminate. Peduncle about 10—15 cm. long, ancipitous,
bialate, the wings prominent at the base but disappearing at about the middle of
the peduncle. Inflorescence short, the rachis covered with the scarious, imbricated
bracts; bracts up to about 1 cm. long. Dorsal вера! about 10 mm. long and 4
mm. broad, lanceolate, obtuse, 5- to 7-nerved, with a short dorsal apicule near
the apex. Lateral sepals similar to the dorsal sepal except somewhat arcuate.
Petals about 10 mm. long and 0.75 mm. broad, linear, 1-nerved. Lip unguiculate;
the lamina about 10 mm. long and 12 mm. broad, subreniform, subentire or ob-
(270)
1946)
FLORA OF PANAMA (Orchidaceae) 339
E. fni
Fig. 123. Epidendrum ellipsophyllum and E. Allenii
(271)
[Vor. 33
340 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 124. Epidendrum anceps
(272)
19461
FLORA OF PANAMA (Orchidaceae) 341
scurely 3-lobed, the terminal lobe inconspicuous and slightly retuse, with 2 ap-
proximate submammillar calluses at the base. Column of the section, about 7
mm. long.
Panama.
COCLÉ: hills north of El Valle de Antón, alt. 1000 m., Allen 2203, 2310.
25. EPIDENDRUM ANCEPS Jacq. Select. Stirp. Am. 224, 2. 138. 1763; Ames,
Hub. & Schweinf. Genus Epidendrum in U. S. & Mid. Am. 56. 1936.
Epidendrum musciferum Lindl. in Hook. Jour. Bot. 1:6. 1834.
Epiphytic herbs up to about 1 m. tall. Stems slender to rather coarse, covered
with the amplexicaul leaf-sheaths, usually ancipitous. Leaves 4—25 cm. long and
1—5 cm. broad, ligulate to elliptic-lanceolate, acute or obtuse, those at the middle
of the stem or above usually largest, the lower ones reduced. Inflorescence usually
long-pedunculate (up to 45 cm.), simple or branched, racemose or subcapitate;
peduncle ancipitous, covered with scarious sheaths. Dorsal sepal 5-10 mm. long
and 2—3.5 mm. broad, elliptic to elliptic-ovate, acute or obtuse. Lateral sepals
6-10 mm. long and 2-4 mm. broad, oblong-lanceolate to ovate, obtuse or acute,
oblique, semi-cochleate. Petals 4-9 mm. long and 0.5-1.5 mm. broad, filiform
to linear-oblanceolate. Lip unguiculate; claw adnate to the column; lamina 3-6
mm. long and 4—7.5 mm. broad, reniform to cordate-suborbicular, 3-lobed; the
lateral lobes rounded, fleshy; terminal lobe subquadrate to oblong, entire, retuse
or doubly retuse; disc fleshy, usually with a low longitudinal callus from base
to apex.
Florida and Mexico through Central America and the West Indies to Brazil
and Peru.
BOCAS DEL TORO: Old Bank Island, Wedel 2081; Bastimentos Island, Wedel 2021.
CANAL ZONE: Gatün Lake, Frijoles, San Juan, Fort Sherman, Powell 40, 41, 3349, 3366,
3367; Barro Colorado Island, Woodworth & Vestal 66A. снікшюсі: Lino Hill, alt. 1200—
1500 m., Powell 200. PANAMÁ: Cerro Campana, alt. 1000 m., Allen 2452; Panamá City,
Kieswetter s. n.
26. EPIDENDRUM BISULCATUM Ames, Sched. Orch. 5:24, Ё. 5. 1923.
Small, branched, epiphytic herbs up to about 50 cm. long. Stems slender,
much branched, covered with the scarious leaf-sheaths when young, becoming
naked with age. Leaves 4—10 cm. long and 0.6—2 cm. broad, elliptic to lanceolate
to oblanceolate, acute. Inflorescence shorter than the leaves, 1- to few-flowered,
borne on a short winged peduncle. Flowers greenish, the lip with a purple cast.
Dorsal sepal 7-10 mm. long and 3—6 mm. broad, oval to oblong-ligulate, acutish.
Lateral sepals 10-12 mm. long and 3-6 mm. broad, oblong to oblong-oval, ob-
lique, acute. Petals 7-10 mm. long and 1.5-2 mm. broad, linear, obtuse. Lip
unguiculate; the claw adnate to the column; lamina 7-10 mm. long and 9-13
mm. broad, suborbicular-cordate, crenulate, obscurely emarginate, fleshy, provided
with 2 (or 3) submammillate calluses at the base which are prolonged into low
(273)
[Vor. 33
342 ANNALS OF THE MISSOURI BOTANICAL GARDEN
А
f @
кабель ano p
AU
COCU mn
оға "a % 7/ я
Ж
Fig. 125. Epidendrum bisulcatum
(274)
1946)
FLORA OF PANAMA (Orchidaceae) 343
fleshy keels and with a low keel passing between the mammillae.
Panama.
HIRIQUÍ: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 122, 322; Palo Alto
Hill, p 1800 m., Powell 280, 28
27. EPIDENDRUM CALIGARIUM Reichb. f. in Gard. Chron. 1869:1110. 1869;
Ames, Sched. Orch. 9:48, fig. 8. 1925.
Caespitose epiphytic herbs up to about 80 cm. long. Stems slender, strict,
covered with loosely appressed sheaths, the sheaths densely maculated with minute,
purple, papillose or tuberculate excrescences. Leaves 1.5—6 cm. long and 0.3—0.6
cm. broad, linear or linear-lanceolate, borne near the apex of the stem. Inflores-
сепсе exceeding the leaves, a densely flowered simple or branched raceme. Sepals
EPIDENDRUM
caligarium
Fig. 126. Epidendrum caligarium
5-6 mm. long and 2.5-3 mm. broad, oval, obtuse or obscurely apiculate. Petals
5-6 mm. long and about 1.5 mm. broad, elliptic-oblanceolate. Lip unguiculate;
the claw adnate to the column; lamina 6-8 mm. long and 8-10 mm. broad, more
or less orbicular in outline, 3-lobed, the lateral lobes deeply lacerate, often cut
nearly to the middle of the lip, each lobe often 4-partite, mid-lobe deeply retuse,
the lateral lobules denticulate or laciniate; disc provided with 2 small antrorse,
submammillate calluses at the base.
Known definitely only from Panama.
VERAGUAS: San Juan, alt. 360 m., Powell 380.
(275)
[Vor. 33
344 ANNALS OF THE MISSOURI BOTANICAL GARDEN
28. EPIDENDRUM Caro ni Schltr. in Fedde Rep. Sp. Nov. Вей. 19:35. 1923.
Erect, caespitose, epiphytic herbs up to about 40 cm. tall. Stems 3-12 cm.
long, slender, covered with the scarious leaf-sheaths, becoming naked with age,
bearing leaves toward the apex. Leaves 2.5-8 cm. long and 0.7-2 cm. broad,
elliptic-lanceolate or oblong. Inflorescence up to 30 cm. long, long-pedunculate,
a simple or branched raceme; peduncle ancipitous, covered with several equitant
scarious bracts. Flowers small, usually purplish with a yellow lip. Dorsal sepal
5-6 mm. long and 1.5—2 mm. broad, lanceolate to oblong, acute or obtuse.
Lateral sepals similar to the dorsal but oblique. Petals 3-5 mm. long and 1-1.8
mm. broad, linear to linear-oblong, slightly arcuate, obscurely denticulate. Lip
unguiculate; claw adnate to the column; lamina 4-5 mm. long and as broad,
ovate-cordate, crenulate, fleshy, the disc with a longitudinal callus well developed
and joining the two erect lateral lobes of the lip.
Mexico, Costa Rica and Panama.
cHIRIQUÍ: Boquete, alt. 1150 m., Davidson 1058.
29. EPIDENDRUM CENTRADENIA Reichb. f. in Walp. Ann. 6: index p. 1163. 1865.
Oerstedella centradenia Reichb. f. in Bot. Zeit. 10:932. 1852.
Epidendrum tenuiflorum Schltr. in Fedde Rep. Sp. Nov. 3:49. 1906.
Caespitose or spreading and proliferous epiphytic herbs up to 0.5 m. or perhaps
longer. Stems slender, covered with the amplexicaul, minutely papillose or tuber-
culate leaf-sheaths, becoming naked with age, simple or branched, bearing leaves
mainly toward the apex of the new growths. Leaves 2-9 cm. long and 0.2-0.8
cm. broad, linear to linear-lanceolate, acute. Inflorescence 1- to few-flowered; flow-
ers lilac-colored. Sepals 7-10 mm. long and 2-3 mm. broad, linear-lanceolate, acute.
Petals 7-10 mm. long and 1.5-2 mm. broad, linear-oblanceolate, obtuse, slightly
oblique. Lip 10-18 mm. long and 9—17 mm. broad, unguiculate; the narrow
claw adnate to the column; lamina 3-lobed, lateral lobes small, about 3 mm. long,
explanate, mid-lobe relatively large, deeply retuse and with the two large divari-
cate lobules cuneate to the junction with the lateral lobes; disc provided with a
small hood-shaped tridentate callus at the base of the lamina. Column with a
large hood-shaped or calyptriform clinandrium.
Costa Rica and Panama.
CHIRIQUÍ: "Province of Chiriquí”, alt. 1200 m., Powell 130.
30. EPIDENDRUM CENTROPETALUM Reichb. f. in Bot. Zeit. 10:732. 1852.
Oerstedella centropetala Reichb. f. loc. cit. 932.
Epidendrum aberrans Schltr. in Fedde Rep. Sp. Nov. 15:206. 1918.
Epidendrum leprosum Schltr. іп Fedde Rep. Sp. Nov. Beih. 19:38. 1923.
Caespitose or repent epiphytic herbs up to 65 cm. tall. Stems slender, covered
with the leaf-sheaths or naked with age, simple or branched, bearing leaves mainly
toward the apex; leaf-sheaths verrucose or tuberculate. Leaves 2.5-8 cm. long
and 0.3—1.5 cm. broad, linear-lanceolate to lanceolate, acute or obtuse. Inflores-
(276)
19461 А
FLORA OF PANAMA (Orchidaceae) 345
cence a simple few- to several-flowered raceme exceeding the leaves; flowers small,
rose to purple. Dorsal вера! 7-9 mm. long and 2-3 mm. broad, elliptic to ob-
lanceolate, obtuse or acute. Lateral sepals 7-10 mm. long and 2-3.5 mm. broad,
oblong or oblanceolate, acute, arcuate. Petals 6-9 mm. long and 1-2.3 mm.
broad, linear-oblanceolate, obtuse. Lip unguiculate, strongly 3-lobed; claw at
the base adnate to the column, free above, fleshy and provided with a raised sulcate
and retuse callus reaching to about the lateral lobes; lamina strongly 3-lobed, the
mid-lobe strongly bilobed, lateral lobes 2-4 mm. long, explanate, oblong, obtuse;
terminal lobe clawed and the two lobules explanate to recurved, the lobules oblong,
obtuse or acute. Column provided with а hood-shaped or calyptriform
clinandrium.
Guatemala, Honduras, Costa Rica and Panama.
CHIRIQUÍ: Chiriquí Volcano, Warscewicz.
31. EPIDENDRUM COCLÉENSE Ames, Hub. & Schweinf. in Bot. Mus. Leafl. Harv.
Univ. 4:6. 1936.
Scandent epiphytic herbs up to about 2.5 m. long. Stems simple or branched,
covered with the amplexicaul leaf-sheaths or naked with age. Leaves 4-10 cm.
long and 4-6 mm. broad, linear or ensiform, acute. Inflorescence short, 1- to 3-
flowered, shorter than the subtending leaves; sheath up to 15 cm. long, subimbri-
cated, scarious, cucullate, exceeding the pedicels of the flowers. Dorsal sepals
about 8 mm. long and 2.5 mm. broad, linear-oblong, obtuse. Lateral sepals about
10 mm. long and 3.5 mm. broad, lanceolate, acute, shortly alate-keeled dorsally
at the apex. Petals about 8 mm. long and 2 mm. broad, oblanceolate, acute. Lip
unguiculate, fleshy; claw adnate to the column; lamina about 6 mm. long and 4
mm. broad, oblong-ovate, simple, obtuse, provided with a small tridentate callus
under the apex of the column, the middle tooth of which is prolonged into a
longitudinal carina.
Panama.
cocLÉ: El Valle de Antón, alt. 500 m., Hunter & Allen 380.
32. EPIDENDRUM CONFERTUM Ames & Schweinf. Sched. Orch. 10:61. 1930.
Epidendrum prostratum Schltr. in Beih. Bot. Centralbl. 36, Abt. 2:407. 1918, non Cogn.
Branched or simple, prostrate epiphytic herbs up to 50 cm. long. Stems slen-
der, densely leaved, covered with the amplexicaul leaf-sheaths. Leaves 8-15 mm.
long and 3-6 mm. broad, oblong-ligulate to ovate, acute to aristate. Inflorescence
usually a single flower at the apex of the stem or branches. Sepals 8—9 mm. long
and 2-2.5 mm. broad, lanceolate-ligulate, acute. Petals about 8 mm. long and
1 mm. broad, linear, obtuse or acute, broadest near the apex. Lip unguiculate;
claw adnate to the column; lamina about 5 mm. long and as broad, orbicular-
cordate, slightly cochleate.
Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 246.
(277)
[Vor. 33
346 ANNALS OF THE MISSOURI BOTANICAL GARDEN
33. EPIDENDRUM CONGESTUM Rolfe in Kew Bull. 1913:29. 1913.
Epidendrum serruliferum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:44. 1923.
Small, densely caespitose herbs up to 7 cm. tall. Stems short, covered with the
broad equitant bases of the leaves. Inflorescence 1- to few-flowered, sessile in the
axils of the terminal leaves. Leaves 8-30 mm. long and 4-8 mm. broad, linear-
oblong to oblong-oval, obtuse, approximate, semi-equitant, spreading, fleshy.
Dorsal sepal 7-9 mm. long and 3—4 mm. broad, lanceolate, acute to acuminate.
Lateral sepals 8-10 mm. long and 2-3 mm. broad, lanceolate, cucullate, alate-
keeled dorsally and the keel denticulate, oblique, connate at the base with the claw
of the lip. Petals 7-8 mm. long and 2-2.5 mm. broad, lanceolate, acute, minutely
denticulate. Lip 7-10 mm. long, long-unguiculate; claw adnate to the column
but exceeding it slightly; lamina 4—5 mm. long and 3—4 mm. broad, lanceolate-
cordate to cordate, acute or acuminate, fleshy, lobes more or less erect. Clinan-
drium not fimbriate.
Costa Rica and Panama.
cocLÉ: vicinity of El Valle, alt. 600-1000 m., Allen 1247.
34. EPIDENDRUM CORIIFOLIUM Lindl. in Jour. Hort. Soc. Lond. 6:218, fig. 1851;
Summerhayes in Bot. Mag. 160: /. 0477. 1937.
Epidendrum magnibracteatum Ames, Sched. Orch. 6. 1922, поп Kranzlin
Epidendrum coriifolium Lindl. var. purpurascens Scher in Fedde Rep. Sp. Nov. Beih.
17:31. 1922
Epidendrum fuscopurpureum Schltr. loc. cit. 32.
Epidendrum palmense Ames, loc. cit. 2:33. 1923.
Epidendrum subviolascens Schltr. loc. cit. 19:219. 1923.
Erect, caespitose or repent, epiphytic herbs up to about 40 cm. tall. Stems
indurated, covered toward the base with sub-scarious sheaths and above by the
amplexicaul leaf-sheaths, bearing 2-4 leaves above. Leaves 7-30 cm. long and
1—4.5 cm. broad, ligulate or ligulate-oblong, obtuse, apex bilobed, keeled dorsally
along the mid-nerve. Inflorescence up to 30 cm. long, with up to 15 flowers, the
peduncle enveloped in the imbricated floral-bracts; bracts 15—65 mm. long and
10—30 mm. broad when spread out, strongly cucullate, obtuse; flowers variable in
size, greenish or the lip purplish. Dorsal вера! 15-25 mm. long and 3.5-6 mm.
broad, lanceolate, acute. Lateral sepals 15-25 mm. long and 5-11 mm. broad,
lanceolate to ovate-lanceolate, acute, oblique. Petals 14-20 mm. long and 1.5-2.5
mm. broad, linear to linear-oblanceolate, acute. Lip unguiculate; the claw adnate
to the column; lamina 12-25 mm. long and 14-28 mm. broad, cordate-reniform,
usually retuse or bilobed at the apex, fleshy, provided with a longitudinal callus-
thickening, entire or serrulate.
Mexico, Guatemala, Costa Rica, Panama, Venezuela, Ecuador and Peru.
HIRIQUi: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 245; province of
Chiriquí, alt. 1200 m., Powell 19, 3304. cocté: region north of El Valle de Antón, alt.
1000 m., Allen 2901
A. variable species that has been much described.
(278)
19451
FLORA OF PANAMA (Orchidaceae) 347
35. EPIDENDRUM CRINIFERUM Reichb. f. in Gard. Chron. 1291. 1871; Hook. f.
іп Bot. Mag. 100: /. 6094. 1874.
Erect, caespitose, epiphytic herbs up to 50 cm. tall Stems slender, strict,
simple, covered with the amplexicaul leaf-sheaths or naked below. Leaves 5-13
cm. long and 0.5-1.5 cm. broad, elliptic-linear to linear-lanceolate, acute, erect
or spreading. Inflorescence a simple few-flowered raceme, peduncle covered with
several large imbricated spathe-like bracts at the base. Sepals 10-25 mm. long
and 2-3 mm. broad, linear-lanceolate to lanceolate, acute or acuminate. Petals
10—20 mm. long and about 1 mm. broad, filiform. Lip unguiculate; claw adnate
to the column; lamina strongly 3-lobed, the lateral lobes semi-ovate with crenate
or laciniate margins and the setae usually recurved, the mid-lobe 5-12 mm. long
and linear or filiform, disc provided with 2 tubercular calluses at the apex of the
column.
Costa Rica, Panama and Peru.
СОСІ.Е: north rim of El Valle de Antón, alt. 600-1000 m., Allen 1670.
36. EPIDENDRUM CRYPTANTHUM Г. Wms. in Am. Orch. Soc. Bull. 11:249, 8. 7.
1942.
Repent, branched, epiphytic herbs up to about 40 cm. long. Stems slender,
covered with the scarious amplexicaul leaf-sheaths, becoming naked with age,
arcuate, continuing the growth from about the middle of the previous year's
growth, unilateral in origin, somewhat swollen toward the apex of each growth.
Leaves 5.5-8 cm. long and 1.7-3 cm. broad, elliptic to elliptic-ovate, acute,
coriaceous, 2-3 at the apex of each growth. Inflorescence about 3 cm. long, a
short, congested subcapitate raceme; bracts conspicuous, ovate or ovate-oblong,
entire or lacerate toward the apex; flowers small, hidden by the subtending bracts.
Dorsal sepal about 10 mm. long and 3 mm. broad, elliptic-oblong, obtuse. Lateral
sepals 10-11 mm. long and 3.5-4 mm. broad, elliptic-oblong or elliptic-oval,
obtuse or acutish, arcuate. Petals about 10 mm. long and 3.5-4 mm. broad,
oblanceolate-oblong, obtuse. Lip short-unguiculate; lamina about 10 mm. long
and 8 mm. broad, obovate, cordate at the base, the apex apiculate, lateral margins
involute; disc fleshy.
Panama.
COCLÉ: hills north of El Valle de Antón, alt. 800-1000 m., Allen 2262.
37. EPIDENDRUM DENTIFERUM Ames & Schweinf. Sched. Orch. 8:42. 1925.
Epidendrum platychilum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:42. 1923, non Schltr.
1921.
Small, repent, epiphytic herbs up to about 10 cm. long. Stems slender, simple
or branched, covered with the scarious leaf-sheaths or becoming naked. Leaves
2-7 cm. long and 0.3—0.5 cm. broad, linear or ligulate. Inflorescence terminal,
few-flowered; flowers relatively large for the plant. Dorsal sepal 8-15 mm. long
(279)
[Vor 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
348
Epidendrum cryptanthum
Fig. 127.
(280)
19461
FLORA OF PANAMA (Orchidaceae) 349
and 2-2.5 mm. broad, lanceolate, acute. Lateral sepals 9-16 mm. long and
2.5-3.5 mm. broad, lanceolate, acuminate, arcuate. Petals 8-15 mm. long, fili-
form or subulate. Lip simple, unguiculate; claw adnate to the column; lamina
7-13 mm. long and 8-14 mm. broad, suborbicular or suborbicular-reniform,
mucronate, provided with 2 small basal calluses.
Costa Rica and Panama.
DARIEN: Cana Cuasi trail, Chepigana District, alt. 2400 m., Terry 9 Terry 1412.
38. EPIDENDRUM DENTILOBUM Ames, Hub. & Schweinf. in Bot. Mus. Leafl. Harv.
Univ. 3:69. 1935.
Erect, caespitose, epiphytic herbs up to about 30 cm. tall. Stems slender,
simple, covered with the scarious leaf-sheaths or naked below. Leaves 6-10 cm.
long and 1.2-1.8 cm. broad, elliptic, acute. Inflorescence simple or branched,
few-flowered; flowers inadequately known. Dorsal sepal about 12 mm. long and
5 mm. broad, concave, narrowly oval. Lateral sepals about 10 mm. long and 6
mm. broad, oblong-oval, oblique, acute. Petals about 11 mm. long and 5.5 mm.
broad, spatulate-oblanceolate, oblique, obtuse. Lip unguiculate; claw short,
adnate to the column; lamina 3-lobed, about 7 mm. long and 8 mm. broad; lateral
lobes small, dentiform, narrowly triangular-lanceolate, explanate, about 1.5—2 mm.
long; terminal lobe largest, semicircular-flabellate, cuneate to the base; disc with
3 thickened, carinate nerves, the central one most prominent and with a mammil-
late callus at the base.
Panama.
CANAL ZONE: hills back of San Juan, alt. sea-level, Powell 342.
The available material of this species is inadequate.
39. EPIDENDRUM DIFFORME Jacq. Enum. Pl. Carib. 20. 1760; Select. Stirp. Am.
225, £. #30: 71163
Epidendrum umbellatum Sw. Nov. Gen. & Sp. Prodr. 121. 1788.
Epidendrum latilabrum Lindl. in Bot. oc 27: Misc. p. 77. 1841.
Epidendrum chlorocorymbos Schltr. in Fedde Rep. Sp. Nov. Beih. 17:30.
ое curvicolumna Ames, Hub. & Schweint. in Bot. Mus. Leafl. Harv. Uis. 3:67.
Caespitose or repent, epiphytic herbs up to 50 cm. tall. Stems fractiflex to
straight, covered with the pergameneous, amplexicaul, occasionally infundibuli-
form leaf-sheaths. Inflorescence a short, few- to many-flowered, subumbellate
raceme; flowers greenish or yellowish, long-pedunculate. Leaves 2-12 cm. long
and 0.5—3.5 cm. broad, variable, from linear-oblong to elliptic-oval, obtuse. Dorsal
sepals 14-35 mm. long and 2-4 mm. broad, linear-elliptic to lanceolate or ob-
lanceolate, acute. Lateral sepals 12-35 mm. long and 3-6 mm. broad, elliptic to
oblanceolate, acute, oblique. Petals 7-30 mm. long and 1-5 mm. broad, filiform
to linear to oblanceolate, acute. Lip 7-18 mm. long and 12—30 mm. broad,
unguiculate; claw adnate to the column; lamina transverse, reniform, broader
than long, obscurely to prominently 3-lobed; mid-lobe usually bifid and longer
(281)
350
ANNALS OF THE MISSOURI BOTANICAL GARDEN
[Vor. 33
Fig. 128. Epidendrum difforme
(282)
19461
FLORA ОЕ PANAMA (Orchidaceae) 351
than the lateral lobes.
From Florida and Mexico through Central America and the West Indies to
Brazil and Peru.
са Вајо Chorro, alt. 1800 m., Davidson 182; Volcan de Chiriqui, alt. 2100
m., Davidson 885; David, alt. 30-80 m., Pittier 2845; Lino Hill, alt. 1200-1500 m.,
Powell Mery Monte Lirio, alt. 1300-1900 m., Seibert 211. COCLÉ: hills north of El Valle
de Antón, alt. 800-1000 m., Allen 2287, 2293; Valle Chiquito, alt. 700—800 m., Seibert
514; San Juan, Chorrera, Pus and Frijoles, Powell 82, 249, 304, 3250, 3202, 3203, 3294,
3389.
A variable and widely distributed species which may include E. Barbeyanum
Krinzl., a species reported from Panama by garden specimen. The two following
varieties are hardly distinct.
39а. EPIDENDRUM DIFFORME Jacq. var. FIRMUM (Reichb. f.) Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 2:55. 1934
Epidendrum firmum Reichb. f. Beitr. Orch. Centr.-Am. 87. 1866.
Epidendrum majale Schltr. in Већ. Bot. Centralbl. 36, Abt. 2:406. 1918.
Distinguished from the species in having the mid-lobe of the lip “ас most little
broader than long, subquadrate.”
Guatemala, Costa Rica and Panama.
CHIRIQUÍ: Finca Lérida to Peña Blanca, alt. 1750-2000 m., Woodson & Schery 208.
39Ь. EPIDENDRUM DIFFORME Jacq. var. SIMULACRUM (Ames) Ames, Hub. &
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 2:57. 1934
Epidendrum simulacrum Ames, Sched. Orch. 6:75. 1923.
А small-flowered variant of the species, said to be distinguished from "'small-
flowered specimens of the typical form of the species in having the mid-lobe of
the lip a little broader than long.”
Panama.
CHIRIQUÍ: Lino Hill, alt. 1200-1500 m., Powell 208.
40. EPIDENDRUM EBURNEUM Reichb. f. іп Gard. Chron. 404. 1867; Hook. f. in
Bot. Mag. 93: ¢. 5043. 1867.
Epidendrum leucocardium Schltr. in Fedde Rep. Sp. Nov. 12:206. 1913.
Slender, caespitose, epiphytic herbs up to about 50 cm. tall. Stems strict,
covered with amplexicaul leaf-sheaths, becoming naked below. Leaves 3—14 cm.
long and 0.8—2.5 cm. broad, elliptic to lanceolate, obtuse or acute. Inflorescence
short, usually exceeded by the leaves, rachis strongly fractiflex; flowers rather
large, white. Dorsal sepal 22-36 mm. long and 2.5—4 mm. broad, linear-lanceolate,
acute or acuminate. Lateral sepals 22-36 mm. long and 3-4 mm. broad, linear-
lanceolate, acute or er slightly oblique. Petals 20-35 mm. long and 1—2
mm. broad, filiform. Lip with claw adnate to the column; lamina 20—30 mm.
long and 22-28 mm. ud orbicular-cordate to subquadrate or transversely
orbicular-oblong, usually apiculate, entire or rarely lobed (one flower seen wit
(283)
[Vor. 33
352 ANNALS OF THE MISSOURI BOTANICAL GARDEN
distinct sinuses) ; disc with 2 mammillate calluses at the base near the apex of the
column.
Panama, near sea-level.
CAS DEL TORO: vicinity of Chiriqui Lagoon, Wedel 8 50, 1625, 2556, 2766. CANAL
ZONE: Gatun Lake, near Colón, Powell 38, 132, 3462; “а few miles from Colón in
swamps,” Henderson. согом: Río Indio de Fató, Pittier 4266.
41. EPIDENDRUM ELLIPSOPHYLLUM L. Wms. in Ann. Missouri Bot. Gard. 28:419,
[. 22, figs. 1-2. 1941.
Erect epiphytic herbs up to about 25 cm. tall. Stems slightly fusiform and
indurated, with only the annual leaves persisting and the sheaths of the older
leaves becoming scarious and disintegrating. Leaves 11-12 cm. long and 2-2.8
cm. broad, elliptic, short-acuminate, usually only 2 or 3 persisting. Peduncles up
to about 4 cm. long, ancipitous, generally covered with imbricated bracts. In-
florescence short, 1-, or at most few-,flowered; bracts up to about 2 cm. long,
cucullate, lanceolate-triangular, acute; flowers green, segments stiff and coriaceous.
Sepals 10—12 mm. long and 3—4 mm. broad, elliptic to oblanceolate, obtuse, 5-
nerved. Petals about 10-11 mm. long and 3-4 mm. broad, oblanceolate, obtuse,
with 3 principal nerves. Lip unguiculate, the claw mostly adnate to the column;
lamina 5-6 mm. long and 5-6 mm. broad, orbicular-ovate іп outline, 3-lobed, the
lateral lobes small and situated toward the base of the lip, with a 2-ridged lamellate
callus plate at the junction of the claw and the lamina. Column slender, about
8 mm. long; clinandrium entire and obtuse; pollinia not seen.
Panama.
cocLÉ: hills north of El Valle de Antón, alt. 1000 m., Allen 2178, 2706.
This species is illustrated in fig. 123.
42. EpIDENDRUM Емркеѕи Reichb. f. in Gard. Chron. п. s. 19:432. 1883; Hook.
f. in Bot. Mag. 128: £. 7855. 1902.
Epidendrum Adolphi Schltr. іп Fedde Rep. Sp. Nov. 3:108. 1906.
Caespitose or repent, epiphytic herbs up to about 30 cm. long. Stems slender,
simple or branched; the amplexicaul leaf-sheaths verrucose and tuberculate.
Leaves 1—4.5 cm. long and 0.5—1.5 cm. broad, ovate to oblong-lanceolate, obtuse,
coriaceous. Inflorescence a simple, few-flowered raceme; flowers white tinged with
purple. Dorsal sepal 8-10 mm. long and 3-5 mm. broad, elliptic to oval, acute.
Lateral sepals 9-11 mm. long and 3.5—4.5 mm. broad, ovate-oblong, acute. Petals
8-11 mm. long and 2.5—5 mm. broad, oblanceolate-spatulate to obovate-spatulate,
obtuse, slightly oblique. Lip unguiculate; claw adnate to the column; lamina
7-12 mm. long and 6—10 mm. broad, 3-lobed; lateral lobes 2—4 mm. long, ex-
planate, subtriangular, obtuse or acute; mid-lobe bifid with the lobules spreading;
disc provided with 2 submammillate calluses under the apex of the column.
Costa Rica and Panama.
CHIRIQU{: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 323.
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19461
FLORA OF PANAMA (Orchidaceae) 353
43. EPIDENDRUM EQUITANTIFOLIUM Ames, Sched. Orch. 4:39. 1923.
Epidendrum equitans Lindl. in Bot. Reg. 24: Misc. p. 44. 1838, non Forst. f. пес
Ruiz & Pavon.
Erect or pendent, caespitose, epiphytic herbs up to about 4 dm. long. Stems
slender, leafy, ancipitous. Leaves 8-27 cm. long and 0.4-1.2 cm. broad, equitant,
ancipitous, linear-lanceolate, acute or acuminate, fleshy. Inflorescence borne on an
ancipitous, winged peduncle up to 25 cm. long and simulating a leaf, 1- to few-
flowered, subcapitate, subtended by a reduced leaf-like bract. Flowers small, dull-
colored. Sepals 12-17 mm. long and 1.5-2.5 mm. broad, linear to linear-oblong
or elliptic, acute or obtuse. Petals 8-13 mm. long and 1-2 mm. broad, linear-
oblanceolate to linear-elliptic. Lip unguiculate; the claw adnate to the column
for about half its length; lamina 7-9 mm. long and 5-6 mm. broad, 3-lobed,
provided with a callus on the claw at the base; lateral lobes semi-orbicular; termi-
nal lobe lanceolate to triangular-lanceolate; disc concave or cochleate.
Mexico, British Honduras, Honduras and Panama.
CANAL ZONE: hills east of Panamá City, Powell 206, 3409. снікюоі: "Province of
Chiriqui”, alt. 900-1050 m., Powell 206.
44. EPIDENDRUM EXASPERATUM Reichb. f. Beitr. Orch. Centr.-Am. 87. 1866.
Epidendrum chondranthum Kránzl. in Vierteljahrschr. Naturforsch. Ges. Zürich 74:136.
1929.
Erect, caespitose, epiphytic herbs up to about 3 dm. tall, normally less. Stems
slender to thickened, simple or branched, covered with amplexicaul leaf-sheaths,
becoming naked with age. Leaves 4—13 cm. long and 0.5—4.5 cm. broad, elliptic
to lanceolate to lanceolate-oblong, obtuse or acute, those on branches smallest and
narrowest. Inflorescences either terminal or lateral or both, the lateral ones
originating opposite the base of a leaf, simple or paniculate. Flowers greenish or
whitish and mottled with red or purple. Dorsal вера! 8-14 mm. long and 3-6
mm. broad, oblong-lanceolate, obtuse or acute, cucullate, fleshy, verrucose to
mammillate dorsally, the mid-nerve prolonged into a subterminal apicule. Lateral
sepals similar but elliptic-oblong and usually oblique. Petals 8-11 mm. long and
2—4 mm. broad, oblanceolate, obtuse. Lip unguiculate, the claw adnate to the
column, fleshy or callus-thickened; lamina 7-13 mm. long and about as broad,
prominently 3-lobed; lateral lobes spreading, originating about opposite the apex
of the column, oblong, oblique, obtuse, 3-5 mm. long, rarely lobulate or dentate;
mid-lobe bilobulate, emarginate, the lobules divergent to explanate, rarely dentate,
obtuse; disc usually provided with a low callus-thickening which extends from
the claw to the isthmus between the lateral lobes and usually is terminated by two
mammillate processes, rarely continuous from base to apex,—often obscurely
verrucose. Column provided with 2 thin stelidia at the apex over the anther.
Costa Rica and Panama.
(285)
[Vor. 33
354 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CHIRIQUÍ: south slope of Volcán de Chiriquí, Boquete District, alt. 2900 m., Terry
8 Terry 1343; trail from Potrero Muleta to Finca Lérida, Woodson 9 Schery 475.
45. Ерремрком HUNTERIANUM Schltr. іп Fedde Rep. Sp. Nov. Beih. 17:33.
1922. |
Pendent or erect epiphytic herbs up to about 30 cm. long. Stems simple,
somewhat compressed, densely leaved. Leaves to 7.5 cm. long and 4 cm. broad,
oblong or broadly oblong, obtuse, unequally bilobed. Inflorescence short, terminal,
4- to 6-flowered, subumbellate; bracts ovate, obtuse to acuminate. Sepals about
1.5 cm. long, oblong-ligulate, subacute, the laterals oblique. Petals about as long
as the sepals but narrower, lanceolate-ligulate, subacute, narrowed at the base.
Lip unguiculate, the claw adnate to the column; lamina about 11 mm. long and
about 10 mm. broad, suborbicular-cordate, very obtuse, base with 2 oblique
calluses, median nerves thickened.
Panama.
CANAL ZONE: Im Uberschwemmungsgebiete des Gatun-Sees, Powell 29.
No specimens were seen of this species. Description taken from the original.
46. EPIDENDRUM IBAGUENSE HBK. Nov. Gen. & Sp. РІ. 1:352. 1816; С.
Schweinf. in Bot. Mus. Leafl. Harv. Univ. 11:229. 1944.
Epidendrum radicans Pavon ex Lindl. Gen. & Sp. Orch. Pl. 104. 1831; Ames, Hub. &
chweinf. Genus Epidendrum in U. S. & Mid. Am. 162. 1936
Epidendrum radicans Pavon var. chiriquense Schltr. in Fedde Rep. Sp. Nov. Beih. 17:39.
14824.
Variable terrestrial or epiphytic herbs with stems from about 0.1 to 1 m. long,
erect, pendent or sprawling. Stems simple or branched, usually slender and leaf-
bearing on the newer growths, provided with long whitish roots which orig-
inate opposite the bases of some of the leaves, or rarely without these roots. Leaves
1.5-12 cm. long and 0.6-3.5 cm. broad, ligulate to oval or oblong to ovate, obtuse,
fleshy, lax or crowded. Inflorescence а lax to short subumbellate or paniculate
raceme borne at the apex of a scape which may be up to 1 m. long; scape slender,
terete, covered with scarious, appressed bracts or becoming naked, often branched.
Flowers variable in size and color, from red to white but most often brick-red.
Dorsal sepal 12-22 mm. long and 4-7 mm. broad, elliptic or oblong to narrowly
obovate, acute. Lateral sepals similar but slightly oblique. Petals 12-20 mm. long
and 5-8 mm. broad, elliptic to obovate, acute. Lip unguiculate, the claw adnate
to the column; lamina 7-17 mm. long and as broad, 3-lobed, suborbicular-cordate
in outline; the lateral lobes large, rounded, larger than the mid-lobe, the margins
entire or usually lacerate; mid-lobe usually oblong in outline, bilobulate with the
lobules slighty divergent, entire or usually lacerate, equalling or exceeding the tips
of the lateral lobes in length; disc provided with 2 flattened submammillate calluses
at the apex of the column and usually with 2 inconspicuous lamellate calluses
extending from the apex of the column toward the apex of the lip.
(286)
19461
FLORA OF PANAMA (Orchidaceae) 355
Mexico to Panama, Venezuela, Colombia, Peru and British Guiana.
BOCAS DEL TORO: without locality, Wedel 12. CANAL ZONE: Quebrada Ancha, alt.
770 m., Dodge & Steyermark 17042; Barro Colorado Island, Woodworth & Vestal 703,
70 CHIRIQUÍ: Volcán de Chiriquí, alt. 2450 m., Davidson 945; valley of Rio Caldera,
alt. 1400—1600 m., Killip 3517; Chiriquí, alt. 1000- 1200 m., Powell 61, hee: 3515; be-
tween Concepción and El Volcán, Peggy White 314; Chiriqui Viejo Valley, Gene White
84; Finca Lérida to Boquete, alt. 1300-1700 m., Woodson, Allen 9 Seibert 1098; vicinity
of Boquete, alt. 1200-1500 m., Woodson 9 Schery 707.
A variable species, both in vegetative and floral structure. Where it occurs it
is often exceedingly common.
47. EPIDENDRUM IMATOPHYLLUM Lindl. Gen. & Sp. Orch. РІ. 106. 1831; Ames,
Sched. Orch. 4:43, figs. 1923. |
Epidendrum lorifolium Schltr. in Fedde Rep. Sp. Nov. Већ. 17:35. 1922.
Epiphytic (or terrestrial) herbs occasionally up to about 1 m. long. Stems
slender, somewhat weak, arising from a rhizome, covered with the pergameneous
leaf-sheaths, becoming naked with age. Leaves 6-20 cm. long and 0.6-3 cm.
broad, ligulate to ligulate-lanceolate, acute or obtuse, coriaceous. Inflorescence
simple or occasionally branched, a more or less compact raceme borne at the apex
of a slender peduncle which may be up to 2 dm. long; flowers rose to purple.
Dorsal sepal 13-20 mm. long and 3.5-5 mm. broad, elliptic to elliptic-oblanceolate,
acute or acuminate. Lateral sepals similar to the dorsal except oblique. Petals
13-18 mm. long and 6-8 mm. broad, elliptic to elliptic-rhomboid, acute. Lip
unguiculate; lamina 7-10 mm. long and 5-9 mm. broad, oblong to oblong-
obovate in outline, obscurely 3-lobed or lacerate-dentate, especially toward the
base; disc with 2 short submammillate calluses at the apex of the column and with
a short lamellate callus between them and extending toward the apex of the lip.
Mexico to Panama, Trinidad, Colombia to Peru, the Guianas and Brazil.
CANAL ZONE: е swamp, Allen 867. PANAMA: Sabana, Pittier 3403; Tapia
River, Juan Diaz region, Maxon 9 Harvey 6608; above Peluca Hydrographic Station,
Hunter & Allen Bea; “Panama”, Powell 138.
This species usually has its roots inhabited by ants. It is said to deteriorate in
cultivation if the ants are exterminated.
48. EpIDENDRUM INCOMPTUM Reichb. f. in Bot. Zeit. 10:733. 1852; Ames in
Bot. Mus. Leafl. Harv. Univ. 4:64, 2. 1936.
Erect or scandent epiphytic herbs up to about 70 cm. long. Stems branched
or rarely simple, bearing leaves near the apex of the stem, covered with the amplexi-
caul leaf-sheaths or naked below. Leaves 3-13 cm. long and 1.5-5.5 cm. broad,
narrowly oblong to oblanceolate-acute, coriaceous. Inflorescence a few- to several-
flowered terminal raceme, about equal to the subtending leaves; flowers greenish.
Dorsal sepal 11-15 mm. long and 3.5-5.5 mm. broad, oblanceolate, acute. Lateral
sepals 11-15 mm. long and 4.5-6 mm. broad, semilunate, acute. Petals 11-14
mm. long and 1.5—2 mm. broad, narrowly oblanceolate, acute. Lip unguiculate;
claw adnate to the column; lamina 7-9 mm. long and 12-17 mm. broad, fleshy,
(287)
[Vor. 33
356 ANNALS OF THE MISSOURI BOTANICAL GARDEN
ae
тӘ
4 Лы S а №.
^ Cd АА S,
ЕРОЕМОКОМ inçomplam * vid сеге. f.
Fig. 129. Epidendrum incomptum
(288)
1946]
FLORA OF PANAMA (Orchidaceae) 357
3-lobed, subreniform in outline, lobes about equal, the laterals divaricate and
rounded, the mid-lobe subtriangular; disc without calluses.
Mexico, Guatemala, Costa Rica and Panama.
CHIRIQUÍ: “Chiriqui”, Warscewicz.
No specimens of this species have been found in Panama since the original was
collected.
49. EPIDENDRUM ISOMERUM Schltr. in Fedde Rep. Sp. Nov. 2:132. 1906.
Pendent or repent epiphytic herbs up to 1 m., or perhaps more, long. Stems
slender, simple or usually branched, flexuose, bearing leaves toward the apex of the
new growths, covered with the chartaceous leaf-sheaths, even when old. Leaves
2-12 cm. long and 0.15-0.4 cm. broad, linear, acute, flat or subterete. Inflores-
cence a single flower borne at the apex of new growths. Flower small, yellowish.
Sepals 11-15 mm. long and 2-3 mm. broad, linear-lanceolate, acute, margins
revolute. Petals 11-13 mm. long and 1.2-1.5 mm. broad, elliptic-linear, acute.
Lip unguiculate, claw adnate to the column; lamina 8—9 mm. long and 2-2.5 mm.
broad, narrowly lanceolate, trulliform or canaliculate, fleshy, acute; disc provided
with 2 short inconspicuous calluses at the base and with 1 sublamellate median
callus extending from the base to about the middle of the lamina.
Mexico, Guatemala, British Honduras, Costa Rica and Panama.
OCAS DEL TORO: between Finca St. Louis and Konkintoë, alt. 10-15 m., Woodson,
Allen 5 Seibert 1886. CANAL ZONE: Rio Medio, Miller 1746.
50. EPIDENDRUM LOCKHARTIOIDES Schltr. in Fedde Rep. Sp. Nov. Beih. 19:39.
1923; Ames, Sched. Orch. 8:46, Ё. 24. 1925
Small, densely caespitose, epiphytic herbs up to about 2.5 dm. tall, usually less.
Stems covered with the subequitant bases of the leaves. Leaves 1—3.5 cm. long,
laterally compressed, approximate, spreading, cultriform, and the sheathing base
subequitant, 5-10 mm. broad. Inflorescence composed of several flowers borne
in the axils of the upper (sheath-like) leaves; flowers inconspicuous, greenish-
yellow or green. Dorsal sepal 6-7 mm. long and about 3 mm. broad, broadly
lanceolate, acute, cucullate. Lateral sepals 6—8 mm. long and 2-3 mm. broad,
cucullate, arcuate, lanceolate, prominently keeled along the mid-nerve dorsally,
the keel usually denticulate. Petals 4-7 mm. long and 0.5—1 mm. broad, linear,
obtuse, slightly arcuate. Lip 5—7 mm. long, unguiculate; claw adnate to the
column; lamina 4—5 mm. long and as broad, cordate-orbicular to subquadrate,
obtuse, often with a short apicule, mid-nerve callus- ере;
Costa Rica and Panama.
cocLÉ: El Valle de Antón, alt. 600-1000 m., Allen 1680, 2206.
51. EPIDENDRUM MICRODENDRON Reichb. f. in Beitr. Orch. Centr.-Am. 84. 1866.
Pendent or repent epiphytic herbs up to about 70 cm. long. Stems slender,
branched, covered with the leaf-sheaths or becoming naked. Leaves 1.5—5 cm. long
and 0.3—0.9 cm. broad, linear-ligulate to elliptic-oblong, obtuse and with the apex
(289)
[Vor. 33
358 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
obscurely and unequally bilobed. Inflorescence a short terminal subfractiflex
raceme of 2-8 flowers; bracts 4-7 mm. long, chartaceous, cucullate, acute. Dorsal
вера! 7-10 mm. long and 1.5-3 mm. broad, lanceolate, acute. Lateral sepals 6.5-9
mm. long and 2.5-3 mm. broad, lanceolate to ovate-lanceolate, acute, oblique.
Petals 5-10 mm. long and 0.7-1.5 mm. broad, linear to elliptic-linear, acute. Lip
unguiculate; the claw short, adnate to the column; lamina 5-8 mm. long, and
when expanded about as broad, 3-lobed but obscurely so, the lateral lobes erect,
rounded, enfolding the short column, mid-lobe lanceolate or narrowly triangular,
acute or acuminate, fleshy, and the apex subterete; disc provided with a small
cucullate callus under the apex of the column which is entire or tridentate at the
apex. Column provided with a horn-like process at its apex.
Costa Rica and Panama.
CHIRIQUÍ: Cuesta de las Palmas, Serra de la Horqueta, alt. 1700-2100 m., Pittier 3220.
The specimen from Panama is not typical—perhaps represents a distinct species.
52. EPIDENDRUM MOYOBAMBAE Krinzl. in Fedde Rep. Sp. Nov. 1:185. 1905; С.
Schweinf. in Bot. Mus. ГеаЙ. Harv. Univ. 11:238. 1944.
Epidendrum subpatens Schltr. in Fedde Rep. Sp. Nov. Beih. 17:40. 1922.
Epidendrum benignum Ames in Sched. Orch. 2:26. 1923
Pendent or ascending, caespitose, epiphytic herbs up to about 8 dm. long.
Stems slender, simple, covered with the leaf-sheaths above, naked at the base.
Leaves 6-16 cm. long and 1.5-4 cm. broad, elliptic-lanceolate to elliptic-ovate,
acute, coriaceous. Inflorescence a terminal few- to many-flowered pendent raceme
up to about 4 dm. long; flowers rather large, greenish to yellowish or whitish.
Dorsal вера! 17-25 mm. long and 5-9 mm. broad, oblanceolate, acute or obtuse.
Lateral sepals 17-25 mm. long and 5-9 mm. broad, oblanceolate to narrowly obo-
vate, oblique, obtuse. Petals 16-22 mm. long and 3-7 mm. broad, linear-oblance-
olate to broadly oblanceolate, obtuse or acute, slightly oblique. Lip unguiculate;
the claw adnate to the column; lamina 15-25 mm. long and 20-30 mm. broad,
3-lobed; lateral lobes spreading or deflexed, suborbicular, obtuse, much larger than
the mid-lobe; mid-lobe bilobulate, the lobules more or less divaricate, smaller than
either lateral lobe; disc provided with 2 small submammillate or mammillate
calluses under the apex of the column and with 1 or rarely 3 lamellate calluses
extending along or near the median line.
Guatemala, Costa Rica, Panama, Trinidad, Colombia, Peru and Brazil.
ZONE: Gatün Lake, Powell 86, 3337, 3558; Barro Colorado Island, Shattuck
216. VERAGUAS: Santiago, alt. 15 m., Powell 86, 3558.
Presumably this was originally confused with Epidendrum patens Sw. It may
prove to be the same. It is a rather showy plant. Powell's numbers for the two
localities are the same, and it is possible that all of the material came from one
original collection.
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19461
FLORA OF PANAMA (Orchidaceae)
359
#
Fig. 130. Epidendrum nocturnum
53. EPIDENDRUM NOCTURNUM Jacq. Enum. Pl. Carib. 29. 1760; Select. Stirp
Am. 225, #. 130. 1763; Ames, Hub. & Schweinf. Genus Epidendrum in U. 5
and Mid. Am. 129. 1936.
Epidendrum nocturnum Jacq. var. panamense Schltr. in Fedde Rep. Sp. Nov. Beih. 17:36.
1922.
Variable, caespitose or subrepent, epiphytic plants up to 1 m. tall. Stems
slender, simple, covered with the scarious leaf-sheaths or becoming naked. Leaves
(291)
[Vor. 33
360 ANNALS OF THE MISSOURI BOTANICAL GARDEN
5-15 cm. long and 0.8-6.5 cm. broad, elliptic, elliptic-oblong, ligulate or oval,
acute or obtuse, coriaceous to fleshy. Inflorescence a 1- to few-flowered terminal
raceme (1 flower open at a time); rachis when present fractiflex; flowers large,
usually white. Sepals similar, 35-90 mm. long and 2-5 mm. broad, from filiform
to linear, acute or acuminate. Petals similar to the sepals but smaller, 35-80 mm.
long and 1-3 mm. broad. Lip unguiculate, claw adnate to the column; lamina
up to 65 mm. long, prominently 3-lobed; lateral lobes subparallel, acute, oblique,
lanciform, 10-40 mm. long; mid-lobe subfiliform, 20-40 mm. long.
Florida, Mexico to Panama, the West Indies and tropical South America.
8 DEL ToRO: vicinity of Chiriqui Lagoon, Wedel 1001, 2938. CANAL ZONE:
hills near Panamá City, Gatün Lake, Frijoles, San Juan, aei nó Paja, Powell 35, 216,
217, 3020, 3055, 3073, 3075. CHIRIQUÍ: “Chiriqui”, alt. 1200-1400 m., Powell 347.
состЕ: hills north of El Valle de Antón, alt. 800-1000 m., Allen 2257.
Occasional throughout the American tropics, usually found at low elevations
but occasionally in the mountains.
54. EPIDENDRUM OBESUM Ames in Sched. Orch. 2:31. 1923.
Coarse, caespitose, epiphytic herbs up to about 6 dm. tall. Stems slender,
covered with the loosely amplexicaul complanate leaf-sheaths or reduced leaves.
Leaves 5-20 cm. long and 3-7 cm. broad, elliptic-ligulate to oval, obtuse, usually
crowded on the stem. Inflorescence a short, few- to several-flowered raceme sub-
tended by a large spathaceous sheath, the flowers long-pedunculate, whitish. Dorsal
sepal 35—55 mm. long and 3-7 mm. broad, linear or linear-elliptic, acute. Lateral
sepals similar to the dorsal except slightly arcuate. Petals 28-50 mm. long and
1.5—4 mm. broad, linear, acute. Lip unguiculate; the claw adnate to the column;
lamina 18-30 mm. long and up to about 15 mm. broad, 3-lobed; the lateral lobes
semiorbicular to subquadrate, explanate, obtuse, about 6-8 mm. long and as broad;
mid-lobe about 14-25 mm. long, linear-lanceolate to very narrowly triangular,
acute; disc provided with 2 fleshy calluses at the base, the apices of which are free
and porrect. Pedicellate ovary up to about 12 cm. long.
Nicaragua, Costa Rica and Panama. Possibly also in Ecuador.
CHIRIQUÍ: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 1288.
The flowers somewhat resemble those of Epidendrum nocturnum Jacq.
55. EPIDENDRUM PANAMENSE Schltr. in Fedde Rep. 12:212. 1913.
Caespitose epiphytic herbs up to about 2 dm. tall. Stems ancipitous, covered
with amplexicaul leaf-sheaths, provided with 2—3 leaves toward the apex. Leaves
8-11 cm. long and 1-1.3 cm. broad, elliptic to lanceolate, acuminate. Inflores-
cence a compact few-flowered distichous raceme terminating a slender ancipitous
peduncle. Sepals about 1.2 cm. long, ligulate, apiculate, the laterals oblique.
Petals about as long as the sepals, linear. Lip unguiculate; claw adnate to the
column; lamina about 5 mm. long and 4 mm. broad, ovate, shortly acuminate,
subcordate at the base, provided with 2 calluses at the base which have a short keel
between them. (Description compiled from tbe original).
(292)
19461
FLORA OF PANAMA (Orchidaceae) 361
Panama.
PANAMA: Cerro de Garagara, Sambü Basin, southern Darién, alt. 550—974 m., Pittier
5035.
Allied to Epidendrum Allenit L. Wms. and to E. Lankesteri Ames.
56. EPIDENDRUM PANICULATUM Ruiz & Pavon, Syst. Veg. 243. 1798; Hook. f.
in Bot. Мар. 94: 4. 5731. 1868; Ames, Hub. & Schweinf. in Bot. Mus. Leafl.
Harv. Univ. 2:67. 1934.
Bre е НВК. Nov. Gen. & Sp. 1:353, 7. 86. 1816; Hook. in Bot. Mag.
“к ш Scb. f. in Gard. Chron. 1871:1678. 1871.
1 1877
Epidendrum Isthmi Schltr. in Fedde Rep. Sp Nov. Beih. 17:34. 1922.
Epidéndrum reflexum Ames & Sekwa, in Sched. Orch. 8:49. 1925.
Caespitose epiphytic or saxicolous plants up to about 1.5 dm. tall, often rank
in growth. Stems simple, erect or spreading, covered with the amplexicaul leaf-
sheaths or naked with age. Leaves 4—25 cm. long and 0.5-7 cm. broad, variable
but essentially linear-lanceolate to oval, acute or acuminate, rarely obtuse, coria-
ceous. Inflorescence from shorter than the subtending leaves to much longer,
from a simple few-flowered raceme to a compound multiflorous panicle, terminal,
usually subtended with a spathe which may be up to 6 cm. long; bracts оп the
peduncle from very short to about 7 cm. long; flowers variable in size and color.
Dorsal вера! 7-16 mm. long and 2-3.5 mm. broad, elliptic to broadly oblanceolate,
acute or obtuse. Lateral sepals 8-16 mm. long and 2.5-4.5 mm. broad, elliptic to
broadly lanceolate, acute or obtuse. Petals 8-15 mm. long and 0.2-2 mm. broad,
usually filiform or filiform-clavellate but occasionally linear-oblanceolate. Lip
unguiculate; claw adnate to the column; lamina 4-10 mm. long and 5-12 mm.
broad, from nearly simple to 3- or 4-lobed (commonly 3-lobed); lateral lobes
from dolabriform to obliquely triangular-ovate or ovate, outer margin entire to
sublacerate; mid-lobe usually strongly bilobulate, but often from truncate to
retuse, often apiculate, lobules variable in form and divergence, linear to oblong
to triangular or spatulate, margin entire or somewhat lacerate; disc with 2 short
sublamellate calluses at the base, often provided with one or more callus ridges
parallel to the mid-nerve.
Mexico to Panama and in South America to Argentina.
CANAL ZONE: Quebrada Ancha, Steyermark & Allen 17100; hills east of Panamá
City, and Rio Indio near the mouth of Chagres River, Powell 104, 323, 3306. CHIRIQUI:
El Boquete, alt. 100-1300 m., Pittier 2073; “Chiriqui”, Powell 233, 3174; Rio Chiriqui
Viejo, alt. 1300-1900 m., Seibert 145; Upper Rio Chiriquí Viejo, White 326. сосІЁ:
vicinity of La Mesa, north of El Valle de Antón, alt. 1000 m., Allen 2380. PANAMA: Río
Tapia, Standley 28137
A widespread and variable species. Only the essential synonymy for Panama
and adjacent Costa Rica has been given above.
(293)
[Vor. 33
362 ANNALS OF THE MISSOURI BOTANICAL GARDEN
/ И 222222 22
А ee 2
Ју == |
Fig. 131. Epidendrum pendens
ИХ,
777
"
57. EPIDENDRUM PENDENS L. Wms. in Ann. Missouri Bot. Gard. 28:421, £. 23.
1941.
Pendent epiphytic herbs about 45 cm. long. Leaves 9-15 cm. long and 2-3.5
(294)
19461
FLORA OF PANAMA (Orchidaceae) 363
cm. broad, elliptic to elliptic-lanceolate, coriaceous, distichous, marginate, gray,
deciduous; leaf-sheaths loose, completely investing the stem, pergameneous, an-
cipitous, the abscission line prominent. Inflorescence of 1 or few large terminal
flowers subtended by a leaf-like sheath up to 3 cm. long; flowers large for the
genus, green. Dorsal sepal about 5 cm. long and 1.2 cm. broad, oblong-oblance-
olate, acute, fleshy, with 5-7 main nerves. Lateral sepals 4.3-4.8 cm. long and
1.5 cm. broad, elliptic, acute, fleshy, with about 7 main nerves. Petals about
4.5-5 cm. long and 2 mm. broad, linear, somewhat narrowed to the base, obtuse
or acute, fleshy, 1-nerved. Lip with the claw adnate with the column to the apex
of the column; lamina about 2.5-3 cm. long and 3 cm. broad, 3-lobed, suborbic-
ular in general outline, with a thickened central nerve and with thickened nerves
or ridges radiating from it and covering the remainder of the lip, lateral lobes
rounded, about 1.5 cm. long, mid-lobe 1-1.5 cm. long and about 1.5 cm. broad,
subquadrate, emarginate. Column 1.5-2 cm. long and 6-8 mm. in diameter at
the apex, fleshy.
Panama.
COCLÉ: vicinity of La Mesa, El Valle de Antón, alt. 1000 m., Allen 2570.
58. EPIDENDRUM PHYSODES Reichb. f. in Gard. Chron. 289. 1873.
Small repent or caespitose, epiphytic or terrestrial herbs up to about 30 cm.
long. Stems simple, slender, densely leaved, prolonged into a long peduncle above.
Leaves 1.5—6.5 cm. long and 0.3-0.8 cm. broad, lanceolate, the margins often
revolute. Inflorescence a short few-flowered raceme (with 1 flower open at a
time?) on the apex of a long peduncle. Dorsal sepal 5-9 mm. long and 2-3 mm.
broad, elliptic to oblong-lanceolate, acute. Lateral sepals 5-9 mm. long and
2.33.6 mm broad, elliptic-oblong to oblong-ovate, somewhat oblique, acute.
Petals 4.5 mm. long and 0.5—0.8 mm. broad, linear to subfiliform. Lip unguic-
ulate; claw adnate to the column; provided with a spur which is 3—4 mm. long,
adnate to the ovary for a part of its length or occasionally for all of its length
but usually the obtuse and rounded tip free; lamina 3-5 mm. long and 4-6 mm.
broad, entire, subreniform or suborbicular, fleshy; disc with 2 mammillate calluses
near the apex of the column and often with a longitudinal callus-thickening along
the median line.
Guatemala, Costa Rica and Panama.
BOCAS DEL TORO: vicinity of Chiriquí Lagoon, Wedel 1126.
This species is atypical in the genus Epidendrum in that it is provided with a
distinct spur which apparently originates from the lip. If generically distinct it
belongs in Lindley's genus Physinga. Suitable material for comparative studies is
not available.
59. EPIDENDRUM PLATYSTIGMA Reichb. f. in Beitr. Orch. Centr.-Am. 85. 1866.
Epidendrum ramonianum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:217. 1923.
Epidendrum gibbosum L..W ms. in Ann. Missouri Bot. Gard. 28:420, £. 21, figs. 7-10.
1941.
(295)
[Vor. 33
364 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 132. Epidendrum platystigma
Simple or branched, repent, terrestrial or epiphytic herbs up to probably more
than 5 dm. tall. Stems covered with the very loose leaf-sheaths or becoming naked
and vernicose, up to 15 mm. in diameter. Leaves 2-15 cm. long and 0.5-2.5 cm.
(296)
19461
FLORA OF PANAMA (Orchidaceae) 365
broad, linear-lanceolate or linear-oblong to ligulate, obtuse and often unequally
bilobulate at the apex, coriaceous, leaves on the sterile or poorly flowered branches
largest, on the heavily flowered branches very small; leaf-sheaths usually some-
what inflated and pergameneous, soon disintegrating. Inflorescences short, few-
flowered, terminal, the rachis fractiflex; bracts up to 2 cm. long, scarious, coch-
leate, acute; flowers greenish or whitish, often marked with red. Dorsal sepal
7-12 mm. long and 4-6 mm. broad, elliptic-oblong, acute. Lateral sepals 7-12
mm. long and 3-7 mm. broad, narrowly obovate to oblong-ovate, obtuse or acute,
apiculate or winged dorsally along the mid-nerve. Petals 8—10 mm. long and 3-5
mm. broad, oblanceolate to narrowly obovate, obtuse. Lip 6-12 mm. long and
5-10 mm. broad, broadly oblong to suborbicular or cordate, obtuse or shallowly
emarginate, fleshy and tumid along the mid-nerve or with several raised nerves,
gibbous at the base, provided with an inconspicuous bi- or trilobate callus under
the apex of the column.
Costa Rica and Panama.
CHIRIQUÍ: Volcán de Chiriquí, alt. 2600 m., Davidson 971; Casita Alta to Serra
Copete, alt. 2300-3300 m., Woodson & Schery 367.
60. EPIDENDRUM POLYANTHUM Lindl. Gen. & Sp. Orch. Pl. 106. 1831; Batem.
Orch. Mex. & Guat. /. 34. 1842.
Epidendrum pergameneum Reichb. f. Beitr. Orch. Centr.-Am. 86. 186 ў
Epidendrum quinquelobum Schltr. in Fedde Кер. Sp. Nov. Beih. 19:125. 1923,
Caespitose epiphytic herbs, often rank-growing, up to about 1.5 m. tall. Stems
erect or ascending, simple, leafy above. Leaves 5-25 cm. long and 0.8—5 cm.
broad, from linear to oval or ovate, acute or acuminate, variable. Inflorescence
terminal or lateral (i.e. flowers or racemes borne from below a leaf) or both,
usually consisting of racemes borne laterally on an elongated peduncle, rarely the
racemes branching and becoming paniculate, rarely consisting of a single short
terminal raceme; flowers extremely variable in size and coloration. Dorsal sepal
6.5-13 mm. long and 2.5-3.5 mm. broad, oblanceolate, obtuse, papillose externally.
Lateral sepals 6.5—16 mm. long and 2.5-5 mm. broad, oblanceolate, obtuse, ob-
lique, papillose externally. Petals 5-12 mm. long and about 0.5 mm. broad, fili-
form or subfiliform. Lip unguiculate; claw adnate to the column; lamina 4-7
mm. long and 4—7 mm. broad, usually broader than long, 3-lobed or rarely almost
entire; the lateral lobes oblong to flabellate, entire or usually undulate, dentate or
lacerate; mid-lobe from subquadrate to oblong, retuse or bilobulate, the lobules
sometimes explanate; disc provided with 2 mammillate calluses near the apex of the
column and with one or more callus-thickened ridges along the median line. Ovary
and pedicel usually papillose.
Guatemala, Honduras, Nicaragua, Costa Rica and Panama. Venezuela and
Brazil.
CHIRIQUÍ: Bajo Chorro, alt. 1400 m., Davidson 457.
А polymorphic species which, however, is usually easy to recognize.
(297)
[Vor. 33
366 ANNALS OF THE MISSOURI BOTANICAL GARDEN
60a. EPIDENDRUM POLYANTHUM Lindl. var. мүореѕ (Reichb. f.) Ames, Hub.
& Schweinf. in Bot. Mus. Leafl. Harv. Univ. 3:75. 1935.
Epidendrum myodes Reichb. f. Beitr. Orch. Centr.-Am. 86. 1866.
Similar to the species except the lateral sepals are said to be only 4-6.5 mm.
long (6.5-16 mm. long in the species).
Honduras, Costa Rica and Panama.
снікюілі: “Chiriqui”, alt. 1200 m., Powell 188, 242, 3398, 3452, 3404, 3472, 3505.
61. EPIDENDRUM PonPAX Reichb. f. in Bonplandia 3:220. 1855.
Epidendrum porphyrophyllum Schltr. in Fedde Кер. Sp. Nov. Beih. 17:37. 1922.
Small, caespitose, epiphytic herbs up to about 8 cm. tall. Stems short, covered
with the scarious amplexicaul sheaths of the leaves. Leaves 7-25 mm. long and
2—7 mm. broad, ligulate to lanceolate or oblong-lanceolate, obtuse or acute, fleshy,
spreading. Inflorescence consisting of a single flower from the apex of the stem,
subtended by a scarious sheath; flowers large for the plant, sepals and petals green-
ish, lip purple. Dorsal sepal 7-14 mm. long and 2-5 mm. broad, lanceolate to
lanceolate-ovate, acute. Lateral sepals 8-14 mm. long and 3-5.5 mm. broad,
lanceolate to lanceolate-ovate, acute, adnate at the base to the claw of the lip and
to the column for about the length of the column. Petals 7-13 mm. long and
0.5—1.5 mm. broad, linear, obtuse, obscurely ciliolate toward the apex. Lip up to
18 mm. long, unguiculate; claw narrow, adnate to the column; lamina 9-15 mm.
long and as broad, suborbicular-cordate, obscurely ciliolate, fleshy, provided with
2 mammillate calluses at the base under the apex of the column.
Mexico, Guatemala, Honduras, Costa Rica, Panama and Venezuela.
AL ZONE: foothills east of ты City, Powell 183. снимоој: “Province of
Chiriqui” , alt. 1200 m., Powell 400
62. EPIDENDRUM Ромел Schltr. in Fedde Rep. Sp. Nov. Већ. 17:38. 1922.
Caespitose epiphytic herbs up to about 30 cm. tall. Stems simple, somewhat
flexuose, covered with leaf-sheaths and ancipitous. Leaves 3.5-16 cm. long and
0.35—1.2 cm. broad, elliptic-linear, acute. Inflorescence up to about 7 cm. long,
a terminal, densely flowered raceme subtended by one or more spathaceous bracts.
Flowers small, whitish with pink or orange markings. Dorsal вера! 2.5-3.5 mm.
long and 1-1.2 mm. broad, narrowly lanceolate to oblong- oval, acute. Lateral
sepals 3-4 mm. long and 1.5-2 mm. broad, oblong-oval to ovate, oblique, acute.
Petals 2—3 mm. long and 0.3—0.4 mm. broad, linear, acute. Lip unguiculate; claw
adnate to the column; lamina 1.5—3 mm. long and about as broad, 3- lobed; the
lateral lobes dolabriform to semiorbicular, entire or crenate or lobulate; mid-lobe
ligulate to subtriangular, acute; disc with 2 small calluses at the base.
Panama.
осі: “Chiriqui”, alt. 1200 m., Powell 228, 3445, 3456. COCLE: hills south of
El Valle de Antón, alt. 600—800 m., Allen 2 2797.
(298)
19461
FLORA ОЕ PANAMA (Orchidaceae) 367
63. EPIDENDRUM PROBIFLORUM Schltr. in Fedde Rep. Sp. Хоу. Beih. 17:39. 1922.
Small, caespitose, epiphytic herbs up to about 15 cm. tall. Stems slender,
short, produced into a long peduncle above, bearing 2-3 leaves. Leaves 3-5 cm.
long and 0.6-1.2 cm. broad, linear-lanceolate to lanceolate, acute. Inflorescence
a few-flowered raceme at the apex of the peduncle; flowers small, purple-brown
with a white lip. Dorsal sepal about 5 mm. long and 1.5 mm. broad, lanceolate,
acute, Lateral sepals about 4.5 mm. long and 2 mm. broad, ovate-lanceolate,
slightly oblique, acute. Petals about 4 mm. long and 1 mm. broad, elliptic, ob-
scurely serrulate, acute. Lip unguiculate; claw adnate to the column; lamina
about 3 mm. long and 4 mm. broad, entire, reniform-cordate, acute, fleshy; disc
provided with a single sublamellate callus near the middle.
anama.
CHIRIQUÍ: “Chiriquí”, Powell 250.
Known to us only by the inadequate specimen cited.
64. EPIDENDRUM PSEUDEPIDENDRUM Reichb. f. Xenia Orch. 1:160, /. 53. 1856;
Hook. f. in Bot. Mag. 97:1. 5020. 1871.
Pseudepidendrum spectabile Reichb. f. in Bot. Zeit. 10:733. 1852, not Focke nor Reichb. f.
Caespitose epiphytic herbs up to about 1 m. tall. Stems slender, simple, leafy
above, naked below. Leaves 6-20 cm. long and 1.5—4.5 cm. broad, oblanceolate,
acute, coriaceous. Inflorescence 1 or more, 1- to 3-flowered terminal racemes borne
on a more or less elongated peduncle, the peduncle covered at the base with several
imbricated bracts; flowers relatively large, sepals and petals greenish, lip orange.
Dorsal вера! 26-30 mm. long and 3-5 mm. broad, oblanceolate, acute. Lateral
sepals 22-32 mm. long and 5-7 mm. broad, oblanceolate, acute, sometimes obtuse.
Petals 22-30 mm. long and 2-3 mm. broad, linear-oblanceolate, obtuse. Lip
unguiculate; the long claw adnate to the column; lamina 13-17 mm. long and
15-22 mm. broad, retuse, transversely oblong or subquadrate to suborbicular-
flabellate, entire (or obscurely 3-lobed), crenulate, dentate or lacerate; disc pro-
vided with 2 short lamellate calluses at the apex of the column and a longitudinal
1- to 5-ridged callus-thickening along the median line, from base to apex.
Costa Rica and Panama.
CHIRIQUÍ: "auf Ficus-Báumen der Cordilleren von Chiriquí”, alt. 1200 m., Warscewicz.
Apparently one of the prettier Epidendrums occurring in Panama.
65. EPIDENDRUM PUDICUM Ames, Sched. Orch. 6:71, fig. 10. 1923.
Small, caespitose, epiphytic herbs up to about 20 cm. tall. Stems slender, ob-
scurely fractiflex, covered with the persistent, pergameneous, amplexicaul leaf-
sheaths. Leaves 15-30 mm. long and 1-3 mm. broad, linear, obtuse, fleshy.
Inflorescence a 1- to few-flowered subumbel. Dorsal вера! 8-10 mm. long and
1.5—2.5 mm. broad, linear-oblanceolate, acute. Lateral sepals similar to the dorsal
but slightly arcuate. Petals about 8 mm. long and nearly 1 mm. broad near the
apex. Lip unguiculate; claw adnate to the column; lamina about 5 mm. long and
(299)
[Vor. 33
368 ANNALS OF THE MISSOURI BOTANICAL GARDEN
7 mm. broad, subtriangular-cordate to reniform, obscurely 3-lobed, acute, pro-
vided with 2 slightly divergent submammillate calluses at the base.
Panama.
CHIRIQUÍ: Palo Alto, alt. 1360 m., Powell 310, 3412.
Closely allied to Epidendrum difforme Jacq.
Fig. 133. Epidendrum pudicum
(300)
19461
FLORA OF PANAMA (Orchidaceae) 369
66. EPIDENDRUM RAMOSUM Jacq. Enum. Pl. Carib. 29. 1760; Select. Stirp. Am.
221, 2. 132. 1763; Ames, Hub. & Schweinf. in Bot. Mus. Leafl. Нагу. Univ.
2:45. 1934.
Epidendrum flexicaule Schltr. in Већ. Bot. Centralbl. 36, Abt. 2:403. 1918.
Epidendrum modestiflorum Schltr. in Fedde Rep. Sp. Nov. Beih. 19:213. 1923.
Erect or scandent epiphytic herbs, perhaps up to a meter or more long. Stems
slender, simple or branched, covered with the leaf-sheaths, becoming naked with
age. Leaves 1-12 cm. long and 0.2-1.5 cm. broad, linear to lanceolate to oblong-
elliptic, obtuse and the apex usually unequally bilobed. Inflorescence short and
few- to several-flowered from a fractiflex raceme; bracts scarious, conspicuous;
flowers small, variable in size. Sepals 4.5—13 mm. long and 1.5—3 mm. broad,
linear-lanceolate to elliptic-oblong, occasionally carinate dorsally. Petals 4.5—13
mm. long and 0.8-2.3 mm. broad, linear to linear-spatulate, acute or obtuse. Lip
unguiculate; claw adnate to the column; lamina 4-9 mm. long and 2-5 mm.
broad, simple or subtrilobate, triangular-cordate to lanceolate-cordate, acute or
obtuse, provided with a longitudinal callus which is bifurcate at the base.
Mexico to Panama, the West Indies and in South America to Brazil and Peru.
кішілі: Lino Hill, alt. 1550 m., гуе 322. COCLÉ: El Valle de Antón, alt.
600- 1000 m., Allen 1244, 2827, 2836.
66a. EPIDENDRUM RAMOSUM Jacq. var. ANGUSTIFOLIUM (Cogn.) L. Wms. in
Ann. Missouri Bot. Gard. 28:422. 1941.
Epidendrum imbricatum Lindl. var. angustifolium Cogn. in Mart. Fl. Bras. 35:171. 1898,
as var. angustifolia.
Epidendrum imbricatum Lindl. Gen. & Sp. Orch. РІ. 110. 1831, non Lam
мае inne Cogn. in Bull. Herb. Boiss. II, 2: ed figs. 1-3. 1902, non Forst. f.
ескер
Epidend rum анон Schltr. in Вей. Bot. бағын 36, Abt. 2:459. 1918.
Epidendrum santaclarense Ames, Sched. Orch. 4:49. 19
Epidendrum ramosum Jacq. var. imbricatum Ames, Hub. & Schweinf. in Bot. Mus. Leafl.
H Univ. 2:47. 1934
Intergrades with the species but usually varies in having thicker stems, fine
maculations on the leaf-sheaths and bracts, the bracts more tightly imbricated
and conduplicate; leaves usually broader and larger.
Guatemala, Costa Rica, Panama, the West Indies, Brazil.
DEL TORO: Río Cricamola, between Finca St. Louis and Konkintoé, alt. 10-15
m., ‘Woodson, Allen 9 Seibert 1808.
67. EPIDENDRUM REPENS Cogn. іп Fedde Кер. Sp. Хоу. 7:122. 1909.
Small, repent or pendent, epiphytic herbs up to about 5 dm. long. Stems slen-
der, much branched, leafy, covered with the persistent leaf-sheaths, slightly an-
cipitous, often flexuose. Leaves 6-25 mm. long and 2-6 mm. broad, linear-oblong
to oblong to ovate, obtuse, coriaceous, usually obtuse and obscurely and unequally
bilobed at the apex. Inflorescence a single flower borne terminally on the stem,
subtended by one or more short scarious bracts; flowers small, inconspicuous,
maroon. Dorsal sepal 7-8 mm. long and about 2 mm. broad, elliptic-oblong or
(301)
[Vor. 33
370 ANNALS OF THE MISSOURI BOTANICAL GARDEN
E
а
———= С
——— —
==
Е
: = , “> рс É E: ii
Fig. 134. Epidendrum rigidum
(302)
1946]
FLORA OF PANAMA (Orchidaceae) 371
lanceolate, acute. Lateral sepals 7-8 mm. long and 1.8-3 mm. broad, elliptic-
oblong to lanceolate-ovate, acute. Petals 6-7 mm. long and 0.7-1.5 mm. broad,
linear, acute. Lip unguiculate; claw adnate to the column; lamina about 4 mm.
long and as broad, triangular-cordate to cordate, simple, fleshy, ecallose or pro-
vided with a short, fleshy, lamellate callus toward the apex.
Mexico, Guatemala, Costa Rica, Panama, the West Indies and Venezuela.
cHIRIQUÍ: vicinity of Cerro Punta, alt. 2000 m., Allen 1530.
68. EPIDENDRUM RIGIDUM Jacq. Enum. РІ. Carib. 29. 1760; Select. Stirp. Am.
232, 85 ТЫ 1763.
Epidendrum cardiophorum Schltr. in Je hang Sp. Nov. 9:214. 1911.
Spathiger rigidus Small, Fl. Miami, p. 55. 1913.
Repent epiphytic herbs up to about 35 cm. long. Stems slender, simple, cov-
ered with the amplexicaul leaf-sheaths, sometimes appearing ancipitous. Leaves
3-13 cm. long and 0.4-2 cm. broad, linear to lanceolate-oblong or oblong, obtuse,
coriaceous, the apex usually unequally bilobed. Inflorescence a strict raceme up to
about 15 cm. long, few- to several-flowered; bracts up to about 15 mm. long,
ovate, acute or obtuse, semi-equitant or usually the base surrounding the peduncle,
usually with a median keel which is decurrent onto the peduncle; flowers usually
greenish. Dorsal вера! 5.5-9 mm. long and 2-3 mm. broad, oblong to lanceolate-
ovate, obtuse. Lateral sepals 5-9 mm. long and 2-4 mm. broad, ovate-lanceolate
to obovate, oblique, obtuse. Petals 5-9 mm. long and 1.2-2 mm. broad, ovate-
lanceolate to obovate, oblique, obtuse. Lip unguiculate; claw adnate to the
column; lamina 4-6 mm. long and about as broad, cordate-orbicular, obtuse or
obscurely retuse, lateral margins revolute; disc fleshy, with 2 mammillate calluses
at the apex of the column.
Florida, Mexico to Panama, the West Indies, tropical and subtropical South
America.
ZONE: near Salamanca Hydrographic Station, Rio Pequeni, Dodge, Steyer-
mark 8 wir s. п.; between Tumba Vieja and Salamanca, Steyermark © Allen 16740.
A widespread species which is surprisingly constant in characters.
68a. EPIDENDRUM RIGIDUM Jacq. var. angustise$mentum Г. Wms. var. nov.!
Vegetatively similar to the species. Dorsal вера! 9-10 mm. long and 2-3 mm.
broad, lanceolate, acute. Lateral sepals 9—10 mm. long and 2.5—3.5 mm. broad,
lanceolate, oblique, acute. Petals 7.5-9 mm. long and 0.5—1 mm. broad, subfili-
form to linear, acute. Lip unguiculate; claw adnate to the column; lamina 5-6
mm. long and 3-3.5 mm. broad, oblong-ovate, obtuse, truncate at the base, fleshy;
disc with two mammillate calluses at the apex of the column and with one or
more inconspicuous callus ridges.
Sepalum rond бағалы қыс acutum. Sepala lateralia lanceolata, obliqua, acuta. Petala sub-
anc" vel lin acuta. Lamina labelli oblongo-ovata, obtusa, basi truncata, carnosa; discus
prope basim bici
(303)
(Vor. 33
372 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Panama.
OCAS DEL TORO: epiphyte, Nances Cay Island, Wedel 586; Water = Wedel 851;
Water Valley, vicinity of Chiriqui Lagoon, Wedel 1708 (түре Herb. es).
The variety differs from the species in its narrower floral segments and dif-
ferently shaped lip.
69. EPIDENDRUM SANCHO Ames, Sched. Orch. 4:48. 1923; loc. cit. 10:70.
Repent epiphytic herbs up to about 4 dm. long. Stems small, branched, often
profusely so, usually consisting of a principal stem and smaller elongated branches,
covered with the verrucose or rugose amplexicaul leaf-sheaths. Leaves 0.6—8 cm.
long and 0.1—0.6 cm. broad, linear or linear-oblong, obtuse, apex usually unequally
bilobed. Inflorescence of terminal few-flowered racemes; rachis fractiflex. Dorsal
sepal 5-8 mm. long and 1.2-2 mm. broad, lanceolate, acute. Lateral sepals 5—8
mm. long and 1.2-2.5 mm. broad, lanceolate, acute. Petals 4,5-7 mm. long and
0.5-1 mm. broad, linear-elliptic or linear, acute. Lip unguiculate; claw adnate to
the column; lamina 3.5—5 mm. long and 2-2.5 mm. broad, triangular-cordate or
narrowly so, fleshy, the basal auricles usually erect; the disc appearing excavated,
provided with a small trilobulate callus at the base.
Costa Rica and Panama.
CHIRIQUÍ: rain forest, Boquete District, alt. 1800 m., Davidson 114, 348.
70. EPIDENDRUM SCHLECHTERIANUM Ames, Sched. Orch. 7:9, fig. r. 1924.
ара discolor Lindl. in Bot. Reg. 18:Ё. 1541. 1832, non Epidendrum discolor Rich.
Epidendrum brevicaule Schltr. in Fedde Rep. Sp. Nov. Beih. 17:30. 1922, non Schltr.
ы congestoides Ames & Schweinf. Sched. Orch. 10:61. 1930.
Dwarf, densely caespitose herbs up to 8 cm. tall. Stems short, covered with
the equitant bases of the leaves. Leaves 1-3 cm. long апа 0.2-1 cm. broad, linear-
oblong to oblong-oval, obtuse, approximate, semi-equitant, spreading, fleshy. In-
florescence of 1 to few sessile flowers in the axils of the terminal leaves, shorter
than the leaves to slightly exceeding them; flowers large for the plant. Dorsal
sepal 8-20 mm. long and 3-4.5 mm. broad, lanceolate to oblanceolate, acute or
acuminate. Lateral sepals 8-18 mm. long and 2-4.5 mm. broad, lanceolate to
ovate-lanceolate, oblique, acute, serrulate-keeled along the mid-nerve dorsally,
connate at the base with the claw of the lip. Petals 8-18 mm. long and 1-2.5
mm. broad, elliptic-linear to linear-lanceolate, entire or denticulate, acute. Lip
8-20 mm. long, long-unguiculate; claw adnate to the column, usually slightly
longer than the column; lamina 4-8 mm. long and as broad, suborbicular-reni-
form to oblong-quadrate, apiculate, lobes more or less erect, fleshy. Clinandrium
fimbriate-denticulate.
Mexico to Panama, Jamaica, Surinam, Brazil and Peru.
CANAL ZONE: near Vigia and San Juan, Dodge, Steyermark & Allen 16562; Rio
Azote Caballo, 66-70 m., Dodge, Steyermark & Allen 16864; San Juan, Powell 18, 3465.
(304)
19461
FLORA OF PANAMA (Orchidaceae) 373
EPIDENDRUM jw
Duos um 4 EZ mn es
Fig. 135. Epidendrum Schlechterianum
71. EPIDENDRUM SCHUMANNIANUM Schltr. in Fedde Rep. Sp. Nov. 9:215. 1911;
Allen in Am. Orch. Soc. Bull. 10:359, figs. 1942
Tall, caespitose, epiphytic herbs up to a meter or more nak Stems up to
about 8 mm. in diameter, leafy above, covered with the maculate or verru-
cose leaf-sheaths, becoming naked and vernicose with age. Leaves 1.5-9 cm. long
and 1-3.5 cm. broad, lanceolate to elliptic-oblong, obtuse or acutish, coriaceous.
Inflorescence simple or usually a lax many-flowered panicle. Flowers multicolored
and rather pretty. Dorsal вера! 8-12 mm. long and 3.5-4.5 mm. broad, oval to
broadly oblanceolate, obtuse or acutish. Lateral sepals 8-12 mm. long and 3.5—5
mm. broad, oblanceolate to oval, obtuse or acute, oblique. Petals 8-15 mm. long
and 5.5-8 mm. broad, obovate or obovate-spatulate, obtuse. Lip unguiculate;
claw adnate to the basal half of the column; lamina 9-15 mm. long and 7.5-12
(305)
[Vor. 33
374 ANNALS OF THE MISSOURI BOTANICAL GARDEN
mm. broad, 3-lobed; the lateral lobes small, oblong, obtuse, explanate; terminal
lobe obovate-cuneate, bilobulate or deeply bifid, anterior margins entire to crenu-
late; disc provided with 2 small submammillate calluses at the base and with 1—3
longitudinal callus-thickenings.
Costa Rica and Panama.
CLÉ: hills north of EI Valle de pee vicinity of La Mesa, alt. 1000 m., Allen
xt wer El Valle, alt. 670 m., Dudak s
72. EPIDENDRUM SCULPTUM Reichb. f. in Bonplandia 2:89. 1854.
Epidendrum colonense Ames, Sched. Orch. 1:14. 1922.
Pendent or repent epiphytic herbs up to about 50 cm. long. Stems simple or
branched, densely leaved, covered with the chartaceous leaf-sheaths or becoming
naked with age. Leaves 2-4.5 cm. long and 1-1.8 cm. broad, oblong or oblong-
lanceolate, obtuse, retuse, coriaceous. Inflorescence with 1—3, usually 2, terminal
flowers subtended by spathaceous bracts; flowers green ог greenish-yellow. Dorsal
sepal 10-14 mm. long and 2.5—3 mm. broad, lanceolate to oblong-lanceolate, ob-
tuse. Lateral sepals 10-15 mm. long and about 3 mm. broad, oblong-lanceolate,
oblique, acute. Petals 9-14 mm. long and 2-3 mm. broad, elliptic-linear to nar-
rowly oblanceolate, oblique, obtuse. Lip unguiculate; claw adnate to the column;
lamina 9-13 mm. long and 5.5-7 mm. broad, lanceolate-ovate, acute, 3-lobed;
lateral lobes small, rounded, reaching to about the middle of the lip; mid-lobe
lanceolate; disc provided with two callus-like thickenings at the base on the lateral
lobes and with two longitudinal ridges along the median line and reaching to about
the middle of the lip.
Possibly Honduras and Costa Rica. Panama, British and Dutch Guiana.
L ZONE: near Vigia and San Juan, Dodge, Steyermark & Allen 16583; Gatün
Lake, Powell 116, 3407, 3410; between Fort Clayton and Corozal, Standley 20000. COLON:
Rio Indio de Fato, Pittier 4265
A specimen from Costa Rica, presumed to be this species, has an entire lip.
73. EPIDENDRUM STROBILIFERUM Reichb. f. in Nederl. Kruidk. Arch. 4:333.
1859
idendrum verecundum Schltr. in Fedde Rep. Sp. pt Beih. 17:42. 1922.
ru ied strobiliferus Small, Man. Southeast. Fl. 390. 1933
Small, repent or caespitose, epiphytic herbs up to about 2 dm. long. Stems
slender, simple or branched, covered with amplexicaul leaf-sheaths. Leaves 8-45
mm. long and 2-10 mm. broad, from ligulate to elliptic-linear to linear-lanceolate,
obtuse or acute, coriaceous, apex usually unequally bilobed. Inflorescence short,
few-flowered, flexuose; bracts 5-8 mm. long, cucullate, obtuse, chartaceous;
flowers small, inconspicuous. Dorsal вера! 3.5—5 mm. long and 1.2-1.5 mm.
broad, linear-lanceolate, acute. Lateral sepals 3.5-5 mm. long and 1.8-2 mm.
broad, ovate-lanceolate, oblique, acute. Petals 3-4.5 mm. long and 0.5-1 mm.
broad, linear-oblanceolate, obtuse or acute. Lip 3-3.5 mm. long and 1.5-3 mm.
broad, lanceolate-cordate to ovate-cordate, acute, ecallose. Ovary with a small
(306)
1946]
FLORA OF PANAMA (Orchidaceae) 375
^N SN | ie
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na
| 222
-NWE WA
CENE-
у:
ie
+t
as
М SP
М А
SMS PS
SS ENTRIES MSS
АА
4
да (ма
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M cov nii nnee
Fig. 136. Epidendrum strobiliferum
(307)
[Vor. 33
376 ANNALS OF THE MISSOURI BOTANICAL GARDEN
subscrotiform sac at the apex near the attachment of the lip.
Florida, Mexico to Panama, the West Indies, Venezuela, the Guianas, Brazil
and Peru.
CANAL ZONE: foothills east of Panama City, San Juan Range, and San Juan, Powell
209, 350, 3435, 3453; near Summit, Standley 25747. PANAMA: Rio Tecümen, north of
Chepo Road, alt. 30 m., Hunter & Allen 2 224.
74. EPIDENDRUM SUBNUTANS Ames & Schweinf. in Sched. Orch. 10:73. 1930.
Caespitose, epiphytic herbs up to about 85 cm. tall. Stems slender, simple,
covered with the amplexicaul leaf-sheaths, naked below. Leaves 4.5-18 cm. long
and 0.8—4 cm. broad, elliptic-lanceolate to oval or ovate, acuminate or acute. In-
florescence terminal, paniculate, many-flowered; flowers relatively small. Dorsal
sepal 5-6 mm. long and 1.2-2 mm. broad, oblanceolate, acute, sometimes papillose
dorsally. Lateral sepals 5-6 mm. long and 1.7-2 mm. broad, oblanceolate to
elliptic-oblanceolate, oblique, sometimes papillose dorsally. Petals 4-5 mm. long
and 0.1-0.5 mm. broad, filiform to narrowly linear-oblanceolate, obtuse. Lip
unguiculate; claw adnate to the column; lamina 2.5-3.5 mm. long and 4.5-5.5
mm. broad, 3-lobed, subreniform in outline; lateral lobes explanate, larger than
the terminal lobe, dolabriform, margins crenate or undulate; mid-lobe subquad-
rate, retuse and with erect to spreading lobules; disc provided with a large oval
central callus made up of 3 parallel ridges, the middle one extending to the apex
of the lip.
Costa Rica and Panama.
Қ yore Bajo series EH District, alt. 1800 m., сее 43, II 13; “Chiriqui”,
8; vicinity of Bajo Mona and Quebrada Саш, alt. .» Woodson %
E bei 541, 554; Bajo Chorro, alt. 1900 m., Woodson & в 648.
Very closely allied to Epidendrum paniculatum Ruiz & Pavon, of which it may
be but a small-flowered variety.
75. EPIDENDRUM TERETIFOLIUM Sw. Nov. Gen. & Sp. Prodr. 121. 1788.
Epidendrum teres Reichb. f. in Bonplandia 3:220. 1855, non Thunb.
Epidendrum teretifolium Sw. var. Powellianum Schltr. in Fedde Rep. Sp. Nov. Beih.
7:41. 1922.
Caespitose or subrepent, epiphytic herbs up to about 35 cm. tall. Stems slen-
der, simple, erect or ascending, covered with the amplexicaul leaf-sheaths, even in
age. Inflorescence a single flower or a few-flowered fascicle borne on the apex of
a peduncle which is up to about 8 cm. long, and subtended by chartaceous sheaths
and 1 or 2 terminal leaves which are borne near the apex of the peduncle; the
peduncle more or less ancipitous; flowers greenish or yellowish, segments fleshy.
Dorsal вера! 8-10 mm. long and about 2 mm. broad, linear-lanceolate, acute.
Lateral sepals 8-11 mm. long and 1.5-2 mm. broad, elliptic to lanceolate, slightly
oblique, acute. Petals 4-6 mm. long and 1.2—1.5 mm. broad, oblanceolate, obtuse,
the apex fleshy and thickened. Lip unguiculate; claw adnate to the basal part of
the column; lamina 6-8 mm. long and 1-2 mm. broad, 3-lobed, elliptic; lateral
(308)
19461
FLORA OF PANAMA (Orchidaceae) 377
lobes small, rounded, erect, at about the middle of the lip; terminal lobe with a
prominent lamellate callus which is higher than the lobe is broad, acute; disc
provided with a small lamellate callus toward the base and a large one on the
terminal lobe.
Mexico, Guatemala, Honduras, Nicaragua, Panama, the West Indies and
Venezuela.
CHIRIQUÍ: Palo Alto, alt. 1200-1400 m., Powell 240.
76. EPIDENDRUM TETRACEROS Reichb. f. in Bot. Zeit. 10:733. 1852.
Much-branched epiphytic herbs, probably up to a meter or more long. Stems
slender, covered with the scarious leaf-sheath bases when young, becoming naked
with age, bearing leaves toward the ends of the stems. Leaves 3-7 cm. long and
0.2—0.8 cm. broad, linear to linear-lanceolate, acute. Inflorescence a few-flowered
raceme, about as long as or shorter than the subtending leaves. Flowers with sepals
and petals greenish, the lip white with purple-red markings. Dorsal sepal about
12-16 mm. long and 4-5 mm. broad, elliptic-lanceolate, acute or acuminate.
Lateral sepals similar to the dorsal but slightly oblique. Petals 12-14 mm. long
and 3-5 mm. broad, linear-oblanceolate, acute. Lip unguiculate; claw adnate to
the column; lamina 10-12 mm. long and 11-14 mm. broad, fleshy, suborbicular,
undulate-crenate, provided with 2 small lamellate calluses under the apex of the
column (hidden by the clinandrium) and a large, thick, central callus made up of
3 or more ridges. Column winged laterally at the apex and with two apical
stelidiar arms above the clinandrium.
Costa Rica and Panama.
CHIRIQUÍ: "Chiriqui", Warscewicz; near El Boquete, Pittier 3115.
77. EPIDENDRUM TRACHYTHECE Schltr. in Fedde Rep. Sp. Nov. 3:249. 1907.
Small, caespitose or repent, epiphytic herbs up to about 25 cm. long. Stems
slender, leafy, simple or usually much branched, covered with the verrucose or
rugose amplexicaul leaf-sheaths. Leaves 1-4.5 cm. long and 0.3—0.5 cm. broad,
ligulate, obtuse, the apex retuse and unequally bilobed. . Inflorescence short, a sin-
gle flower, or usually a few-flowered flexuose raceme exceeded by the subtending
leaves; bracts up to about 5 mm. long, chartaceous, cucullate, oblong to ovate.
Flowers small, inconspicuous, greenish-yellow to white. Dorsal вера! 3—3.5 mm.
long and 1.5-2 mm. broad, oblong to oval, obtuse or acute. Lateral sepals 3-3.5
mm. long and 2,2-3 mm. broad, oblong-ovate to suborbicular, slightly oblique,
acute or obtuse. Petals 3—3.2 mm. long and 0.7-0.8 mm. broad, linear-oblong to
oblanceolate, acute or obtuse. Lip unguiculate; claw adnate to the lower % of
the column; lamina 2-2.5 mm. long and 3-3.5 mm. broad, transversely oval to
subreniform, often short-apiculate; disc provided with a bilobate submammillate
callus near the middle, the lobes sometimes somewhat separated.
Guatemala, Costa Rica and Panama.
CHIRIQUÍ: Bajo Chorro, Boquete District, alt. 1800 m., Davidson 124.
(309)
е,
[Vor. 33
378 ANNALS OF THE MISSOURI BOTANICAL GARDEN
78. EPIDENDRUM TRIANGULABIUM Ames & Schweinf. in Sched. Orch. 10:75 1930.
Small, caespitose or repent, epiphytic herbs up to about 20 cm. long. Stems
slender, simple or branched, covered with the loose, keeled, amplexicaul leaf-
sheaths, naked with age. Leaves 1.3-2.5 cm. long and 0.5-1.2 cm. broad, elliptic
to oval, obtuse, coriaceous, apex obscurely bilobed. Inflorescence a single terminal
or lateral flower. Flowers apparently cleistogamous. Dorsal sepal about 12 mm.
long and 3 mm. broad, lanceolate, acute. Lateral sepals about 12 mm. long and
3 mm. broad, lanceolate, acute, oblique. Petals about 12 mm. long and 2 mm.
broad, elliptic, acute, oblique. Lip unguiculate; claw adnate to the column;
lamina about 6 mm. long and 9 mm. broad, sagittate-triangular, the apex of the
lateral extensions and the terminal point acute.
Costa Rica and Panama.
COCLÉ: vicinity of El Valle de Antón, alt. um 1000 m., Allen 1245. ?PANAMÁ: near
top of Cerro Campana, alt. 800-1000 m., Allen 2223.
We have seen no completely satisfactory material of this species.
79. EpIDENDRUM Warszewiczu Reichb. f. in Bot. Zeit. 10:732. 1852; in Xenia
Orch. 1:69, Ё. 26. 1854.
Small, erect, epiphytic herbs with relatively large flowers. Stem slender, few-
leaved. Leaves (about 12-15 mm. long?) linear-ligulate, obtuse, apex bilobed.
Inflorescence reflexed, many-flowered (illustration shows five). Perianth segments
oblong, acute (in illustration apparently lanceolate, acute or acuminate); lip
dilated in front, minutely denticulate, 3-lobulate (illustration shows: about 30
mm. long and 17 mm. broad, this presumed to be natural size, obovate, obscurely
3-lobed, denticulate margins); disc with 2 erect teeth (calluses) at the base and
3 elevated longitudinal lines.
Costa Rica and Panama according to Warscewicz.
PANAMÁ: "Costa Rica, Veraguas, Chiriqui", Warscewicz.
The collection locality of the single specimen known to exist was given by
Reichenbach, in the second publication cited, as we have shown it. The descrip-
tion was taken from the original publication and from the plate cited.
80. EPIDENDRUM LIVIDUM Lindl. in Bot. Reg. 24: Misc. p. 51. 1838; Ames, Hub.
& Schweinf. Genus Epidendrum in U. S. & Mid. Am. 118. 1936.
Costa Rica, Venezuela and Colombia. Reported from Panama in cultivation.
28. DIACRIUM Lindl. ex Benth.
Diacrium Lindl. ex Benth. in Jour. Linn. Soc. Lond. 18:312. 1881; Benth. &
Hook. Gen. Pl. 3:526. 1883; Pfitzer in Engler & Prantl, Nat. Pflanzenf. II,
6:146. 1888. |
Epidendrum subg. Diacrium Lindl. in Hook. Jour. Bot. 3:81. 1841.
Caespitose epiphytic or semi-epiphytic herbs. Stems pseudobulbose, solid or
hollowed by insect action. Leaves at or near the apex of the pseudobulbs, few.
(310)
19461
FLORA OF PANAMA (Orchidaceae) 379
==
АЕ
>
==.
222
2222
CGE PEA
р „Зәбт
SS
2222
Fig. 137. Diacrium bilamellatum
Inflorescence terminal, racemose, borne at the end of an elongated peduncle, few-
to several-flowered. Sepals similar, free, spreading. Petals similar to the sepals.
Lip free from the column and spreading, 3-lobed, provided with two horn-like
processes toward the base which are hollow and open from the underside of the lip.
Column short, footless, produced laterally into wings. Anther terminal, opercu-
late, incumbent, 2-celled, the cells divided by a longitudinal septum; pollinia 4,
laterally compressed, each one provided with a granular, viscid appendage.
Diacrium is closely allied to Epidendrum, from which it is easily, if super-
ficially, distinguished by the hollow horn-like processes on the lip. A small genus
of four or five species.
(311)
[Vor. 33
380 ANNALS OF THE MISSOURI BOTANICAL GARDEN
1. DIACRIUM BILAMELLATUM (Reichb. f.) Hemsl. іп Godm. & Salvin, Biol.
Centr.-Am. Bot. 3:222. 1883.
Epidendrum bilamellatum Reichb. f. in Walp. Ann. 6:345. 1862.
Diacrium bilamellatum var. Reichenbachianum Schltr. in Fedde Rep. Sp. Nov. Beih. 17:47,
in text. 1922.
Coarse, caespitose, epiphytic herbs up to about 60 cm. tall. Stems swollen,
solid or hollowed by insects, cylindric to fusiform, provided with 1-4 leaves at the
apex. Leaves 6-20 cm. long and 0.6-2.5 cm. broad, ligulate to ligulate-lanceolate,
obtuse, fleshy. Inflorescence a short several-flowered raceme borne near the top
of an elongated peduncle; peduncle up to 45 cm. long, provided with amplexicaul,
chartaceous sheaths. Flowers pedicellate, one or few open at one time, perhaps
often cleistogamous. Dorsal вера! 12-15 mm. long and about 6 mm. broad,
oval, acute. Lateral sepals 11-16 mm. long and 5-6 mm. broad, elliptic-oval to
lanceolate-ovate, acute. Petals 11-14 mm. long and 7-8 mm. broad, broadly
obovate, short-unguiculate, acute. Lip free from the column and spreading;
lamina 11-14 mm. long and 4-6 mm. broad, lanceolate-ovate in outline, 3-lobed,
the lateral lobes small and often obscure above the middle of the lip, apices
rounded; disc provided with two large lamellate horn-like processes near the
middle, these processes hollow, 2-4 mm. high.
Guatemala, British Honduras, Costa Rica, Panama and Venezuela.
BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Wedel 2100. CANAL ZONE: Chagres,
Behr; open woods east of Panamá City, Powell 67; Culebra, Pittier 3406. PANAMA:
swamp between El Jagua Hunting Club on Río Jagua and El Congor Hill, Hunter 9 Allen
473; San José and Trapeche Islands, Pearl Islands, Miller 1800, 1000. WITHOUT LOCALI-
TIES: Ames s. n.; Cowell 160; Jobnston 703; Killip 3394. |
This species is said to have its stems often inhabited by ants. It is a lowland,
coastal species.
29. CATTLEYA Lindl.
САТТІЕҮА Lindl. Collect. Bot. 7. 33 © t. 37. 1824.
Epiphytic herbs with thick, fleshy or pseudobulbose stems, 1- to 2-foliate at
the apex. Leaves coriaceous or fleshy, most often thick. Inflorescence terminal,
usually subtended by a spathaceous bract; raceme simple; flowers often large and
showy, usually few. Sepals subequal, free. Petals usually broader than the sepals.
Lip sessile, broad, erect, free from or slightly adnate to the column, sides erect
and usually enfolding the column. Column long, wingless, often somewhat
arcuate; anther terminal, operculate, incumbent, each cell with a longitudinal
septum; pollinia 4, parallel, ceraceous, broad, somewhat compressed.
One variety in Panama, about 30 in Central and South America. Cattleya is
the most widely cultivated of the orchids. It hybridizes easily with Laelia, Brassa-
vola and Sophronitis and the hybrid genera formed by these crosses.
(312)
19461
FLORA OF PANAMA (Orchidaceae) 381
|
ul
'
Fig. 138. Cattleya Skinneri var. autumnalis
1, CATTLEYA SKINNERI Batem. var. AUTUMNALIS Allen in Ann. Missouri Bot.
Gard. 29:345. 1942.
Epiphytic or semi-terrestrial herbs up to 4 dm. tall. Pseudobulbs 15-30 cm.
long and 1-3 cm. in diameter, cylindric to fusiform, 2-, rarely 1-, leaved. Inflores-
сепсе a simple few-flowered raceme; spathe 5-10 cm. long, pergameneous. Leaves
10-15 cm. long and 3.5—5.5 cm. broad, elliptic to oblong-lanceolate, obtuse,
coriaceous. Sepals 3-5 cm. long and 0.8-1.4 cm. broad, elliptic to broadly ob-
lanceolate, acute. Petals 3-5 cm. long and 1-2.5 cm. broad, lanceolate to nar-
rowly ovate or oval, obtuse or acute. Lip 3—4.5 cm. long and 2-3 cm. broad,
oval to obovate, retuse, usually darker in color than the sepals and petals, enfolding
the column.
Panama.
(313)
[Vor. 33
382 ANNALS OF THE MISSOURI BOTANICAL GARDEN
PANAMA: vicinity of Bejuca, alt. 15 m., Allen 2668; east of city, Powell 16. CANAL
ZONE: Сайоп Quebrada, Pittier 6828,
The Panamanian variety of a species distributed from Mexico to Costa Rica
and the only known Cattleya in Panama. Autumn flowering, and the flowers
smaller than in the species.
30. LAELIA Lindl.
LAELIA Lindl. Gen. & Sp. Orch. Pl. 115. 1831; Benth. & Hook. Gen. Pl. 3:533.
1883; L. Wms. in Darwiniana 5:74. 1941.
Schomburgkia Lindl. Sert. Orch. /. 10. 1838.
Epiphytic herbs often with showy flowers and ancipitous, terete, solid or
hollow pseudobulbs. Sepals subequal, free, spreading, flat or undulate. Petals
similar to the sepals, sometimes broader. Lip free from or slightly connate with
the column, 3-lobed (or nearly simple), the lateral lobes enfolding the column or
more or less explanate; disc smooth or lamellate; column long or short, winged or
wingless. Anther operculate, incumbent, each cell divided into two by a septum
and with each cell sometimes again divided by a more or less perfect septum;
pollinia 8, 4 in each cell of the anther, ovate or compressed laterally, ceraceous.
The genus is to be found rather widely over tropical and subtropical America,
where it is often exceedingly abundant. Some species are prized by orchid growers
and are useful in hybridization. There are about 50 species in the genus.
a. Pseudobulbs less than 8 Á long; sepals and petals р1апе...................... . L. RUBESCENS
aa. — bulbs more than cm. long; sepals and petals үтікте” ог
. L. LUEDDEMANII
1. LAELIA RUBESCENS Lindl. in Bot. Reg. 26: Misc. p. 20, Ё. 41. 1840.
Laelia peduncularis Lindl. loc. cit. 28: Misc. p. 9. 1842.
Caespitose epiphytic herbs up to 60 cm. tall. Pseudobulbs 1.5—7 cm. long and
1.5—4 cm. broad, oval to orbicular, ancipitous, 1-, rarely 2-,leaved. Leaves 4—20
cm. long and 2-4.5 cm. broad, elliptic, elliptic-lanceolate to ovate, obtuse, cori-
aceous. Inflorescence a long-pedunculate, few- to several-flowered raceme, termi-
nal or rarely appearing laterally оп undeveloped bulbs; bracts up to 1 cm. long,
chartaceous, cucullate; flowers small to fairly large, white, or white with the base
of the lip lavender to pink, or lavender throughout. Sepals 2—4.5 cm. long and
0.2-0.7 cm. broad, linear-lanceolate to elliptic, obtuse or acutish. Petals 2—4.5
cm. long and 0.3—1.5 cm. broad, elliptic or lanceolate, obtuse or acute. Lip
1.5-3.5 cm. long and up to 1.5 cm. broad, shorter than the sepals and petals, 3-
lobed, provided with several inconspicuous longitudinal ridges; lateral lobes short,
rounded and obtuse, enfolding the column; terminal lobe oblong-oval, blunt or
retuse.
Mexico, Guatemala, Salvador, Nicaragua, Costa Rica and Panama.
CHIRIQUÍ: “М. №. corner of Chiriquí, near the Gulf of Dulce," cultivated by Powell
64.
(314)
19461
FLORA OF PANAMA (Orchidaceae) 383
139. Laelia rubescens
(Flowers a. есін аге normally resupinate)
2. Larria LuEDDEMAND (Prill.) Г. Wms. in Ann. Missouri Bot. Gard. 27:282.
1940.
Li iei s Lueddemanii Es in Jour. Soc. Imp. Hort. Paris 8:275. 1862; Rolfe in
t. Mag. 138: 1. 8427.
Large, caespitose, M herbs up to a meter or more tall when in flower.
Pseudobulbs 18-30 cm. long and 2-3 cm. broad, apparently somewhat ancipitous,
largest toward the top, bifoliate. Leaves 20-35 cm. long and 4-6.5 cm. broad,
lanceolate or elliptic, obtuse, coriaceous. Inflorescence a long-pedunculate, several-
(315)
[Vor. 33
384 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
to many-flowered raceme up to 75 cm. or more long; bracts up to 6 cm. long,
linear to linear-lanceolate; flowers brown or maroon with a purple lip, medium-
sized, long-pedunculate. Sepals and petals 2.5-5.5 cm. long, undulated, linear or
spatulate, obtuse or acutish. Lip 1.3-2 cm. long and 1-1.3 cm. broad, 3-lobed,
with 3 or more inconspicuous longitudinal carinae; lateral lobes rounded, obtuse,
appressed to the column; terminal lobe orbicular-ovate, acute, recurved.
Costa Rica, Panama and possibly Venezuela.
RIQUÍ: near city of David, at sea-level, po 170. COCLE: mountains beyond
La PR alt. 400-600 m., Hunter 9 Allen
Laelia tibicinis (Batem.) L. Wms., in Darwiniana 5:77. 1941, has been reported
from Panama but we have seen no specimens nor authentic record of it from the
country.
31. BRASSAVOLA R. Br.
BRassAvOLA К. Br. іп Aiton, Hort. Kew., ed. 2, 5:216. 1813.
Epiphytic herbs with more or less thickened, 1- to 2-leaved stems. Sepals free,
equal, spreading, linear or linear-lanceolate, often long and acuminate. Petals
similar to the sepals. Lip sessile, erect, more or less enfolding the column, base
short or long and narrow, expanded into a broad lamina which is plane, rarely
concave or somewhat cucullate. Column erect, usually shorter than the claw of
the lip, more or less bialate, footless; anther operculate, incumbent, each cell with
an obscure longitudinal septum; pollinia 8, 4 in each cell of the anther, broadly
ovate and laterally compressed, ceraceous.
A genus of about 15 species. Flowers often large and showy.
a. Flowering scapes subtended by a aa „often s long as or longer than
the leaf; — usually several-flowere . B. NoposA
aa. Flowering scape from the dM not ae by a leaf; inflores-
cence usually 1- снн 2. В. ACAULIS
BRAssAVOLA Noposa (L.) Lindl. Gen. & Sp. Orch. Pl. 114. 1831; in Bot. Reg.
17: #. 1465. 1832.
Epidendrum nodosum L. Sp. Pl. 953. 1753.
Brassavola venosa Lindl. in Bot. Reg. 26: ж р. 20. 1840.
Brassavola scaposa Schltr. іп Orchis 13:77. 1919.
—
B
Erect or pendent epiphytic herbs up to about 40 cm. long, usually less; flowers
often showy. Stems or pseudobulbs 2-12 cm. long, cylindric, slender, 0.2-0.5 cm.
in diameter, usually covered with а pergameneous sheath, 1-leaved. Leaves 6-23
cm. long and 0.3—2.5 cm. broad, either terete or flattened, if flattened then usually
linear-elliptic, acute. Inflorescence borne at the end of the stem, either shorter or
longer than the subtending leaf, few- to several-flowered. Sepals 4-9 cm. long
and 0.2—0.4 cm. broad, linear, acute. Petals 4-9 cm. long, 0.05—0.2 cm. broad,
filiform to linear, acute. Lip 4.5-8 cm. long and 2.5-4 cm. broad; base unguicu-
late and cucullate, enfolding the column; anterior portion broad, cordate to
orbicular-cordate, acuminate or at least apiculate, having several prominent veins.
(316)
1946]
Fig. 140. Brassavola nodosa
(317)
385
[Vor. 33
386 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Mexico to Panama, the West Indies and northern South America. Common at
lower elevations in Central America.
BOCAS DEL TORO: Shepherd Hill, Wedel 2; vicinity of Chiriqui Lagoon, Wedel 1413,
1005, 2644. CANAL ZONE: nt Қ Fort Sherman, Allen 2844; еи Field, Powell
25; between Tumba Vieja and Salamanca, Steyermark & Allen 16753. AMA: Vicinity
of La Palma, Pittier 6680; ce. Island, Allen 2608; Taboga Island, Killip 3181;
Hacienda La Joya, Dodge et al 16903; Isla Taboga, Woodson, Allen & Seibert 1487;
Cativo, Lee 5.0. VERAGUAS: Cerro ‘eed de Cristo, alt. 1500 m., Powell 422.
2. BRASSAVOLA ACAULIS Lindl. in Paxt. Flow. Gard. 2:152, fig. 216. 1851-52.
Brassavola lineata Hook. in Bot. Mag. 79: t. 4734. 1853.
Pendent epiphytic herbs up to about 80 cm. long. Stems 2-7 cm. long,
0.2-0.5 cm. in diameter, cylindrical, unifoliate. Leaves 40-80 cm. long, 0.3-2
cm. in diameter, terete, fleshy. Inflorescence borne from the rhizome on short
peduncles (stems?), 1-, rarely 2-, flowered, much shorter than the leaves. Sepals
and petals 7-8 cm. long, 0.3—0.6 cm. broad, linear, acute. Lip 5.5-6.5 cm. long
and 2.5-3.5 cm. broad; base unguiculate, cucullate, enfolding the column; an-
terior part orbicular to orbicular-cordate, acute or acuminate, prominently veined.
Panama, reported also from Costa Rica and Guatemala.
CHIRIQUÍ: "province of Chiriquí,” alt. 1200 m., Powell 102; “Chiriquí province,”
Svibla s. n.; Boquete District, alt. 1200 m., Davidson 68 3.
32. SCAPHYGLOTTIS Роерр. & Endl.
SCAPHYGLOTTIs Poepp. & Endl. Nov. Gen. & Sp. 1:58. 1835.
Cladobium Lindl. Nat. Syst. Bot. 446. 1836 (or 1835?).
Hexadesmia Brongn. in Ann. Sci. Nat. Bot. II, 17:44. 1842.
Caespitose or repent epiphytic herbs. Stems indurated or pseudobulbose,
simply branched, or fasciculately branched (superimposed) above. Leaves 1-3
from the apex of the segments of the stem or pseudobulbs, thin and grass-like to
coriaceous or fleshy and terete. Inflorescence a fascicle or a raceme, terminal on
the growths or segments of the stem. Sepals subequal, erect. Petals similar to
the sepals but usually smaller. Lip articulated from the tip of the column-foot
or subarticulated, straight, geniculate, reflexed or recurved, clawed or clawless,
entire or 3-lobed, often emarginate. Column short, wingless or winged or auricu-
late, produced into a more or less distinct foot at the base; anther terminal,
operculate, incumbent, the loculae provided with longitudinal septae; pollinia 4
or 6, if 6 then all of equal size or two smaller than the others, ceraceous, laterally
compressed (at least 4 of each set compressed).
A difficult but interesting genus of some 40 or 50 species. The generic name
Scaphyglottis should, in all probability, be substituted by Cladobium Lindl. The
original generic description of Scaphyglottis obviously applies more to Scaphyglottis
parviflora Poepp. & Endl. than to any of the other component species. Of the
species described in the original publication most are Maxillaria, and the generic
description applies best to those species which belong to the genus Maxillaria.
(318)
19461
FLORA OF PANAMA (Orchidaceae) 387
a. Inflorescence about as long as or longer than the subtending leaves........ 1. 5. MICRANTHA
aa. Inflorescence much shorter than the subtending leaves, commonly less
than half as lon
b. Lip stg short, ни dentiform lateral lobes originating about the
middle of the 12. 8. BILINEATA
bb. Lip without бай acute, dentiform lateral lobes.
c. Column Lo ovided with stelidiar arms or teeth near the apex or
near the middle.
d. Stelidiar arms near the apex of the column
e. Lip entire or at most obscurely ти 5. 5. LAEVILABIA
е. Lip trilobate, the Hie: lobes distinct 7. S. AMETHYSTINA
"E Stelidiar arms near the middle of the column 9. S. MESOCOPIS
cc. Column е ng 5 stelidiar arms or teeth near the apex or near the
middle, etimes auriculate.
d. Lip о ах at the middle or toward the base
e. Leaves about five times longer than broad, linear-elliptic to
elliptic to Bis hie 3. S. LINDENIANA
ee. Leaves at least ten times longer than broad, linear to linear-
lanceola
f. Lip Шы lanceolate, broadest below the middle.............. 11. 5. TENELLA
ff. Lip obovate to oblong, broadest across the auriculate e
the middle . 5. MINUTIFLORA
dd. ч eri t above the middle, usually near the ap
ves relatively short and broad, mostly es than 6 cm.
long 4. 8. CUNEATA
ee. Leaves азалы long and narrow, vie a mostly more
than 6 cm. long (if Ка then only 1-2 . broad).
f. Lip 3- Lt and with the lobes about а in length.......... 6. 5. LONGICAULIS
ff. Lip simple or obscurely 3- yes if lobed then rhe lateral
lobes туф than the mid-lo
5. Lip ке ог аи S terminal lobe broad,
rounded and apiculate . 5. ACOSTAEI
gg. Lip not pandurate nor subpandurate, terminal lobe
retuse
8. S. ВЕНВИ
1. SCAPHYGLOTTIS MICRANTHA (Lindl.) Ames & Correll in Bot. Mus. Leafl. Harv.
Univ. 10:85. 1942.
Hexadesmia micrantba Lindl. in Bot. Reg. 30: Misc. 2. 1844; Reichb. f. Xenia Orch.
1:70, #. 50, figs. Ш, 6-10. 1856
Small, caespitose, epiphytic herbs up to about 20 cm. tall. Stems pseudo-
bulbose, fusiform, stipitate, simple or rarely superimposed, 1- or usually 2-leaved
at the apex. Leaves 3-12 cm. long and 0.15—0.5 cm. broad, linear, obtuse, usually
bilobed at the apex. Inflorescence a terminal, slender, few- to many-flowered
raceme about as long as or exceeding the leaves in length; flowers small. Dorsal
sepal about 1.5 mm. long and 0.75 mm. broad, oval, acute. Lateral sepals 1.5—2
mm. long and 1-1.5 mm. broad, ovate, acute. Petals 1.5-2 mm. long and about
1 mm. broad, oval or ovate, obtuse. Lip 1.2-2 mm. long and 1.5-2.5 mm. broad,
subreniform, trilobate,—in natural position the lateral lobes erect and surrounding
the column and the terminal lobe deflexed.
Guatemala, Salvador, Honduras, Nicaragua, Costa Rica and Panama.
“VERAGUAS and CHIRIQUÍ”: Warscewicz.
The smallest-flowered species of the genus.
(319)
[Vor. 33
388 ANNALS OF THE MISSOURI BOTANICAL GARDEN
2. ScaPHYGLOTTIS AcosTAEI (Schltr.) С. Schweinf. in Bot. Mus. Leafl. Нагу.
Univ. 10:27. 1941.
Hexadesmia Powellii Schltr. in Fedde Rep. Sp. Nov. Већ. 17:27. 1922, поп Scaphyglottis
Powellii Schltr.
Hexadesmia Acostaei Schltr. loc. cit. 19:295. 1923.
Small, slender, densely caespitose, epiphytic herbs up to 30 cm. tall. Pseudo-
bulbs 2.5-12 cm. long and 0.1-0.6 cm. in diameter, the upper part swollen and
fusiform, the lower part forming a slender stipe, 1- or usually 2-, leaved. Leaves
5-14 cm. long and 0.1-0.3 cm. broad, linear, obtuse, the apex usually bilobed.
Inflorescence a short, terminal, few-flowered raceme, much exceeded by the leaves.
Flowers small, usually white with a dark lip. Sepals 3-4 mm. long and 1.4-1.8
mm. broad, lanceolate to ovate-lanceolate, acuminate. Petals 3-4 mm. long and
1.4-2 mm. broad, lanceolate or lanceolate-oblong, acute. Lip 3-4 mm. long and
2.5-4 mm. broad, oblong or oblong-subquadrate, pandurate, acute or retuse and
apiculate.
Costa Rica and Panama.
CHIRIQUÍ: Volcán de Chiriquí, alt. 2100 m., Davidson 867; slope of Cerro de la
Horqueta, alt. 1200-1700 m., Maxon 5300; alt. 1500 m., Powell 225.
3. SCAPHYGLOTTIS LINDENIANA (Rich. & Gal.) L. Wms. in Ann. Missouri Bot.
Gard. 28:423. 1941.
Hexadesmia fasciculata Brongn. in Ann. Sci. Nat. Bot. II, 17:45. 1842, non Scaphyglottis
Hexadesmia и Lindridena Rich. & Gal. in Ann. Sci. Nat. Bot. ІП, 3:23. 1845.
Hexadesmia pacbybulbon Schltr. in Fedde Rep. Sp. Nov. Beih. 17:26. 1922.
Small to large and coarse epiphytic herbs up to about 50 cm. tall. Stems
usually stipitate below, swollen into an ellipsoidal or fusiform pseudobulb above,
bifoliate. Leaves 5-25 cm. long and 1.5-5 cm. broad, linear-elliptic to elliptic to
lanceolate to oblanceolate, acute, coriaceous. Inflorescence a fascicle of flowers
borne at the apex of the pseudobulb. Dorsal sepal 7-11 mm. long and 1,5-2 mm.
broad, elliptic, acute. Lateral sepals 7-11 mm. long and 2-3 mm. broad, lanceo-
late, arcuate, acute. Petals 6-10 mm. long and 1.5-3 mm. broad, elliptic-
lanceolate, acute. Lip 6.5-10 mm. long and 2.5—4.5 mm. broad, oblanceolate to
narrowly obovate, obscurely 3-lobed, obtuse, apiculate.
Mexico, Guatemala, Costa Rica and Panama.
uf: "Province of Chiriquí,” alt. 1500 m., Powell 220, 3149. состЕ: hills
o of El Valle de Antón, alt. 800-1000 m., Allen 2201; mountains beyond La Pintada,
alt. 400—600 m., Hunter & Allen 601
Scaphyglottis falcata C. Schweinf. is possibly only a somewhat larger-flowered
form of this variable species.
4. ScAPHYGLOTTIS CUNEATA Schltr. іп Вей. Bot. Centralbl. 36, Abt. 2:398. 1918.
Scaphyglottis prolifera (R. Br.) Cogn. and other authors as to plants, not according to
name-bringing synonym
Tetragamestus gracilis Schltr. loc. cit. 400.
(320)
19461
FLORA OF PANAMA (Orchidaceae) 389
Scaphyglottis Wercklei Schltr. in "T Rep. Sp. Nov. Beih. 19:28. 1923.
Scaphyglottis gracilis Schltr. loc.
ни IM Lindl. ex C. Schweinf. in Bot. Mus. Leafl. Harv. Univ. 11:181. 1944,
и caespitose or repent, epiphytic herbs up to about 25 ст. long. Stems
pseudobulbose, usually superimposed; pseudobulbs slender and cylindric to nar-
rowly fusiform, usually bifoliate at the top of each segment of the growth.
Leaves 1-6 cm. long and 0.1-0.7 cm. broad, linear or ligulate to elliptic-oblong,
obtuse, usually retuse. Inflorescence a single flower or a few-flowered fascicle
borne at the apex of the segments of the stem. Dorsal бергі 3.5—4.5 mm. long and
1.2-1.5 mm. broad, elliptic to elliptic-oval, acute, cucullate. Lateral sepals 4-5
mm. long and 1.5—2.5 mm. broad, linear-oblong, oblique, acute, forming an іп-
conspicuous mentum at the base. Petals 3.5-5 mm. long and 0.5-0.6 mm. broad,
linear to linear-ligulate, obtuse or acute, slightly oblique. Lip 4-6 mm. long and
2.5—4 mm. broad toward the apex, unguiculate, cuneate-flabellate, the apical part
semi-orbicular, obtuse, entire or trilobate; disc with a small flabellate or bipartite
callus toward the apex of the claw or rarely ecallous. Column without stelidiar
arms.
Guatemala, British Honduras, Honduras, Nicaragua, Costa Rica and Panama,
the West Indies and in South America to Brazil and Bolivia.
ZONE: r Río Medio, Miller 1747; San Juan Range, Powell 349; hills north
of Frijoles, SL 20057. cocLÉ: vicinity of El Valle de Antón, alt. 600-1000 m.,
5. SCAPHYGLOTTIS LAEVILABIA Ames in Proc. Biol. Soc. Wash. 34:154. 1921, as
S. laevilabium.
Similar to Scaphyglottis cuneata except the disc of the lip always ecallous and
the column with two small stelidiar arms about opposite the stigmatic cavity.
Panama.
PANAMÁ: Juan Díaz, alt. 50 m., Killip 3113; foothills east of Panamá City, Powell 211.
6. SCAPHYGLOTTIS LONGICAULIS S. Wats. in Proc. Am. Acad. 23:286. 1888.
Scaphyglottis unguiculata Schltr. in Fedde Rep. Sp. Nov. 12:206. 1913.
Slender, caespitose, epiphytic herbs up to about 30 cm. tall. Stems slender, the
pseudobulbs cylindric, simple or rarely superimposed, bearing 1 or 2 leaves at the
summit of the segments. Leaves 4-16 cm. long and 0.1—0.3 cm. broad, linear.
Inflorescence one-flowered, or a few-flowered fascicle at the apex of the stem or of
segments of the stem. Dorsal sepal 5-7 mm. long and about 1.5 mm. broad,
linear-oblong to linear-oblanceolate, acute, somewhat cucullate. Lateral sepals
5.5-7 mm. long and 1.5-2 mm. broad, linear-oblong, acute, forming a distinct
mentum at the base. Petals 5-7 mm. long and about 0.7-1 mm. broad, linear-
oblanceolate, obtuse or acute. Lip 5-7 mm. long and 3-4 mm. broad near the
apex, unguiculate, cuneate-flabellate, trilobate at the apex; disc ecallous. Column
sometimes with lateral teeth at the apex.
Guatemala, Honduras, Costa Rica, Panama and Colombia.
(321)
[Vor. 33
390 ANNALS OF THE MISSOURI BOTANICAL GARDEN
CANAL ZONE: Barro Colorado Island, Kenoyer 251; hills east of Panama City, Powell
105, 3392; hills north of Frijoles, Standley 27664; Rio Indio, Steyermark & Allen 17442.
OLÓN: Loma de la Gloria, Pittier 7000. DARIEN: vicinity of El Real, alt. 15 m., Allen
944.
7. SCAPHYGLOTTIS AMETHYSTINA (Reichb. f.) Schltr. іп Beih. Bot. Centralbl. 36,
Abt. 2:456. 1918.
Ponera amethystina Reichb. f. in Saunders Ref. Bot. 2: /. 03. 1869.
Scaphyglottis brachiata Schltr. in Fedde Rep. Sp. Nov. 9:432. 1911.
Caespitose epiphytic herbs up to about 25 cm. tall. Stems simple or superim-
posed, segments cylindric to fusiform, often distinctly stipitate, bearing 1 or
usually 2 leaves at the apex of each segment. Leaves 3-12 cm. long and 0.4-1 cm.
broad, linear, linear-lanceolate to elliptic, obtuse, the apex retuse and unequally
bilobed. Inflorescence a few-flowered fascicle borne at the apex of the segments.
Dorsal вера! 4.5-6 mm. long and about 1.5 mm. broad, elliptic to oblanceolate,
acute, cucullate. Lateral sepals 5-7.5 mm. long and 1.5-2 mm. broad, linear-
oblong to lanceolate, acute, forming a short mentum at the base. Petals 4.5—5
mm. long and 0.8-1.2 mm. broad, linear-oblong and unguiculate to oblanceolate,
acute. Lip 6-8 mm. long and 3-5 mm. broad, cuneate, obovate, 3-lobed, the
lateral lobes large, rounded, situated above the middle of the lip; mid-lobe sub-
quadrate, obtuse, rarely emarginate, exceeding the lateral lobes. Column with 2
stelidiar arms near the apex.
Costa Rica and Panama.
CANAL ZONE: Chagres River, sea-level, Powell 358. снікюші: vicinity of San Felix,
alt. 0-120 m., Pittier 5103.
8. ScAPHYGLOTTIS Венкп (Reichb. f.) Benth. & Hook. ех Нет]. іп Godm. &
Salvin, Biol. Centr.-Am. Bot. 3:219. 1883; C. Schweinf. in Bot. Mus. Leafl.
Harv. Univ. 7:186. 1939.
Ponera Вертй mie f. in Bonplandia 3:220. 1855.
Ponera albida Reichb. f. in Beitr. Orch. Centr.-Am. 103. МеН
Scaphyglottis Ро йа Schltr. іп Fedde Rep. Sp. Nov. 3:47.
Scaphyglottis albida Schltr. in Већ. Bot. Centralbl. 36, Abt. 2: yes 1918.
Caespitose, epiphytic or pendent herbs up to about 45 cm. long. Stems simple
or superimposed; segments cylindric to narrowly elliptic or narrowly fusiform,
l-leaved or usually 2-leaved at the apex, at least the lower segment stipitate
Leaves 5-26 cm. long and 0.15—0.45 cm. broad, linear, obtuse, obscurely bilobed
at the apex. Inflorescence a single flower or а few-flowered fascicle or one or
more much abbreviated few-flowered racemes borne at the apex of the segments
of the stem. Dorsal sepal 3-4 mm. long and 1-1,5 mm. broad, elliptic-oblong to
elliptic-oval, obtuse, cucullate. Lateral sepals 3.5—4.5 mm. long and 1,2-2 mm.
broad, linear-oblong to oblong, acute. Petals 3-4 mm. long and 0.5-1 mm.
broad, linear to linear-lanceolate, obtuse, more or less unguiculate. Lip 3.5-5 mm.
long and 2-3.5 mm. broad, narrowly obovate, unguiculate, entire to obscurely
3-lobed, obtuse and rarely shallowly retuse. Column with a narrow wing.
Guatemala, British Honduras, Honduras, Nicaragua, Costa Rica, Panama and
Colombia.
(322)
19461
FLORA OF PANAMA (Orchidaceae) 391
CANAL ZONE: Chagres, Behr; near Gatin, Hayes; hills east of Panama City, Juan
Díaz Range, San Juan Range, La Pita, McComber Hill, Frijoles, Juan Diaz, Powell 126,
359, 363, 367, 360, 373, 374, 378, 3506; Barro Colorado Island, Woodworth & Vestal
592. CHIRIQUÍ: Lino Hill, Powell 385. PANAMA: Río Tecümen, Hunter & Allen 229.
VERAGUAS: Bahia Honda, Taylor 1507.
It is possible that the historical type of Scaphyglottis Bebrii is not the species
commonly interpreted as such. We have not seen the original specimen collected
by Behr. А record from the Reichenbach herbarium (not from the type) does
not match too well.
9. ScaPHYGLOTTIS MEsOCOPIS (Endr. & Reichb. f.) Benth. & Hook. ex Нет].
in Biol. Centr.-Am. Bot. 3:220. 1883
Ponera mesocopis Endres & Reichb. f. Xenia Orch. 2:222, Ё. 200. 1874.
Scapbyglottis Powellii Schltr. in Fedde Rep. Sp. Nov. Beih. 17:28. 1922.
Caespitose, erect or repent, epiphytic herbs up to about 50 cm. long. Stems
pseudobulbose, cylindric or usually fusiform, commonly stipitate; segments super-
imposed, 1- to 2-leaved at the apex, the lower segment often bearing a fascicle of
stems at its apex. Leaves 3-17 cm. long and 0.2-0.7 cm. broad, linear, obtuse,
usually bilobate at the apex. Inflorescence a few-flowered fascicle or а much-
abbreviated raceme borne at the apex of the segments of the stem. Dorsal вера!
5.5-7 mm. long and 1.8-2.5 mm. broad, elliptic to elliptic-oblong, acute. Lateral
sepals 6-8 mm. long and 1.8-2.5 mm. broad, linear-oblong to lanceolate-oblong,
acute or obtuse, slightly arcuate, forming a short mentum at the base. Petals 5-7
mm. long and 1.4-2 mm. broad, linear-oblong to elliptic-lanceolate, acute or
obtuse, short-unguiculate. Lip 7-10 mm. long and 3.5—4.5 mm. broad, oblong-
subpandurate, usually broadest near the apex, cuneate to the base and short-
unguiculate, obtuse and emarginate, obscurely denticulate; disc with one or more
the median lines raised and papillose. Column with two stelidiar arms originat-
ing at about the middle.
Costa Rica and Panama.
CANAL ZONE: hills east of Panamá City, Powell 134. cHrriguf: vicinity of Boquete,
near Río Caldera, alt. кр m., Allen 2282; n alt. 1200 m., Davidson 1283;
"Province of Chiriquí," alt. 000 m., Powell 134. CLÉ: vicinity of La Mesa north of
El Valle de Antón, alt. me m., Allen 2316; region odi of El Valle de ye alt. 800
m., Allen 2926.
—
ЕТ
The Powell record for the Canal Zone is doubtless in error, for the same num-
ber is used for the specimen from Chiriqui.
10. SCAPHYGLOTTIS MINUTIFLORA Ames & Correll in Bot. Mus. Leafl. Harv. Univ.
10:83, 20. 1942.
Caespitose epiphytic herbs up to about 75 cm. tall. Stems slender, cylindric
to slightly fusiform, superimposed, 1- to 2-leaved from the apex of the segments.
Leaves 5-18 cm. long and 0.3-1 cm. broad, linear, acute or obtuse, retuse and
unequally bilobed at the apex. Inflorescence a few-flowered fascicle, borne at the
apex of segments of the stem. Dorsal sepal about 2.4 mm. long and 1.3 mm.
(323)
[Vor. 33
392 ANNALS OF THE MISSOURI BOTANICAL GARDEN
==
з,
Lope Ca PRESS =.
==—=——
==
GOR С
SCAPHYGLOT TIS NX red
minutiflora Ames $ Соё “ .
Fig. 141. Scapbyglottis minutiflora
(324)
1946]
FLORA OF PANAMA (Orchidaceae) 393
broad, oval, acute. Lateral sepals about 2.5 mm. long and 1.4 mm. broad, elliptic-
oblong, acute, forming an inconspicuous mentum at the base. Petals about 2.2
mm. long and 0.6 mm. broad, narrowly elliptic-oblong, acute or obtuse. Lip
2.5-4 mm. long and 1,5-2.5 mm. broad, obovate to oblong, obtuse or obscurely
retuse, provided with small, semi-orbicular, auriculate lobes near the middle.
Guatemala, British Honduras and Panama.
COCLÉ: hills north of El Valle de Antón, alt. 800-1000 m., Allen 2258.
It is possible that the Panamanian material is different from the Guatemalan
type.
11. SCAPHYGLOTTIS TENELLA L. Wms. in Ann. Missouri Bot. Gard. 28:423,
t. 24. 1941
Caespitose, erect or spreading, epiphytic herbs up to about 50 cm. long. Stems
slender, cylindric or becoming swollen and fusiform above, simple, bifoliate at the
apex. Leaves 20-23 cm. long and 0.7-1.6 cm. broad, linear to linear-lanceolate,
acute, the apex minutely bilobed. Inflorescence a 1- to few-flowered fascicle at
the apex of the stem. Dorsal sepal 8-9 mm. long and 3-4 mm. broad, lanceolate,
acute. Lateral sepals 8-9 mm. long and 3-4 mm. broad, lanceolate, acute. Petals
8-9 mm. long and 3-3.5 mm. broad, oblong-lanceolate, acute or obtuse. Lip 8-9
mm. long and 4—5 mm. broad, rhombic-lanceolate but somewhat constricted
above the middle, fleshy.
Panama.
cocLÉ: hills north of El Valle de Antón, alt. 800-1000 m., Allen 2276.
12. SCAPHYGLOTTIS BILINEATA (Reichb. f.) Schltr. in Већ. Bot. Centralbl. 36,
Abt. 2:456. 1
Ponera bilineata Reichb. f. Beitr. Orch. Centr.-Am. 88. 1866.
Erect or spreading, caespitose, epiphytic herbs up to about 30 cm. long. Stems
slender, simple or superimposed, cylindric, 1- to 2-foliate at the apex of the seg-
ments. Leaves 6—9 cm. long and 0.7—1.1 cm. broad, ligulate to linear-lanceolate,
obtuse. Inflorescence a 1- to few-flowered fascicle borne at the apex of the seg-
ments. Dorsal sepal about 7 mm. long and 2 mm. broad, elliptic or oblanceolate,
obtuse. Lateral sepals about 8 mm. long and 2.5 mm. broad, lanceolate-oblong,
slightly arcuate, acute. Petals about 7 mm. long and 0.8 mm. broad, linear, ob-
tuse. Lip about 9 mm. long and 3.5 mm. broad, cuneate-unguiculate to the base,
3-lobed; the lateral lobes originating above the middle, dentiform; the mid-lobe
suborbicular-flabellate, slightly retuse; disc fleshy, especially the basal portion,
with the median line callus-thickened toward the apex. Column auriculate at
the apex.
Costa Rica and Panama.
ZONE: Quebrada Lopez, alt. 30 m., Allen 2139. cocté: hills north of El
Valle "ig peu, alt. 800-1000 m., Allen 2263.
May be expected to be more variable than the description indicates.
(325)
[Vor. 33
394 ANNALS OF THE MISSOURI BOTANICAL GARDEN
SCAPHYGLOT TIS
tenella 26.
Fig. 142. Scaphyglottis tenella
(326)
19461
FLORA OF PANAMA (Orchidaceae) 395
OBSCURE SPECIES
SCAPHYGLOTTIS DOLICHOPHYLLA Schltr. in Fedde Rep. Sp. Nov. Beih. 17:28. 1922.
Panama.
This species is based on Powell 126 of which I have seen two specimens. One
of these is Scaphyglottis Bebrii, while the other is very similar but recalls in some
respects Schlechter’s analysis of S. dolichophylla.
33. PLATYGLOTTIS L. Wms.
PrATvcLOoTTIs L. Wms. in Ann. Missouri Bot. Gard. 29:345, 7. 34. 1942.
Caespitose, epiphytic herbs from a very short rhizome. Stems slender, non-
pseudobulbose, indurated, simple. Leaves alternate, distichous, coriaceous, plane,
deciduous; leaf-sheaths tightly enfolding the stem. Inflorescence terminal (or
subterminal?), racemose. Sepals subequal; dorsal sepal free; lateral sepals adnate
to the short column-foot at their bases and with it forming an inconspicuous
mentum. Petals similar to the sepals or broader. Lip articulated to the apex of
the column-foot; lamina broadly ligulate, entire. Column short, clavellate, wing-
less, produced into a short foot at the base; anther terminal, operculate, incumbent,
6-loculate; pollinia 6, equal, 4 basal and laterally compressed in one series, 2
terminal in a second laterally compressed series, ceraceous.
A single species known only from the vicinity of El Valle de Antón, Coclé
Province, Panama.
1. PLATYGLOTTIS CORIACEA L. Wms. in Ann. Missouri Bot. Gard. 29:347, f. 34.
1942.
Epiphytic, caespitose herbs up to about 4.5 dm. tall Stems 5-8 mm. in
diameter, slender, leafy, becoming naked with age. Leaves 2.5—6 cm. long, 1-1.5
cm. broad, narrowly ligulate, obtuse, obscurely and unequally bilobed at the apex,
coriaceous, distichous. Inflorescence a short, few-flowered raceme, terminal or sub-
terminal; bracts 7-20 mm. long, elliptic-lanceolate, acute, cucullate; flowers rather
large for the tribe, sepals and petals lavender, the lip lavender with greenish mar-
gins. Dorsal вера! about 10 mm. long and 3.5 mm. broad, lanceolate, acute or
acuminate, 5-nerved. Lateral sepals about 10 mm. long and 3 mm. broad, lance-
olate, acute or acuminate, slightly oblique, 5-nerved. Petals about 10 mm. long
and 3.5 mm. broad, elliptic-oblanceolate, somewhat oblique, obscurely constricted
near the apex, 5- to 7-nerved. Lip about 10 mm. long and 5 mm. broad toward
the apex, broadly ligulate, subpandurate, entire, apex rounded, the margins thin
and plicated laterally; disc fleshy, with a pair of inconspicuous, subumbonate
calluses at the base.
Panama.
cocLÉ: north of El Valle de Antón, alt. about 1000 m., Allen 2036.
(327)
[Vor. 33
396 ANNALS OF THE MISSOURI BOTANICAL GARDEN
yc 4 E и
~ 20/4 асва Ио, WS.
i. m ИТ,
AN
Бір. 143. Scaphyglottis coriacea
(328)
19461
FLORA OF PANAMA (Orchidaceae) 397
34. ISOCHILUS R. Br.
IsocuiLus К. Br. in Ait. Hort. Kew. ed. 2, 5:209. 1813; Correll in Bot. Mus. |
ГеаЯ. Harv. Univ. 10:1. 1941.
Caespitose, epiphytic herbs with slender
stems and distichous leaves. Inflorescence a
terminal, often unilateral raceme. Sepals sub-
equal, free or connate into a short tube at the
base, sometimes more or less subsaccate at the
base. Petals about as long as the sepals. Lip
subequal to the petals, simple, adnate to the
base of the column or column-foot, often semi-
terete, wingless; column-foot very short and
inconspicuous; pollinia 4, ceraceous, elongated
and laterally compressed.
Closely allied to Ponera and Jacquiniella.
According to Correll, the genus consists of two
species, both variable.
1. IsocHiLUS Major Cham. & Schlecht. in
Linnaea 6:60. 1831; Correll in Bot. Mus.
Leafl. Harv. Univ. 10:9. 1941.
Isochilus chiriquensis Schltr. in Fedde Rep. Sp. Nov.
1 29.
Caespitose, erect or spreading, slender, ері-
phytic herbs. Stems simple, slender, leafy.
Leaves 2-10 cm. long and 0.2—0.8 cm. broad,
linear to linear-elliptic, obtuse, emarginate and
unequally bilobed at the apex. Inflorescence
secund, usually several- to many-flowered;
flowers subtended by conspicuous chartaceous
bracts. Dorsal вера! 7-11 mm. long and 3—4
mm. broad, elliptic-oblong, acute or obtuse.
Lateral sepals 8-12 mm. long and 3—4.5 mm.
broad, oblong-lanceolate, acute, keeled toward
the base, and the keel decurrent onto the ovary.
Petals 7-11 mm. long and 2-3 mm. broad,
oblanceolate, acute or obtuse. Lip 8—12 mm.
long and 1.5—3 mm. broad, elliptic-linear to
Fig. 144. Isochilus major
(329)
[Vor. 33
398 ANNALS OF THE MISSOURI BOTANICAL GARDEN
linear-oblong to oblanceolate, acute or obtuse,
M Ў entire or obscurely lobed, geniculate and fleshy
De at the base.
cH (>> Mexico, Guatemala, Honduras, Costa Rica,
OR Уз Panama and Jamaica.
снікюоі: Rio Quebrada, alt. 1700 m., Killip
3532; “Province of Chiriqui,” alt. 1200 m., Powell
08, 3380; Bajo Mona, alt. 1500-2000 m., Woodson,
Allen t$ Seibert 1003; vicinity of Bajo Mona, alt.
1500 m., Woodson 9 Schery 518.
35. BOTHRIOCHILUS! Lemaire
BoTHRIOCHILUs Lemaire, Illustr. Hort. 3:Misc.
p. 31. 1856; L. Wms. in Bot. Mus. Leafl.
Harv. Univ. 8:145. 1940.
Epiphytic herbs with the stems reduced to
ovoid or lageniform pseudobulbs. Pseudobulbs
several-leaved at their apices, the petioles of
the leaves simulating stems. Inflorescence lat-
. eral from the base of the pseudobulbs. Dorsal
sepal free. Lateral sepals adnate to the column-
foot and with it forming a mentum. Petals
similar to the dorsal sepal but usually shorter.
Lip simple or obscurely lobed, about as long as
the petals, shortly and sharply declined at the
base or even saccate, if saccate the sac some-
times didymous. Column erect, long and slen-
der, wingless or nearly so; column-foot about
as long as the column or longer; pollinia 8,
ceraceous. Ovary wingless.
A single species is known from Panama.
1. BoTHRIOCHILUS MACROsTACHYUs (Lindl.)
L. Wms. in Bot. Mus. Leafl. Harv. Univ.
8:148. 1940
Coelia ӨМетогіесіуе pro in Benth. Pl. Hartw.
92. 3 Hook. Bot. Mag. 79:4. 4712.
1853; Reichb. f. Beitr. Orch. Centr.-Am. 41.
1866.
Fig. 1To this genus belong all of the "NON previously re-
Botbriocbilus macrostacbyus ferred to Coelia, except the type specie
(330)
19461
FLORA OF PANAMA (Orchidaceae) 399
Coelia macrostachya Lindl. var. genuina Reichb. f. loc. с
Coelia macrostachya Lindl. var. integrilabia Reichb. f. loc. cit.
Epiphytic herbs up to about 90 cm. tall. Pseudobulbs up to about 8 cm. long,
ovoid to semi-globose, often somewhat compressed, surrounded with scarious sheaths
at the base, bearing 3-6 leaves at the summit. Leaves 30-100 cm. long and
0.8—2.5 cm. broad, linear to elliptic-ligulate, acute or acuminate, plicate, sheathing
at the base. Inflorescence a densely flowered, more or less elongated raceme, lateral,
up to about 50 cm. long, the scape somewhat flexuous, covered with chartaceous
bracts; the rachis provided with conspicuous linear to lanceolate, acute or acumi-
nate bracts up to about 7 cm. long. Dorsal вера! 7.5-11 mm. long and 3-5 mm.
broad, elliptic-ovate to ovate, acute, rugose dorsally. Lateral sepals 10-14 mm.
long and 4-5 mm. broad, oblong-lanceolate, acute, free but forming a short ob-
tuse mentum, rugose dorsally. Petals 7-11 mm. long and 3-5 mm. broad, elliptic-
ovate to rhombic-obovate, acute. Lip 11-14 mm. long and 4-6 mm. broad,
oblong-pandurate, angled or dentate-lobed above the middle, terminal part of the
lip reflexed, lanceolate, base of lip forming a shallow didymous spur.
Mexico, Guatemala, Honduras and Panama.
CHIRIQUÍ: “Chiriquí Cordilleren,” alt. 1800 m., Warscewicz.
The Panamanian record is a specimen cited by Reichenbach.
36. POLYSTACHYA Hook.
PorysracHvA Hook. Exot. Fl. 2: . 103. 1825; Kranzlin in Fedde Rep. Sp. Nov.
Beih. 39:1—136. 1926.
Epiphytic or rarely terrestrial, caespitose, pseudobulbose (Panamanian species)
herbs. Inflorescence a raceme or panicle. Dorsal sepal free. Lateral sepals larger
than the dorsal, forming a mentum at their bases. Petals much smaller than the
sepals, usually linear. Lip non-resupinate, nearly entire or usually 3-lobed; disc
of the lip usually with a conspicuous callus and often covered with fragile, artic-
ulated or farinaceous cells. Column short, with a prominent foot; pollinia 4, or
4 joined into 2.
Kranzlin, in his monograph, lists 188 species of Polystachya, mostly African.
1. Р. CEREA
. 2. P. MASAYENSIS
a. Ovary and rachis of the inflorescence glabrous
aa. Ovary and rachis of the inflorescence pubescent
1. PoLYsTACHYA CEREA Lindl. in Bot. Reg. 26: Misc. p. 86. 1840.
Caespitose, epiphytic herbs up to about 60 cm. tall, commonly 10-15 cm. tall.
Stems very short, pseudobulbose; pseudobulbs ovoid to fusiform or cylindric,
usually enclosed in bracts or the leaf-sheaths, bearing 2-5 leaves. Leaves 3-26
(331)
[Vor. 33
400 ANNALS OF THE MISSOURI BOTANICAL GARDEN
cm. long and 0.4-3 cm. broad, variable, linear to oblong-elliptic to oblanceolate,
acute or obtuse, the sheaths enclosing the peduncle at the base. Inflorescence
terminal, the peduncle covered with scarious sheaths, bearing a single terminal
raceme or 1 or more short lateral racemes from the nodes of the peduncle. Dorsal
sepal 3-3.5 mm. long and 1.6-2 mm. broad, lanceolate, ovate to oblong-ovate,
acute. Lateral sepals 3-4.5 mm. long and 1,5-3 mm. broad, lanceolate-ovate to
broadly ovate, acute or acuminate. Petals 2-2.5 mm. long and 0.5-1 mm. broad,
obtuse or acute, slightly arcuate. Lip 2.5-3.5 mm. long and 2-3 mm. broad, ob-
long or oblong-ovate, 3-lobed; lateral lobes small, usually rounded, erect; mid-lobe
large, truncate, apiculate or usually retuse; disc provided with an umbonate or
mammillate callus at the base and with the terminal part covered with evanescent,
farinaceous pubescence. Ovary glabrous.
Mexico, Central America, the West Indies, tropical South America.
CANAL ZONE: hills east of Panamá City, NM 122, 3426; Barro Colorado Island,
Shattuck 226. снікюоі: opposite чир. alt. 1200 m., Powe 343. COCLE: vicinity of
El Valle, alt. 600-1000 m., Allen 781, 1171. PANAMA: FHaicende La Joya, alt. 50-300 m.,
Dodge et al 16000; Sabanas near Chepo o, alt. 30 m., Hunter 9 Allen 28. PROVINCE NOT
GIVEN: "hills," Powell 122, 3439; between Fort ыда and Corozal, Standley 20100.
2. POLYSTACHYA MASAYENSIS Reichb. f. in Bonplandia 3:217. 1855.
Similar to Polystachya cerea Lindl. but usually somewhat smaller in size, vege-
tatively, and with the rachis of the inflorescence and the ovary downy-pubescent.
Mexico, Nicaragua, Costa Rica and Panama.
CANAL ZONE: Chagres, Fendler 334. COCLÉ: hills north of El Valle de Antón, =
1000 m., Allen 2319; Río Antón, alt. 500 m., Hunter & Allen 380. cortów: Cat
Porto Bello trail, alt. sea-level, Powell 366.
Perhaps best treated as a mere variety of P. cerea Lindl.
37. GALEANDRA Lindl.
GALEANDRA Lindl. in Lindl. & Bauer, Illustr. Orch. Pl., Gen. 7. 8. 1832.
Terrestrial or epiphytic herbs. Stems leafy, becoming thickened and pseudo-
bulbose. Leaves distichous, plicate. Sepals free, equal, spreading. Petals similar
to the sepals, sometimes broader. Lip entire or bilobed, adnate to the base of the
column, with a conspicuous spur at the base; lateral lobes usually erect and en-
folded over the column. Column erect, short; anther terminal, operculate, in-
cumbent, imperfectly 2-celled; pollinia 4, ceraceous, often in pairs.
There are 3 species of Galeandra in Mexico and Central America, 2 of which
occur in Panama—About 20 species in South America.
a. Plants epiphytic; lip large, with ызды 4-5 cm. long 1. С. BAUERI
Plants terrestrial; Пр about 2 cm. lon ... 2. С. JUNCEA
1. GALEANDRA BAUERI Lindl. in Lindl. & Bauer, Illustr. Orch. РГ, Gen. £. 8.
1832; Batem. Orch. Мех. & Guat. 7. 10. 1840; Lindl. in Bot. Reg. 26: 7. 40.
1840; L. Wms. in Ann. Missouri Bot. Gard. 26:284. 1939.
Galeandra Batemanii Rolfe in Gard. Chron. III, 12:431. 1892.
(332)
19461
FLORA OF PANAMA (Orchidaceae) 401
Fig. 146. Polystachya masayensis
(333)
[Vor. 33
402 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Fig. 147. Galeandra Baueri
(334)
1946]
FLORA OF PANAMA (Orchidaceae) 403
Caespitose, epiphytic herbs up to about 50 cm. tall. Stems pseudobulbose,
fusiform to ovoid, covered with the leaf-sheaths or becoming naked, bearing up to
about 8 leaves. Leaves 8-26 cm. long and 0.8-2 cm. broad, linear-elliptic to
elliptic, acute or acuminate, plicate. Inflorescence a terminal few-flowered raceme;
flowers relatively large and pretty. Dorsal вера! 15-19 mm. long and 3-5 mm.
broad, oblanceolate, acute. Lateral sepals 16-20 mm. long and 4-6 mm. broad,
lanceolate or elliptic-lanceolate, acute, slightly arcuate. Petals 16-20 mm. long
and 4-6 mm. broad, elliptic to elliptic-oblanceolate, acute. Lip, including the
spur, 40-50 mm. long and 30-40 mm. broad, infundibuliform, spur just about
equalling the lamina in length; lamina suborbicular-rhombic, emarginate.
Mexico, British Honduras, Guatemala, Honduras, Panama and French Guiana.
A variety in Peru.
s DEL TORO: in swamp near Almirante, Nash 1962; Shepherd Island, vicinity of
Chiriqui а Wedel 2712
2. GALEANDRA JUNCEA Lindl. беге. Orch. sub. 2. 37. 1840; Cogn. in Mart. FI.
Bras. 34:306. 1895
Strict terrestrial herbs up to about 50 cm. tall. Stems pseudobulbose, the
pseudobulbs 1.5—2 cm. long, borne on the surface or just under the surface of the
ground, simple, invested by the leaf-sheaths. Leaves up to about 20 cm. long and
0.5 cm. broad, ensiform, conduplicate. Inflorescence a simple (or branched?)
few-flowered raceme. Sepals about 13 mm. long and 3.5-4 mm. broad, lanceolate,
acute. Petals about 12 mm. long and 3.5 mm. broad, oblanceolate, acute. Lip
17-20 mm. long and about 16 mm. broad, obscurely trilobate, spurred; lamina
when expanded broadly ovate in outline, with 2 small lamellate, pubescent calluses
near the middle, and toward the apex a dense patch of pilose pubescence arranged
in three lines; spur short, about 5 mm. long, directed outward and backward.
Panama, Venezuela, the Guianas and Brazil.
PANAMA: Isla Taboga, alt. 0-350 m., Allen 1268; Taboga Island, Cheeseman 483.
The description is drawn from the Panamanian material cited.
38. EPIDANTHUS L. Wms.
Еррлмтноѕ L. Wms. in Bot. Mus. Leafl. Harv. Univ. 8:148. 1940.
Small, simple or branched, epiphytic herbs with slender, leafy, repent or caespi-
tose stems lacking pseudobulbs. Leaves distichous, jointed at the base, plane or
terete, linear or subfiliform; leaf-sheaths persistent on the stem. Inflorescence a
terminal, distichous, fractiflex raceme. Sepals similar, free, reflexed or spreading.
Petals narrower than the sepals, with a callus at the base. Lip simple or 3-lobed,
adnate to the base of the column. Column short, terete, adnate to the lip or
callus of the lip for its entire length; anther terminal, incumbent; pollinia 2,
ceraceous.
A small Mexican and Central American genus of three species, only one of
(335)
[Vor. 33, 1946]
404 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN
which is known from Panama. It has been confused with Epidendrum, with
which it has little affinity.
1. EPIDANTHUS PARANTHICUS (Reichb. f.) L. Wms. in Bot. Mus. Leafl. Harv.
Univ. 8:150.
Epidendrum paranthicum Reichb. f. in Bot. Zeit. 10:732.
Epidendrum Sancti Ramoni Kranzl. in Vierteljahrschr. Noti Zürich 74:137. 1929,
Small, caespitose, epiphytic herbs up to 30 cm. tall. Stems slender, slightly
fractiflex. Leaves 0.8—2.5 cm. long and 0.05—0.2 cm. broad, linear, semi-terete,
obtuse, deciduous. Inflorescence a terminal, fractiflex, few- to several-flowered
raceme; flowers small, white or yellow. Sepals similar, 2-3 mm. long and 1-1.5
mm. broad, lanceolate to ovate-lanceolate, acute, the laterals often slightly
cucullate. Petals 2-2.5 cm. long and about 0.5 mm. broad, linear-lanceolate,
acute. Lip 2.2-3 mm. long and 2-2.5 mm. broad, unguiculate; the claw adnate
to the column; lamina cordate-orbicular to reniform, long-apiculate.
Mexico, Guatemala, Honduras, Costa Rica and Panama.
CHIRIQUÍ: Вајо Chorro, Boquete District, alt. 1800 m., Davidson 121.
(То be continued in Part ІП, Fascicle 4)
(336)
GENERAL INDEX TO VOLUME XXXIII
New scientific names of plants and the final members of new combinations are printed
in bold face type; synonyms and page numbers having reference to figures and plates, in
italics; and previously published names and all other matter, in ordinary type.!
A
Aceto-carmine stain in studies of yeast
cultures, 253,
Aceto-orcein stain in studies of yeast cul-
tures, 253, 260, 282, 288
Acostaea, 84
Acupula, Mexico, maize grown near, 239
American Carboniferous floras, Contribu-
tions to our knowledge of. VIII. Another
Medullosa from Iowa, 141; ome
petrified seeds from Iowa, 291
aize in Mexico—A
ican Carboniferous floras. VIII. Another
Medull from Iowa, d Lee
Lenz i
knowledge of American Carboniferous
Some petrified seeds from
Archeological investigations of maize, 161,
168
Autlan, Mexico, varieties of maize common
at, 149, 158
B
Blastodendrion, 264
Bothriochilus, 398
Brassavola, 384
Brazilin stain in studies of yeast cultures,
253, 256, 278, 28
C
"Cacahuzintle" variety of maize, 149, 166
Carboniferous floras, American, Contribu-
i knowledge of IL An-
other Medullosa from Iowa, 141; ІХ
Chalco, Mexico, maize from, 234
Chihuahua, Mexico, maize from, 247
r Woodson
ы іп =. ANNAL
of the "Flor
and Schery’s “Flora
of Panama” only the
s Index, since a complete а will be appended at the end of each volume
Coal фен а of, Жр 291; map show-
in where found, 143
ERR enum. 303
Conostoma, 291; oblongum, 291, di Spr
i micropylar region
>
e
<
an
қз
о
la]
"38
&
3
~
Меди Поза from
Cycad forest of E. (m Hills, N. D.,
Cytological study of yeast wb г
cerevisiae), 249
D
Dent corn, 148, 153, 1
Des Moines, Iowa, map showing coal mines
near, 143
Diacrium, 378
E
Ecological factors in maize variation, 162
Elleanthus, 22
"Elote" varieties of Mexican maize, 149,
164, 174 (pl. 7); showing кезе. Бе-
tween, and common varieties, 105
dM кез deposit, near Gilboa, N.
«y 309
Кыйан, 403 К
Epidendrum, 315; ciliare var. Oerstedii,
327; > apt var. angustisegmentum, 371
Eryth dh
Ethnological Eds of maize, 161, 162
F
Ferns, fossil, 141,
Feulgen udies of yeast cul-
, 288
Fixations in yeast cultures, 252
Flemming's triple stain in studies of yeast
cultures, 256, 278
plant families and new entities are
(405)
406
Flora of Panama, Part ІП, Fasc. 2, 1, Fasc.
Fossil ferns, 141, 3
Fossil forest, The i 309; views of,
311, 312
Fossil T of Arizona, the Black Hills,
N. Florissant, Colo., and Gilboa, N.
To 309; of Yellowstone National Park,
309
G
Galeandra,
Gallatin e forest, The, 309; views of,
311, 312
Genetics іп ma of denting, 171; of
multiple Theon E 171
Giemsa technique in study of yeast cultures,
see Robinow's Giemsa technique
Guadalajara, Mexico, maize researches in,
148
Guatemalan Big Grain maize, 770, 171
H
Habenaria, 11
Heidenhain's haematoxylin in studies of
yeast cultures, 256, 250, 286
Hexisea, 139
Huehuetoca, Mexico, maize grown near,
238, 240
Huichol Indians, varieties of maize grown
by, 161,
Hybrid yeasts, 250
I
Another Medullosa from, 141; map
pre coal mines near Des NAM 143;
me petrified seeds from, 2
шы 397
J
Jalisco, Mexico, maize grown іп, 140, 153,
158, 159, 176-20
Johansen’s — violet stain on yeast cul-
tures, 253, 250,
Jurassic oa forest of North Dakota, 309
K
Kamaraspermum Leeanum, 296, longitudi-
nal section of seed, 206, 300, 301, 306,
transverse section, 208, 200, 306
Kern, El M., N. Andrews
me pet-
rified seeds from Iowa, 291
[Vor. 33
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Kernen, Jules A., Henry N. Andrews and.
I toour knowledge of Amer-
ican Carboniferous е Қы» Another
Midulloes from Iow
Kuleshov's studies on ы МҸ 160
L
Laelia, 382
Lenz, Lee y N. UN and. The
Gallatin fosil T
Lepanthes, 8
Lepiocarpon, 303
Liparis, 1
M
“Magnicorp” in yeast cell: explanation of
term, 254, 254; fixation of, 257; stain-
Maiz, 147; amarillo, 749, 185, 198, 200,
2 ; amarillo del cerro,
140; de tierra fria, 227; del cerro, 216;
del pais, 167; dulce, 149, 167, 174 (pl.
8), 188; espiga blanca, e луу =:
4), 211, 244; reventador, 140, 161, 168,
175, 170, 180; rosquera, 213; tabloncillo,
140; tampenquefo, 749; tulekenio, 210;
umado, 749; valenciana, 107; vaquerefo,
1,2
Maize іп ме хісо--А preliminary survey,
147; Appendix, 175
Maize, Mexican: Central Mexican type,
159; characters used to study, 152; classi-
fication of, 169; collections, 175; denting
, 148, 153, 171; "elote" varie
164, 174 (pl. 7), relation between com-
r
о
206-227; from
Toluca, 140, 158, 163, 220-233; genetics
of, 171; Mexican Narrow Ear,
(pl. 2); Mexican Pyramidal,
170, 171, 174 (pl. 4, pl. 5); Mount
Yellow, 163, 170, 171, 174 (pl. 6); races
1946)
INDEX
of, 160, 171; variation in, 147, 158;
varieties of, 149, 164, 175-247
Malaxis, 128; Wendlandii, 136
Masdevallia, 73
Medullosa, Another, from Iowa 1
Medullosa, 141; anglica, 142, var. ioensis,
142, stelar mm of, 145, I4Ó; sect.
РД
anglorota, 144; Thompsonii, 142
Metepec maize, 2 5
Methyl violet stain in studies of yeast cul-
tures, 253. 256, 27
Methylene blue-eosin stain in studies of
yeast cultures, 252, 258, 28.
Mexican Narrow Ear maize, 160, 174(pl. 2)
Mexican Pyramidal maize, 160, 168, 170,
171, 174 (pl. 4, pl. 5)
Mexico: Maize in—A preliminary survey,
147; Appendix, 175; approximate cross-
section of central, where maize studies
were made, 159
Mexico-Toluca maize, 163, 220-23
Michoacan, Mexico, study of maize grown
in, 153, 158, 163, 206-22
Mitrospermum, 295; compressum, 295
Mountain Yellow maize, Mexican, 163, 170,
171
М
Nagel, Lillian. А cytological “т of yeast
(Saccharomyces cerevisiae),
Narrow Ear varieties of ма maize,
168; 171. 174
O
TR P ik Part III, Fasc. 2,
‚ Hasc
P
Paleontological investigations in Iowa coal
mines, ud in petrified forests, 309
Palmorchis, 3
Panama, Td of, Part III, Fasc. 2, 1, Fasc.
95-215
“Рагуісогр” in yeast cell, explanation of
term, 254; staining, 25
Petrified seeds from Iowa, Some, 291
Phragmipedium, 8
Physosiphon, 7
Platyglottis, 395
Pleurothallis, 85; um var. echinata, 120
Pogonia, 17; 1
M Metu:
Ponthieva,
Popcorns, аа 148, 140, 161, 168
407
Pop-dent corn, 148; grown at Toluca,
Mexico, 140, 170
Prescottia,
Pteridosperms: fossil stems of, 141
Purificacion, Mexico, varieties of maize at,
140
о
R
Me. ом јер 292; cyclocaryon, 295;
spina 292, diagram of seed, 203,
cellular detail, 204, ү: жа ә section,
304
Robinow’s Giemsa technique in studies of
yeast cultures, 252, 255, 260, 276, 280,
282, 284, 286, 288
S
Saccharomyces cerevisiae, Cytological study
of, 249; studies of buddin
284, 286, 288; techniques, 255, 260,
terminology of cell parts, 254, 2
San Francisco, Mexico, maize grown in, 228
Scaphosepalum, 8
Scaphyglottis, 386
Schery, Robert W., Robert E. Woodson
and. Flora of Panama. Part III, Fasc. 2,
Te hase. /2, 315
Seeds, Some petrified, from Iowa, 291
Selenipedium,
Sequoia magnifica, petrified stump of, 311
Shuler Coal Mine, near Des Moines, Iowa,
map showing location of, I
Sobralia, 27; amabilis, 30; callosa, 31
Sporulating cultures d Saccharomyces cere-
visiae, - 284, 286, 288
Spiranthes, 4
Staining Eu employed in study of
де 5% 251
Stelis,
Stems, а fern, 141
Stenoptera, 39
укоб. 264
Т
Tempskya, 310
Tepotzotlán, Mexico, maize grown in, 236,
237
Thompson, Frederick O., coal balls received
from, 141, 291
Toluca, Mexico: Mountain Yellow maize
408
grown іп, 163; Mexican Pyramidal maize
from, 170; varieties of maize grown near,
49, А
Toluidine blue solution, effect of, оп bud-
ding yeast cultures, 263, 263
U
Urbandale Mine: coal-ball plants in, 141,
291; map showing location of, 143
V
Vanilla, 20
Variation in Mexican maize, 146, 156, 156,
163; grids used for diagramming, r50,
I51; methods of studying, 152
W
ics Louis O. 5-7 5 Рапатпа,
art ІП, Fasc. 2, 1, Fas
[Vor. 33, 19461
ANNALS OF THE MISSOURI BOTANICAL GARDEN
Woodson, Robert E., and Robert W. Schery
and collaborators. Flora of Panana, Part
III, Fasc. 2, 1, Fasc. 5, 315
bd
Yeast (Saccharomyces cerevisiae), A cyto-
Yeasts, wild, studies of budding cultures
of, 264, 288
Yellowstone National Park, petrified forests
of, 309, 510
Жы
Zea Mays, 148; races of, 160
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